La publicacin del presente libro cuenta con la autorizacin de Da.

Conchita
Ramrez Gomez, hermana del ilustre profesor autor de este libro.
Como podremos comprobar a travs de su curriculum y de las reseas a las que se
hace referncia, as como en el contenido del presente documento ha sido uno de los
grandes cientficos y miclogos del siglo XX, habiendo sido declarado el Miclogo
nmero 1 en 1985 en el congreso internacional de Cientficos celebrado en Blgica.
Los honores todos para un hombre que naci en el pas y en la poca equivocada.
El resto de los pases con inters en su rama de investigacin se disputaron su
sabidura e intervencin hasta los ltimos das de su vida.
Dedicado a Da. Conchita Ramrez, mi mejor amiga. Cuya esperanza es que el
esfuerzo y saber de su hermano estuviera al servicio de toda la humanidad.
Motivacin que gui el trabajo y todo el esfuerzo del sabio a lo largo de los aos
de dedicacin absoluta a la investigacin.

CURRICULUM VITAE
Actualizado al ao 1984
CARLOS RAMIREZ GOMEZ
Realizado por l mismo en vida.

1978 - CARLOS RAMIREZ GOMEZ

Profesor de Investigacin del Instituto Jaime Ferrn de Microbiologa del C.S.I.C.
CURRICULUM VITAE
1948 1950 -

LICENCIADO EN CIENCIAS NATURALES,

Universidad Comptutense
DOCTOR EN CIENCIAS NATURALES, Universidad Complutense

HISTORIAL CIENTIFICO EN EL CONSEJO SUPERIOR DE INVESTIGACIONES

CIENTIFICAS.
1948.Contratado por el Instituto "Jose Celestino Mutis de Farma -cognosia para dirigir el
Laboratorio de Histologa Vegetal para el estudio de plantas cauchferas, hasta octubre de 1950.
1950.- En octubre pasa al Instituto "Jaime Ferrn" de Microbiologa, realizando trabajos de identificacin
de mohos y leva -duras en esputos de tuberculosos, en calidad de Becario. En Diciembre de dicho ao es
nombrado Jefe de la Subseccin de Micologa General de dicho instituto.
1952.- Se traslada a Holanda pensionado por el Consejo para ampliar estudios sobre Mohos y Levaduras en
el Centraalbureau voor Schimmelcultures en Delft y Baarn.
1954.- Es nombrado Colaborador Cientfico en Ciencias Biolgicas.
1956.- Pensionado por el Consejo, se traslada a Paris para ampliacin
de estudios sobre Hongos superiores y sus toxinas en el Laboratoire de Cryptogamie del Musum National
d'Histoire Naturelle de Paris. Alli realiza un trabajo sobre la ecologa de las levaduras que viven sobre
hongos superiores.
1957-58.- Se traslada a Ginebra con la consideracin de pensionado
por dos aos, en cuya Universidad y en el Laboratoire de Botanique
Gnerale, con el Prof. F. Chodat, realiza investigaciones sobre Fisiologa de hongos, descubriendo los
Ritmos Circadianos en la respiracion de las levaduras.
1959- Con la misma consideracin de Pensionado, se traslada por un ao a la Universidad de Zurich, al
Laboratorium von Allegemeine Botabik,
con el Prof. U. Leopold, realizando investigaciones sobre Gentica de hongos,
principalmente sobre la estructura intragentica de los cromosomas de la levadura Schizosaccharomyces
pompe, trabajo que fue objeto de una comunicacin en 1963 en XI International Congress of Genetics,
celebrado en la Haya en Septierabre de 1963.
1959.- Es nombrado Investigador Cientifico del C.S.I.C.. del Patronato "Alonso de Herrera"
1960-63.- Habiendo obtenido un Postdoctorate Fellowship del National Research Council of Canada, se
traslada a la McMaster University de Hamilton, Ontario, Canada, en el Laboratorio de Botnica, con el
Prof. J.J. Miller, donde por el espacio de cuatro aos realiza investigaciones sobre la esporulacin,
su fisiologa, su cariologa en las levaduras, llegando a fotografiar por primera vez en estos hongos
los cromosomas y las divisiones
mitticas.
1964-67.- Con un contrato del ejrcito de los E.E.U.U., se traslada a Ridgefield, Connecticut, al New
England Institute for Medical Research, donde durante cuatro aos realiza investigaciones sobre las
interaccines entre las bacterias patgenas y los microhongos en sistemas abiertos y los mecanismos de
resistencia y susceptibilidad a las enfermedades. En dicho Instituto fue nombrado Jefe del Laboratorio
de Micologa.
1968 - Regresa al Instituto "Jaime Ferrn", donde vuelve a ocupar su puesto de Jefe del Laboratorio de
Micologa General y Aplicada.
En el verano de dicho ao se traslada a Hamilton, a la MacMaster University, para familiarizarse con las
tcnicas de Autoradiografa, obteniendo el Titulo de usuario de Sustancias Radioactivas de la Comisin
de Energa Nuclear de Espaa.
1970 - En octubre de dicho ao es invitado oficialmente por el Prof. Scott del Departamento de Qumica
de la Universidad de Yale, New Haven, Connecticut (USA), para participar en un proyecto sobre produccin
de vitamina B12 por

microorganismos.
1971 - Nombrado Profesor de Investigacin del C.S.I.C.
1972 - se traslada a Lille, donde participa a un curso de un mes en el Instituto Pasteur dirigido por
el Prof. Buttiaux, sobre Microbiologa de los Alimentos.
Seguidamente y por tres meses, se traslada al Institituto de Microbiologa Tcnica y de los Alimentos
(TNO) de Holanda en Zeist, para realizar estudios sobre micotoxinas y su deteccin en los alimentos y
piensos.
(Para aos siguientes vanse las pginas de publlcaciones, etc.

RESUMEN DEL HISTORIAL

Contratado dos aos (1948-1950)
Becario del C.S.I.C: cuatro aos (1950-1953)
Colaborador Cientfico: cinco aos (1953-1957)
Investigador Cientfico: Doce aos (1957-1968)
Profesor de Investigacin: Once aos (1968-1984)
TOTAL : Treinta y cuatro aos,

COLECCIN PERSONAL DE HONGOS (MOHOS Y LEVADURAS) EN EL LAB0RAT0RI0

DE MIC0L0GIA GENERAL Y APLICADA.
Desde el ao 1950 en que ingreso en el Instituto "Jaime Ferrn" emprend la tarea de conservar en coleccin
permanente los mohos y levaduras aisladas e identificadas, procedentes de diversos sustratos. Esta
coleccin alcanzo en 1956 la cifra de 2.000 cepas entre mohos y levaduras. Desgraciadamente, desapareci
al ausentarme al extranjero, pues fue llevada a la Universidad de Salamanca sin mi autorizacin.
Al regresar al Instituto en 1968, reanude la tarea de reconstruir una nueva coleccin de hongos, que pasa
en la actualidad la cifra de 7.000 cepas, entre mohos y levaduras procedentes de toda clase de sustratos
(aditivos de alimentos, especias, comerciales, tierras, aire, agua, hospitales, etc.), tanto de Espaa,
como del extranjero (U.S.A., Chile, Argentina, Tierra del Fuego, Sahara, etc.)
Para evitar las continuas resiembras las conservo liofilizadas y en tierra esteril.

TRABAJ0S PUBLICAD0S CLASIFICADOS POR MATERIAS SOBRE

ECOLOGA MICOLOGICA
47 trabajos publicados: 9 en Microbiologa Espaola y los restantes en revistas
extranjeras.

SOBRE FISIOLOGA DE HONGO.

3 trabajos en Microbiologa Espaola y 7 en revistas extranjeras.

SOBRE GENETICA DE LEVADURAS

Un trabajo en los Proceedings del IX INTERNATIONAL CONGRESS OF GENETICS. La Haya
, 1963,

SOBRE CARIOLOGA DE MOHOS.

Uno en revista extranjera y 3 en Microbiologa Espaola.

SOBRE

TECNICAS MICROFUNGICAS.

Uno en Microbiologa Espaola y cuatro en revistas extranjeras

SOBRE FERMENTACIONES.
Uno en Microbiologa Espaola.

SOBRE METABOLISMO DE HONGOS.

Cuatro trabajos: 2 en Microbiologa Espaola y dos en revistas extranjeras.

SOBRE INTERACCINES LEVADURA-BACTERIA EN CULTIVO

CONTINUO.
Dos trabajos en Microbiologa Espaola.

SOBRE MICOTOXINAS
Cinco trabajos: dos en revistas espaolas y tres en extranjeras.

SOBRE RADIACIONES GAMAS Y SU INFLUENCIA EN LA ESPORULACION

Un trabajo en revista extranjero

80 TOTAL DE TRABAJOS PUBLICADOS HASTA LA 1984:

27 en revistas espaolas, el resto en revistas extranjeras.

TRES MONOGRAFIAS

"COMPILATION OF DESCRIPTIONS OF NEW CANDIDA SPECIES WITH KEYS TO ALL SPECIES

OF THE GENUS DESCRIBED UP TO DATE" 1974. Micr. Esp. : 27: 15-80.

LOS PSILIDOS DE ESPAA" Bol. Real. Soc. Esp. Hist. Nat. Vo. 55, 56, y 57, 210
pg. 1955, 1956, 1959.

"MANUAL AND ATLAS OF THE PENICILLIA" 1982. Elsevier PuB. Co Amsterdam. XV + 874
pgs.T

TRABAJOS EN COLABORACIN
X) Sa colaboraciSn con LA SECCIN DE MICRQBIOLOGA TECNICA, se estaa reali-zaado estuciios sobre
el pool de amiaoScidos eh los HALOBACTSRIUM, con el fia de tener mas datos para peraiitir situar
estos microorgaaismos ek el lugar que les corresponda taxonomicamente*
2)
Coa la SEC-ION DE FIJACION DEL NITROGENO se esta estudiando el papel que
desempenaa LQ5 HQKGQS SUPERIORES en la utilizacion total de las ligainas, prod
ducto dc desehcho de las papeleras*
3)
Ea colaboracion coa un medico Alerg6logo de las Palmas de Gran Canaria, se
esta 11 vando ua ESTUDIO ECOLOGICO DEL MEDIO AMBIENTE de esta Capital, en
cuaato a la MICOFLO.U CONTAMINANTS DEL AI3E Y SU INFLUENCIA p LAS ALS3fi*AS |
7/ Referencias de publicaclones o Tesis doctorales en relacin con el Proyecto:
Sobre tazonoma de mohos y ecologa mlcolgca ha pubilcado veinte trabajos. Sobre fisiologa
da mohos ha pubilcado diez trabajos. Sobre accin de las radiaccines gamma sbre la esporulacin
pubilcado un trabajo. Sobre metabolismo de Mohos ha pubilcado cuatro trabajos.Sobre
Micotoxinas ha presentado dos trabajos en el Congreso Nacional de Mlcrobiologa da Granada
(1973), dos en el de Salamanca (1975), uno en el International Symposium on Mycotoxins in
Foodstuffs da Paris (1976). De los trabajos pubilcados, 44 tratan de temas de Micologa General
y Aplicada. Dos Tesis doctorales presentadas y una on preparacin
tratan de la micoflora micotoxignica contaminante de produccin destinados a la alimentacin
humana.
Repercusin
Trataremos do reunir en este apartado, aunque no exaustivamente la repercusn Internacfonal
de nuestros trabajos, que se hace bien patento
en las numerosas menciones en revistas extranjeas y en tratados de nuestra especialidad:
1954,* J.Bofdn et F,Abadie.Bull.Soc.Mycol.France,70: 353-383;1957, J.Boidin et J.M. Adzet,
Bui|iSec, MymlFrwice, 73:331*342; 1958, "The Chemistry and Biology of Yeasts, A.H. Cook
(editor); 1959 Van der Walt,
J.P. Antony Van Leewenhoek 25: 458-464; 1962, Artagaveitia-Allende, R.C.
y C. Leganani.
Micopathologa et Mycologa Appl. 18: 140-146; 1962, L. de mONTEMAYOR
y B. Heredia de Ganero, Mycopatholga et Mycologa Appl. 18: 1-52; 1962,
Artagaveita-Allende, B.C. Mycopathologa et Mycologa Appl. 25: 299-302; 1962, M.
W.Miller, H. J. Phaff and H.e.Synder. Mycopatologa Appl. 16: 1-18; 1965 J. Boldin
J. Abadie et Y. Lehoday. Bull. Soc. Mycol. France 81: 5-23 1965, M.C.Pignal et J. Boldfn, Dull.
Soc.Mycol,France SI; 197-226; 1965, D, Yarrow..Ant, van ' oe Leewenhoek 3It 405-467;
f965,R*C#Artagaceftfe>AMende,Myoop*thologa et Mycologfa Appl* 2S;237*273; 196> J.Bofdfn,J,
Plgnal et M.Besson, Bull, Soc.Mycol.France 8tj566*60@; 1966 "The Fungi", An advanced Treatise,
Ed,G, C *Afnsworth and A#S#Sueeman,Vot* I pag 135-I4tj 1966, G,ffiurtan en "The Fungi", Vol,
II pgs.367; 1967* J. J, Miller, Bull,Sec,Hist, Natur.Toulouse,Francef 103;3a7*339; 1964,
J.Bo!dln,F.Abadle et Y, Leho* dey et M.C Filial,Buff.Soc,MycoUFrence, 80*397*438j W7,R.Moniro*
cher, Bvtil Soc.Mycol.Framce 83*641-730 J 1968, F. Abedle,My wathologle et Mycologfa Appl.
36i81*t3j 1968, MlNor^M* liter notional Atomic Ertorgy Agency,Vlnna pg. 181-10*1 J* J* Milter
and G^offman-Ostenhoff, ZeftachrM fur AHgemelne Mfkroblofogfe, 44t63*294f If 69, J,J. Ml Her
on "The Yeasts", EoiA^H.Rdfi and J, S.HarHaon, Aeectemlc Press* Vol I Pfl. 93-104*334*381} 1970,
"The Yeasta* A Teconomic Study, Lodde*, J.Ed.pgs ||fiIM,)7fi.861l1Pti.lofri 13SU

LISTA DE ESPECIES NUEVAS DE MICROHC

)NGOS PUBLICADAa

17:214-218, 1082 19: 08-102, 1984 19 98*102. 1984

(Paris) Mycol. rparia)
Pichia pseudopolymorpha*, Pichia rhodanensis
1952.- Sporotrichum sanguineum p-.., Mycol# (Pmrim) 1954.- Saccharomyces sociasi Rv,
Myeol. Candida boidinii Rav.
1953
Pichia strasburgensis
Endomycopsis balearica Debaryomyces toletanus Pichia chambardii
Myeal. (P,ri). 18il49-l56. 1983 Rsv. Mycol. (Paris), is, 149-186. 1983 Rev. Mycol.
(P.ria). 18t 149-156. 1963
1954
Bull
Mir. Ep. 7:107T110^".1954 Mler. Ep, 7illllX4# 1954
Men*. Soo. Una, Lyon. 23:151.152
1957.-Geotrichum zambettakesii
Sporobolomyces albidus Candida anomala
Rsv. Mycol. (Paris).25 433-436, 1954 Mler. Esp. 10: 215-247, 1957 Mlcr. Esp. 10:
215-847. 1957
Sporobolomyces boleticola Micr. Eap. 10: 215-247, 1957 Torulopsis buffoni Mlcr. Eap.
10: 215-247,'1957
37-41, 1986
191-200, 1956
Mlcr. Eap. 9j 147-162, 1956
1956.- Sporobolomyces hispanicus Mier. Esp. 9; Endomycopsis ovetensis Mlcr. Esp. 9;
Torulopsis holmii, var. acidi-lactici
1972.- Candida iberica
Can. J. Microbiol. 18: 1778-1780, 197S Rav.
Candida levantina
Mycol. (Paris) 40: 108-110, 1972
1978.- Torulopsis navarrensis Mycopatholoffia 63s 81-63, 1978
Penicillium hispanicum
Torulopsis pampelonensis *J ,Can. J. Micron. 24: 433-435, lira Blastobotrys navarrensis
Myeopatholojia. 63: 41-45. Penicillium fagi Mycopathologi*. 63: 57-59, 1978
(an pranaa) (on pranaa)
(an prensa)
Mycopathologa
Penicillium grancanariae Myeopathologa Penicillium palmensis Mycopathologa
1978.- Halobacterium vallismortis
Bacteria extricta hal6ila.
Can. J. Microbiol.
(*n prem
En total 26 especies nuevas de hongos (22 levadura-y 4 mohos) y una nueva especie de bacteria. A estas
especies nuevas hay que afla4
dir las siguientes especies nuevas de Penicillium qu estan en prensa: P. turolense, P. murcianum, P.
ovetense, P. asturjanum, P. onobense, p. castellonense, P. aragonense, P. aurantio-flammiferum, P.
gallaicum p. granatense, "Penicillium ilerdanum, P. gerundense, P. valentinum, P. alicantinurn, P.
malacaen.se, P. tarraconense, P. vasconiae y P. novae-caledoniae Smith, var. album, nov. var.
lo que suman en total 40 especies nuevas de microhongos.

1984
CARLOS RAMIREZ
PUQLICACIONES
1949.- Ramirez C. Nota sobre hall
.azgo de "Acarapis woodit" (Acarioais de las abejas) en la isla de Malloroa. Bol. Real. Soc.
Hist. Nat, Madrid. - Ramirez, C, Socias, A, e
|^ 1955
/^1956. W 1956.
/ 1956
I 1952
j 1952 4 1953
1953.
C 1953. 1 1953.-? 1954.-^ 1954.< W.9 54.I 1954.)Vl955.47 611-118. Gil. Presencia de Aspergillus y Penicillium en esputos y orina de tuberculoses.
Micro- biologa Esp. 5: 113-121.
1
Ramirez, C. Une nouvelle espece de Sporotriohum. le Sporotrichum sanguineum. Revue de Mycologie. Paris. 17j 214-218
Trois nouvelles especes de levures isolees de liqueur tan-nantes. Revue de Mycol. Paris. 18'!
149-158.
Ramirez C. Introduccin de una modificacin al metodo de Wickerham para determinar el poder
fermentativo de la rafi-nosa por las levaduras. Estratto degli atti del VI Congresso
Internazionale di Microbiologa. Roma. Septiembre. I: 872-873 Ramirez, C. Contribucion a la
tincin de ascosporas y clami-dosporas de las levaduras. Microbiol. Esp. 6: 239-246.
Estudio de nuevas especies de levaduras ai&ladas de distintos sustratos. Microbiol. Esp. 6:
249-253.
Deux nouvelles especes de levures isolees de divers substrats. A Revue de Mycologie (Paris).
19: 98-102.
Ramirez, C., et J. Boidin. Saccharomyces chambardii, nouvelle J
espece de levure isolee de liqueur tannante. Bull. Mens. Soc.
Linn. Lyon. 23: 151-152.
Ramirez, C. Socias, A, y Pelaez Campomanes, F. Una nueva eg- i pecie de Debaryomyces. El
Debaryomyces toletanus. Microbiol. Esp. 7: 111-114.
Ramirez, C, Genestar, R., y Socias A. Una nueva especie de "7 Endomycopsis: Endomycopsis balearica
aislada de concentrado de tanino de encina. Microbiol. Esp. 7: 107-110. Ramirez, C. "Los Psilidos
de Espafia". I. Bol. Real. Soc. Esp. Hist. Nat. 53: 151-218.
Ramirez, C. y Socias A. Estudio cariologico de algunas levaduras ascosporogenas y
anscosporogenas. Microbiol. Esp. 8: 389417.
Ramirez, C. "Los Psilidos de Espafia". II. Bol. Real Soc. Hist.
Nat. 54: 63-106.
Ramirez C, Pelaez Campomanes, F. El Sporobolomyces hispani^jis
so. nov., nueva especie de Sporobolomyces aislado de extracto
de corteza de encina. Microbiol. Esp. 9: 37:41.
Ramirez, C. y Sierra, G. Una nueva variedad de Torulopsis holmii (Jorgensen) y estudio de sus productos metabolicos.
Microbiol. Esp. 9: 147-162.

II 1956 /? 1957
tf 1957. 7" 1957.
$1 1959.
^ 1959.
-?3 1959.
Zh 1960.2;,-1960. 1963. -Jl 1964.^^T 1964.-4^1965.-%t> 1968.-}( 1968.-3 ^1969.13 1970.-3^1972..- Ramirez, C, y Pelaez Campomanes, F. Una nueva especie dP
Endomyces: Endomyces ovetensis. MierobioX. Esp. 9: 3.91-200 ' *
. - Ramirez, C. Contribucin al estudio de la ecologa de las
levaduras. I. Levaduras aisladas de hongos carnosos. Microbiol ,.
Esp. 10: 215-247. "-"-croDiol. )|
elemen-Microbiol.
Real Soc.
Ramirez, C. Nota sobre algunas modifieacione* introducldas a la tincin de nucleos de
hongos. Microbiol. Esp, 10: 201-203
Ramirez, C. y Socias, A. Cromosomas ligados y Mitosis tales en los nucleos de Aspergillus
y Penicillimjj Esp. 10: 259-286.
Ramirez, C. "LOS PSILID0S DE ESPANA". III. Bol Esp. Hist. Nat. 57: 5-87.
Ramirez, C, et Chodat, F. Etudes sur la periodicite des echan-ges gazeux chez une levure:
Kloeckera apiculata. Microbiol. Esp. 12: 265-296.
Ramirez, C. Methode rapide et simple pour determiner le degre
de fermentation du raffinose par les levures. Ant. van Leeuwenhoek. 25: 445-448.
Ramirez, C. Une nouvelle diagnose d'une levure isolee d'Amanita JTJ livido-palescens:
Geotrichum zambettakesii (Ramirez) comb nov. Revue de Mycol. (Paris). 25: 433-436.
Ramirez, C, and Miller, J. J. Observations on vegetative nuclear division inScJccharomyces
cerevisiae. Can. J. Microbiol. 8: 603-608.
Ramirez, C, and Miller, J. J. Accumulation of free proline during yeast sporo^enesis.
Nature. 197: 722-723. Ramirez, C. Studies on the causes of reciprocal inhibition of
pseudomycelium formation between parallel colonies of yeasts of the genus Candida. Can.
J. Microbiol. 10: 931-933. Ramirez, C, and Miller, J. J. The metabolism of yeast
spo-rulation. VI. The changes in amino acid content during sporo-genesis. Can. J.
Microbiol. 10: 223-231.
Miller, J. J., Kingsley, V., and Ramirez, C. Comparative sensitivity to Gamma radiation
of yeast mitosis and meiosis. Radiation Bot. 5: 384-390.
Ramirez, C, and Ransom, J. P. The growth of Candida albicans and Salmonella typhimurium
in mixed cultures. Microbiol. Esp.
21: 77-88.
Ramirez, C, Ransom, J. P., and Pitkin, D. R. The in Vitro Interaction of Candida albicans
and Salmonella typhimurium. Microbiol. Esp. 21: 88-91.
Ramirez C. Antibacterial action of non-volatile substances extracted from Artemisia
tridentata Nutt. ssp. tridentata. Can. J. Microbiol. 15: 1341.
'Uamivez, C, Rodriguez, D., and Lanzuela, F. Rapid diagnosis of Candida albicans. Mycopath.
et Mycol. Appl. 40: 10?-111. Ramirez C, and Gonzalez, C. Candida iberlca sp nov. A new
U species of yeast Isolated from Spanish sausages* Can. J. Micro** biol. 18: 1778-1780.

3 ^ 1974 Si 1974 311976 3^1976.

j^ 1977. It (? 1978.
|i? 1978.
Ijl 1978.
Jj> 1978.
4j If 19 78.-^ 1978.4<J* 1978 /,"? 1978
P# 1978. l( 1979.
3
Inhibition of germinatioi
.- Ramirez, C. in.iiuiiion oi germination and sporulation (com
diation in Neurospora sitophila Shear and Dodge by CO . My."
copath. et Mycol. Applicata. 52: 173-175. 2*
m
. Ramirez, C. A Compilation of descriptions of new Candida species with keys to all species of the
genus described up to date. Microbiol. Esp. 27: 15-80.
.- Lopez-Aydillo, N. R. y Ramirez, C. Probable etiologa de la Alopecia en los enfermos latiricos
espafloles. II. Trab. Inst Cajal Inv. Biol. 68: 333-349.
. Sesma, B., Ramirez, C. y Rodriguez Burgos, A. Estudio de la flora microbiana durante la maduracin
de embutidos. Caracterlsticas de las levaduras aisladas del salchich6n de Pamplona. Rev.
Agroquiraica y Tecnol. de los Alimentos. 16: 539~544. I
Guergue, J. M. and Ramirez, C. Incidence of aflatoxin potencial contamination in Spanish sausages.
Ann. Nutr. Alim. 31: 485-488.
Ramirez, C. and Moriyon, I. Torulopsis navarrensis sp. nov. i . A new species of yeast isolated
from an acid washed brown soil in the province of Navarra (Spain). Mycopath. 63: 61-63.
Martinez, A. T., and Ramirez, C. Penicillium fagi sp. nov. isolated from beech leaves. Mycopath.
63: 57-59.
Sesma, B. and Ramirez, C. A new species of Blastobpthrys: Blasto-it
bothrys navarrensis sp. nov. Mycopath. 63: 41-45. Ramirez, C. and Martinez, A. T. Torulopsis
pampelonensis sp. nov.1^ A new species of yeast isolated from beech forest soil. Can. J. Microbiol.
24: 433-435.
Ramirez, C. , Martinez, A. T. , and Ferrer, S. Three new species "* **
of Penicillium. Mycopath. 66: 77-82.
Lopez-Aydillo, N. R. y Ramirez, C. Probable etiologa de la
alopecia en los enfermos latiricos espaholes III. Resultados
de las investigaciones sobre toxinas en los hongos que se desarrollan en la superficie de las semillas de Lathyrus sativus
(Almorta). Arch. Neurol. Madrid. 41: 461-486.
Martinez, A. T. and Ramirez, C. Microfungal biomass and number of propagules in an andosol during
the year and in the whole profile. Soil Biol, and Biochem. Australia. 10: 529-531. Gonzalez, C.
Gutierrez, C. and Ramirez. C. Halobacterium i| vallismortis sp. nov. A new amylolytic and
carbohydrate- metabolizing extremely halophilic bacterium. Can. J. Microbiol.
V
24: 710-715.
Martinez A. T. et Ramirez, C. Quelques moisissures du genre Penicillium dignes d'etre notees. Rev.
Mycol. (Parlfc). 42: 97-102 Martinez A. T. and Ramirez, C. Study of the paicrofungal co mmunity
of an andosol. J. Ecol. (Australia). 67: 305-319.

4
b 1979.Martinez, A.T., and Ramirez, | Ecological importance of antibiosis in soil. Production of antibiotic
by species of Penicillium. Anales Edaf. Agrobiol 3s-159-166.
!T t 1980.Torres, J., Guarro, Suarez, G, Sufie , N., C^lvo, M.A.
and Ramirez, C. Morphological changes in strains of'LpergilluB flavus Link ex Fries, and A. parasiticus SpeaTeT
related with aflatoxin production. Mycopathologa: 72*'
171-174.
Tl 1980
i'^> 1980.
| r 1980.
2T & 1980.
Calvo, M.A., Guarro, J., Su&rez, G., and Ramirez, C. Airborne Fungi in Barcelona City (Spain). I.
A two-year study (1976-1978). Mycopathologa. 71: 89-93. S'3 1980.- Calvo, M.A., Guarro, J. Suarez,
G., and Ramirez, C. Airborne Fungi in the air of Barcelona, Spain. V. Studies of the spore content
of air in dwellings. Ann. Allergy. 44; 228-234.
Ramirez, C, and Martinez. A.T., and Berenguer, J. Four ^ ^ new species of Penicillium isolated from
the air. Mycopathologa 72: 27-34. Ramirez, C., and Martinez, A.T. Some new species of Penici- ^
Ilium recovered from the atmosphere in Madrid and from
other substrata. Mycopathologa. 72: 181-191.
Guarro, J., Calvo, M.A., and Ramirez, C. Soil Ascomycetes
(Sphaeriales) from Catalonia, Spain. I. The genus Chaeto-mium Kunze ex Fresenius. Nova Hedwigia.
28: 191-216.
iT 7 1980.- Calvo, M.A. , Guarro, J., Suarez, G., and Ramirez, C. Airborne Fungi in the air of Barcelona
(Spain). 111= The ge-nusAspergillus Link ex Fries. Mycopathologa. 71: 41-43.
$ <i 1981.
$T / 1980.- Calvo, M.A. , Guarro, J., Suarez, G., and Ramirez, C. Airborne Fungi in the air of Barcelona
(Spain). IV. Various isolated genera. Mycopathologa. 71: 119-123. Ramirez, C, and Martinez, A.T.,
Seven new species of Penicillium and one variety of Penicillium novae-caledoniae Smith. Mycopathologa. . 74: 35-49.
o
1981./
1981.(^
1982.^ 2
1982.Ramirez, C, and Martinez, A.T. Four new species of Penicillium isolated from different substrata.
Mycopathologa 74: 163-171.
Ramirez, C, Gonzalez, C, and Gutierrez, C. Rhodotorula matritense sp. nov., isolated from red pepper.
Mycopathologa. 74: 173-175.
Martinez, A.T., Calvo, M.A., and Ramirez, C. Scanning electron microscopy of Penicillium conidia.
Ant. v. Lee-wenhoek. 48: 245-255, 1982.
Ramirez, C. "MANUAL AND ATLAS OF THE PENICILLIA". Elsevier Biomedical Press B.V. Amsterdam. XV +
874 pp.

&H
1983.
if
1983.
U
1984.
61
1984.
xpensive method for>
Ramirez, C. A convenient and ine preparing fungal conidia for scanning
electron Mycopathologa 83: 17-18. m
Martinez, A.T. and Ramirez, C. Rhizosphaera oudemansii (Sphaeropsidales)
associated with needle cast of Spanish
Abies- pinsapo. Mycopathologa. 83: 175-182.
Ramirez, C, and Gonzalez, C. Penicillium flavido-stipj-tatum sp. nov., isolated from sand dunes at
the Arches National Monument, Utah, U.S.A. Mycopathologa 88:3-7
Ramirez, C, and Gonzalez, A. Rhodotorula grinbergsii I sp. nov. isolated from decayed wood in the
evergreen rainy Valdivian forest of southern Chile. Mycopathologa 88: 51-53.
i^ 1984. 1 1984.
1984.- Ramirez, C, and Gonzalez, A.. Two new species and one variety of nitrate-utilizing,
mycelial Candida isolated from decayed wood in the evergreen rainy Valdivian forest of
bfrSoutsMign Chilpa I M$j{fcop&thologa& 88 : 55-60.
3^
Ramirez, C., and Muntanola-Cvetovic, M. Penicillium ju-goslavicum sp. nov., isolated from sun
flower seeds (Helianthus annuus L.). Mycopathologa 88: 65-69 Ramirez, C., and Gonzalez, A.
Pichia scaptomyzae sp. nov. isolated from Scaptomyza multispinosa Malloch (Diptera) in southern
Chile. Mycopathologa. 88: 79-81.
If 1984.- Ramirez, C, and Gonzalez, A. Five new filamentous, glucose-fermenting Candida
isolated from decayed wood in the evergreen rainy Valdivian forest of southern Chile. Mycopathologa. 88: 83-92.
73 1984
71 1984.- Ramirez, C, and Gonzalez, A. Two new, amycelial Candida isolated from decayed wood
in the evergreen Valdivian forest of southern Chile. Mycopathologa. 88: 99-103. Ramirez, C. Validation
of Candida citrica sp. nov. (Furu-kuwa, Matsuyoshi, Minoda & Yamada). Mycopathologa. 86: 125.
7 4 1984.
Ramirez, C., and Gonzalez, A. Candida santjacobensist and
Candida ancudensis, two new species of yeasts isolated
from decayed wood in the evergreen rainy Valdivian forest
of southern Chile. Mycopathologa 88: 105-109.
Jf 1984.Ramirez, C, and Gonzalez, A. Two new filamentous, non
fermenting Candida and a new Schizoblastosporion isolated
isolated from decayed wood in the evergreen Valdivian
forest of southern Chile. Mycopathologa 88: 165-171.
li 1984.
Ramirez, C, and Gonzalez, A. Two new filamentous, fermenting ^
Candida isolated from decayed wood in the evergreen Valdivian
forest of southern Chile. Mycopathologa. 88: 185-189.

1984.- Ramirez, C, and Gonzalez, A

S, r*om dec of southern
Candida ous yeast anamorph
: "- isolated from decayed
Three new, non-fermenting and nitrate-negative Candida
1 / 184
wood. Antony van Leeuwenhoek.
wood in the evergreen rainy Valdivian forest Chile. Mycopatholo&ia 87: 175-180, Dill, Ingrid, Ramirez, C,
and Gonzalez, A.. lignophila sp. nov., a new basidiomycet from south Chilean rotting 50: 219-225.
1 i 1985.
C
Mycopathologi;
Ramirez, C, Dill, Ingrid, and Gonzalez, A. Emendation ^ Candida shehatae (Buckley & Van Uden). 89: 119-121.
$0 1985
Ramirez, C, and Gonzalez, A. Two new species of Apiotri- <3 chum isolated from decayed wood in the evergreen
rainv Valdivian forest of southern Chile. Mycopathologa.
1985.- Ramirez, C, and Gonzalez, A. Two new filamentous, non-fermenting Candida isolated from decayed wood
in the rainy evergreen Valdivian forest of southern Chile. Mycopathologa.
1985.- Ramirez, C., and Gonzalez, A. Rhodotorula nothofagi sp. * V nov. , isolated from decayed wood in
the evergreen rainy Valdivian forest of southern Chile. Mycopathologa. ^ I * Pll-'O
^2 1985.- Ramirez, C. Two new species of Penicillium from Tierra *" del Fuego (argentine and Chile).
Mycopathologa. ^f j* 9 S- /<*"
#4 1985
AjltMl
7jT1985-am
Ramirez, C. A simple apparatus for obtaining fine agar slices of undisturbea^f^ng11^ cultures
for^micrc|photo-graphy/J . Microbiol. Methods. H ' 16 1 " lor''
Ramirez, C. Species concept in Penicillium based on Morphological characeters. Revue de Mycologie.
Paris
1985
(Y^ot<*Hu ,- M^CM^^i 13 ? "7 Ramirez C.The Penicillus. In Adv. in Penicillium and Aspergillus
systematics. R.A. Samson & J.I. Pitt Ed.Amsterd
102: 65-69. 1985.- Ramirez, C. Revision of recently described Penicillium. In Adv. in Pen. and As.
systematics. 102: 135-142.
1985.- Ramirez. C. Are the species of Penicillium as imperfect as we think? In Adv. in Pe_n. and
Asp, systematics. 102:
445-451.

CONGRESOS .wINTERNACIONALES
1953.- Ramirez, C. Introduction de una modificacin al metodo de
Wickerham para determinar el poder fermentativo de la rafino-sa por las levaduras. Estratto degli
Atti dei VI Congresso Internazionale di Microbiologa. ROMA. Sep. I: 872-873
1963.- Ramirez, C, Friis, J., and U. Leupold. Allelic recombination and complementation in
adenine-requiring mutants of Schizosaccharomyces pombe. Proceedings of the XI Internal Congress
of Genetics. THE HAGUE, The Netherlands. Sept. I: 7.
1973.- Gonzalez, C., Gutierrez, C., and Ramirez, C. Effet du pH, du NaCl et de la temperature sur
la synthese et sur l1action des esterases chez Halobacterium. Proceedings of the 1st International
Congress of Bacteriology. JERUSALEM. ISRAEL. Sept.
1974.- Ramirez, C., Gutierrez, C, and G onzalez C. , The G^enus Detail
ryomyces: Revision of the species includec in the genus under t the light of new criteria. IV
International Symposium on Yeasts. VIENNA, AUSTRIA. June. I: E-8
1974.- Ramirez, C., and Moriyon I. New sppcies osf yeast isolated from an acid washed brown soil.
IV International Symposium on Yeasts. VIENNA, AUSTRIA. July. I: E-5. 8-12 JULY.
1976.- Ramirez, C, and Guergue, J. M. Incidence of Aflatoxin potential contamination in Spanish
sausages. Ill International Symposium on Mycotoxins in Foodstuffs. PARIS, FRANCIA. 16-18 Sept.
I: 10.
1977.- Ramirez, C, and Martinez, A.T. A new species of yeast
isolated from beech forest soil. V. International Specialized Symposium on Yeasts. KEZTHELY,
HUNGRIA. Sept. 12-15.
1978.- Ramirez, C, Martinez, A. T., and Ferrer Soledad. Three
new species of Penicillium. XII INTERNATIONAL C0NGR&# OF MICROBIOLOGY. MUNICH, ALEMANIA FEDERAL.
3-8 Sep. J; C*Z1
1981.- (Rarxtibtteztf JCU;, and Martinet, A. T. Contribution to the
Preparation Techniques of conidia for scanning electron microscopy. IV INTERNATIONAL CONFERENCE
ON CULTURE

19-25. P
C. adn Ramirez, C. The THE HAGUE. 15-17 Sept.
COLLECTIONS. BRNO, CHECOSLOVAQUIA. July
1981.- Gonzalez, C., Gutierrez of indicators organism*
TIONAL SYMPOSIUM OF THE FEMS

si

Snificane
I INTERNA1983.- Ramirez, C. amd Gonzalez, A. New species of yeasts isolated from Chilean "palo podrido"
in the rainy VaXdivian Forest IX INTERNATIONAL SPECIALIZED SYMPOSIUM ON YEASTS. SMOLEMICE
CHECOSLOVAQUIA. Abril 18-22. Abstracts: 25-26, -<'.
1983.- Ramirez, C. Ill INTERNATIONAL MYCOLOGICAL CONGRESS, 28-Agosto 3 Sept. TOKYO, JAPON.
Invitado a presentar una ponencia sobre "SPECIES CONCEPT IN PENICILLIUM BASED ON
MORPHOLOGICAL CHARACTERS", en el Symposium on Taxonomy of Food-borne Fungi. Abstracts: 247.
1984.- Ramirez, C. Two new species of Penicillium found in Tierra del Fuego. V INTERNATIONAL
CONGRESS OF CULTURE COLLECTIONS. BANGKOK, THAILAND. 2 5-Noviembre al 1 de Diciembre.
1983,- Martinez, A.T. y C. Ramirez. Effect of wil<fcfire on soil mi-crofungi in a Spanish
coniferous forest. Ill INTERNATIONAL MYCOLOGICAL CONGRESS. 28 Ag.-3 Sept. TOKYO, Japon. Panel
y Abstracts: 541
+ + + + +
1982.- Ramirez, C. Invitado por la USSR ACADEMY OF SCIENCES a la presentacin de su
libro"MANUAL AND ATLAS OF THE PENICILLIA". MOSCU, URRS. Noviembre 1982.
1983.- Ramirez, C. Invitado por el Comite Organizador del III INTERNATIONAL CONGRESS OF
MYCOLOGY, 28 Agosto- 3 Sept. como ponente en el Symposium on Taxonomy of Food-borne Fungi,
presentando la ponencia "SPECIES CONCEPT IN PENICILLIUM BASED ON MORPHOLOQMAL CHARACTERS".
TOKYO 1^ 1985.- Invitado a partidipar en el FIRST INTERNATIONAL PENICILLIUM
AND ASPERGILLUS N.A.T.O. A.R. WORKSHOP. Amsterdam 6-10 mayo.
presentando varias ponencias publicadas en "Advances in PeniciIlium Systematica" (Samson*R.A. and Pitt, J.I. Eds.)483 pp
London and New York. Plenum Publishing Co.

CONGRESOS NACIONALES
9
1971.- Solana, A, , Mo ran. C Mnfin? r D' , XT.. , "Ulan, u, iviunoz, c, Ramirez, C, y Gonzalez, C. "Aislamiento e identificacion
de levadu-ras de muestras de salchichones curados", in CON GRESO NACIONAL DE
MICROBIOLOGA. Barcelona, 2-3 Junio. Resumenes. I; 15.
1971.- Munoz, C., Moran, C, Solana, A,, Gonzalez, C, y Ramirez, C. "Poblacin bacteriana
en distintas muestras de salchichones curados". Ill CONGRESO NACIONAL DE MICROBIOLOGA.
Barcelona, 2-3 Junio. Resumenes. I: 129.
1973.- R amirez, C., Echarri, L., y Sanz, J. "Aislamiento y clasificacion de la micoflora
de especias comer-ciales". IV CONGRESO NACIONAL DE MICROBIOLOGA. Granada. Septiembre.
Resumenes. I: 7
1973.- Sesma, B., Ramirez, C., y Rodriguez-Burgos, A. "Estudio de las diferentes
especies microbianas que condicionan la maduracion de algunos embuti-dos." IV CONGRESO
NACIONAL DE MICROBIOLOGA. Granada. 8 de Septiembre. Resumenes. I: 20.
1975.- Gutierrez, C, Gonzalez, C, y Ramirez, C. "Nuevas aportaciones a la fisiologa
y bioquimica de la fa-milia Halobacteriaceae. V CONGRESO NACIONAL DE MICROBIOLOGA.
1-4 Octubre. Salamanca. Resumenes. I: 54.
1975.- Martinez, A. T. y Ramirez, C, "Estudio de la Comu-nidad microfungica de un suelo
Andico. Conslflflera-ciones generales. Estudio taxonomico y ecologico del genero
Penicillium. V CONGRESO NACIONAL DE MICROBIOLOGA, Salamanca, 1-4 Octubre. ResumenesI:
C-69
1976.- Gutierrez, C, Gonzalez, C. y Ramirez, C. "Asimila-cion de hidratos de carbono
por bacteri&s halofilas estrictas valoradas por auxanogramas". Ill REUNION CIENTIFICA
DE LA SECCIN REGIONAL DEL NOROESTE DE LA SOCIEDAD ESPANOLA DE MICROBIOLOGA. Oviedo.
4-5 Junio. Resiimenes. I: 124.
1977 _ Martinez, A. T. y Ramirez, C. Importancia ecologica en los medios naturales del
fenomeno de la antibiosis: produccin de antibi6ticos por cepas de Peni~

AU t^ I ^A
UHL

lAAA

-''^ * fff iff;

un Andofuelo
1 argo *A,
Santia1977. -

iium. VI CONGRESO NACIONAT nt, . _____ go de Compostela. 6-9 j^8 MIC*BI0L0GIA. SanJ
Martinez, A.T. y Ramirez, C mMM
dad de los hongos microsc6picos en ** l* Com"**~
Variaciones a lo largo del perfil "" 7M~w,*,aei0' Wm aficrf, VI CONGRESO NACIONAL DE MICROBIOLO^*
de Compostela: 6-9 Julio.

1977. -

nov
gfierrez, C, Gonzalez, C. y Ramirez, I vi CONGRESO NACIONAL DE MICROBIOLoga. Santiago de Compostela
"Halobacterium vallismorti SD
bacteria halofila extrema, asimiladora de hidratos de carbono". Resumenes. I: 111.
1979

Martinez, A.JT., y Ramirez, C. "Tipo de ornamenta cion superficial de los conidios de Penicill
VII CONGRESO NACIONAL DE MICROBIOLoga. Cadiz, 24 Septierobre al 1 de Octubre .

1984.-

Gonzalez, A., Ramirez, C. y Grinbergs, J. "Palo po drido". Una transformacion biologica de la madera
animal, i II REUNION CONJUNTA DE MICO-LOGA. 31 Mayo-2 Junio, Bellaterra (Barcelona).

TESINAS DIRIGIDA3 Y PHmam^n^

||
1)
Aislaraiento y Clasificaci^n H I ~ il
,y ittbuxcacion de la flora contarolnante de espe
cias comerciales (Canela y plmienta negra), 1973. ECHARRI PRIM, LUIS. Universidad de
Navarra.
Estudio de la Micoflora de una tierra parda bajo robledal. 1973. MORIYON URIA, IGNACIO.
tfjiverslclad de Navarra.
Estudio de la Micoflora presente en un Andosuelo Espaol .
1973. MARTINEZ FERRER, ANGEL, T, Universidad de Navarra,
4)
Estudio de la Micoflora presente en el piment6n picante,
1974. FERRER ORTEGA, MARIA SOLEDAD. Universidad Complutense
Madrid.
5)
Aislamiento y clasificacion de mohos y levaduras aislados de
piraenton comercial.
1974. GOMEZ CEA, MANUEL. Universidad de Navarra.
6)
Produccin de aflatoxinas por la micoflora contaminante de
especias comerciales.
1974. GUERGUE HUALDE, JUAN MANUEL. Universidad de Navarra.
7)
Levaduras de la fermentacion de aceitunas: Negra de Aragon,
Ojiblanco, Manzanilla fina y Manzanilla Carrasquefia.
1975. RUBIO RABANERA, ROSARIO. Universidad de Navarra.
+ + + TESIS DIRIGIDAS
1)
Contribucion al estudio de las levaduras de taninos.
1955. GENESTAR SERRA, RAFAEL. Universidad Complutense, Madrid
Contribucion al estudio de las levaduras de vinos de Jerez. 1962. ZAJARA LOPEZ, JUAN.
Facultad de Farmacia, Universidad de Sevilla.
Levaduras en la fermentacin de aceitunas verdes. 1965. GONZALEZ CANCHO, FERNANDO.
Facultad de Farmacia. Universidad de Sevilla.
Estudio de la comunidad microbiana durante la maduracion del salchichon elaborado en
Pamplona.
1975. SESMA VEA, BEGOftA. Universidad de Navarra.
r

/tV*/^- -*** S^A. f*^A\*~C4+

ahXi^i /?6
sU
4L
CM. C*> *-H.

TESTS DIRIGIDAS (continuacin)

5) Estudio de la taxocenosis microfungica de una . ,,,,,, ,,.,
ra
bajo robledal.
P^rda
1976. MORIYON URIA, IGNACIO. Universidad de Navarra.
6). Estudio ecologico de la micocenosxs de un Andosuelo bale bosque de haya. 1976. MARTINEZ
FERRER, ANGEL TOMAS, Universidad de Navarra.
7) Contribucion al estudio de la micoflora atmosferica de la ciudad de Barcelona.
1978. CALVO TORRAS, MARIA DE LOS ANGELES. Facultad de Farmacia. Universidad de Barcelona.
8)
Estudio taxonomico y toxicologico de la micoflora presente
en los almidones de uso farmaceutico.
1977. GUARRO ARTIGAS, JOSE. Facultad de Farmacia, Universidad de Barcelona.
9)
. .
Micoflora contaminante y micotoxmas en especias comerciales.
1984. BERENGUER SOLER, JOAQUIN. Universidad Complutense, Madrid,
10)
Flora micologica levaduriforme de las pudriciones de la madera
denominadas "palo podrido" de los bosques de la pluviselva
valdiviana del sur de Chile.
1984. GONZALEZ BERCERRA, ALDO ENRIQUE. Facultad de Ciencias Biologicas. Universidad
Autonoma de Madrid.
MC.

au &*&%*- ^vt-^
' I/A**? lf-*> gC-V **>
SL*-4A^L^M &^> ht *-k & %
&\^

H /&U. -U. f\-~'&'7

CONSIDERACIONES ESTADISTICAS
sobre mi dedicacion exclusiva y total a i Tnwo tigacion durante el periodo de 15 aos (1968-iqa/n
Horas de trabajo obligatorias al afio: Horas de trabajo obligatorio
en 15 aos:
Horas que trabajo por ano:
Horas que he trabajado en 15 aos:
Horas extras por ano: Horas extras en 15 aos:
^Cuantos aos se necesitarian para traba-jar 51.960 horas de modo
normal?
Labor real en aos que he realizado hasta
ahora:

1.880 horas 28.200 1

5.344
80.160

i
u

3.464

51.960
t !
27 aos y 6 meses
42 aos.

80.160
2.8
Ritmo de trabajo que supone
28.200 Lo que supone un ritmo de casi tres veces el normal
2 7 aos y 6 meses.
Afios extras que he regalado al Consejo en 15 aos :
42 aos.
Afios oficiales de trabajo que fcubiesen sido necesarios para realizar la labor
efectuada en estos 15 aos:
2.010
;En que ano nos encontrariamos actualmente si toda esa labor hubiese sido hecha
de modo normal?
89 aos.
,Que edad tendria?
59 millones 400.000 pt
^Cuanto me deberia el Consejo por todas las horas extra en 15 aos?

Contents

Preface

xi

Abbreviations of International Culture Collections

. . . . . . . . . . . . . . . . . . . . . . .

xiii

Part 1 General discussion

Chapter 1

Introduction

Typification of the genus Penici/lium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Cultural and morphological criteria employed in the delimitation and

description of species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Morphology and terminology of conidiogenous structures . . . . . . . . . .

Chapter 2

Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

15

Examination of cultures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

17

Scanning electro n microscopy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

21

Camera lucida drawings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

22

Preservation of penicillia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

23

Part 11

The manual proper

Chapter 3

The Monoverticillata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

Chapter 4

The Asymmetrica

201

Chapter

Asymmetrica-velutina sub-section

345

Chapter 6

Asymmetrica-Ianata sub-section

Chapter

Asymmetrica-funiculosa sub-section

437

447

....................

27

419

Chapter 8

Asymmetrica -fasciculata sub-section . . . . . . . . . . . . . . . . . . . .

Chapter 9

The Biverticillata-symmetrica or Lanceolata . . . . . . . . . . . . . ..

507

Chapter 10

The Polyverticillata

. . ..

609

Chapter 11

Addendum..... ......... .... ...... . . .. ... .. . .. . . .. ...

619

Gonorol blbllogro phy .. , , . , . , . , , . , , , . , , , , . , .. , ,

Chapte r 1
,
Chapt er 13 Ac copt od spoclos ond varlotlos , .. , , , . ,
Chaptor 14 Speci es of dubious ident ity
Chapter 15

,.
,
.

Disposition of species accepted by Pitt (1979 b)

631
641
647
649

Check Iist of synonymies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

653

Colour plates - explanation, legends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

665

Colour piates

673

Alphabetic list of the colour plates of the Penlcllliurn species and varieties .

801

Scann ing electron micrographs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

805

Index to the scanning electron micrographs of conidia of the different

species and varieties of Penicillium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..
Glossary
Author index
Subject index

845
849

o.

861
865

Preface

'1'111': PURPOSE OF THIS 'MANUAL AND ATLAS OF THE PENICILLIA ' IS TO SUMMARIZE

as cxha ustivc ly as po ssibl e th e cu r rc nt information reg arding th e identification

nnd characi erization of all rccogni zabl e spec ies 01' Penicillium published to date,
a lld lo supply th e lat est tcchnical ad va nces av ailable as additional criteria in
lh e idcntifi ca tion of th e speci es , in orde r to facilitate this diffi cult task 1'01' those
who are not highly specialized in my cological taxonomy, applying new a pproach cs whi ch rest fundamentally on the principIe that a good illustration is
worth more than a thousand words . 1'0 ac h ieve this ende a vou r, th e book is desig'lIcd to provide th e followin g.
(a) Full colour plates , reproduced from photographic transparencies, of the
cu ltu ral ch a racte r istics of fun gal colon ies grown under sta nda rd conditions ,
(hu s providing an immediate ex a m ple whi ch may be a fir st approach in id entilicalion Ior th e less speci alized , inter est ed , workers .
(b) Line drawings of th e most rel evant mi croscopic features of eac h species of
Penicillium , represented by standard scale camera lucida drawings , whi ch may be
uscd as templates for comparison with the strain under study .
(c) Scanning electron mi crographs of mature con id ia showing th e fine detail
of th e surface ornamentation of th ese structu re s.
(el) K eys and full d escriptions of th e type sp ecies for th e ultimate id entification a nd characte riza tion of all recognizabl e Penicillium sp ecies e1escribeel up to
date .
This book is directeel mainly towards my cologists , mi crobiologists,
bio chemists, soil scientists and organic che m ist s in hi gh er educ a tion , industrial
laboratories (ph armaceutical , chem ical industries a nd food industries) a nd
gove rn me n t research laboratories; it will als o interest undergraduate and postgraduat e students at universities a nd polytechnic schools.
The a u th or is indebted to Doctors Angel T. Martinez, Angeles Calvo, a nd
J os Guarro for th eir colla bo ra tion in preparing th e Penicillium con id ia for
scanning mi croscop y ; to th e D epartment of Electron Microscopy of the
U niv ersity of Barcelona , Spain, and to th e D epartment of Electron Microscopy

Xll

of the Institute of Soil Sciences of the National Research Council of Spain,

Madrid, Spain, for their generous cooperation in the preparation of the
scanning electron micrographs.
Very special thanks are extended to Dr Shun-ichi Udagawa for his information on Japanese Penicillium species, to Professor Lorenzo Vilas for kindly
translating many Russian papers, and finally to Dr Carmen C . Gonzalez for
her encouragement.
CARLOS RAMIREZ
Madrid, May 1980

Abbreviations of International
Culture Collections

A T e c :
III> UN :

111 (; :
III1IM :
IIKM :

IIIU . :

C IIS :

CC M :

<:I-: H:
C I. M R:

CM I :

CMT :
CS IR :

DAOM :

Am eri can T ype Culture Collection , Ro ckville , MD, USA .

Botany D ep artment , University of Nottingham , UK .
Botanisches Institut, Greifswald , GDR .
Botanisches Institut , H ann .-Mnden , FRG .
Department of Type Cultures, Institute of M icrobi ology, USSR
Acad em y of Scien ces, M oscow , U SSR .
Butterwick R esearch Lab oratories , W elwyn , H erts , UK .
C entraalbu reau voor Schim me lcultures, Baarn , The Ne the r
lands.
C zecho slova k Collection of Micr o-or ganisms, J .E. Purkyn e
University , Brno , C zech oslovaki a.
Centre d'tudes du Bou chet , Le Bou chet , France
The C en tr al Laborat o ry , South M an churia R ailway Co .,
Dairen , M anchuria.
C om monwea lth Mycological In stitute (formerl y IMI: Imperi al
Mycological Institute) , K ew , UK .
Centro di Micolo gia del T erreno , T orino , Italy.
Council for Scientific and Industrial R esearch, Pretor ia , South
Afri ca.
D epartment of Agriculture (Mycology), Ottawa , Canada .

EST .
AG R .

NAe. :
ETI-I:

FI E:
FGSe :

FP RL:

II Aee :
IAM :
IARI :

Estacao Ag ro n m ica Nacion al , Sacavm, Portugal.

Eidgen ossisch e T echnische Hochs ch ule , Institu t f r spezielle
Botanik , Z rich , Swit zerl and .
Forstbotanisches Institut , Eb erswalde , G D R .
Fungal Ge ne tics St ock Centre, Humboldt Sta te College, Ar cat a ,
C A, U SA .
Forest Products R esearch Laborat ory , Princes Ri sborough ,
Bucks . , UK.
Hindustan Antibiotics L td., Pimpri , Poon a , India.
In stitute of Applied Mi crobiology , University of T okyo , Japan.
Indian Agri cultural R esearch In stitute , New D elhi , India .

rcr:

Imperial C he rnical Indu stries, W clwyn , Hcrts, U K .

11'0 :

ln st itut c or Fcrm cnt aj io n, Osa ka, .Japan.

Institut d 'Hygicn c , Srra sbo u rg , Fru nce ,
Instituto 'Jaime Ferrn' de Mi crobiologa , M adrid , Spai n.
see C M I.
In stitute of Microbiology , Rutgers , The State U niversity, N ew
Brunswick , NJ , U SA.
In stituto de Micologia , Universid ad e , R ecife , Brazil.
Centro de Biologi a , In stituto G ulbe nkia n de C ienc ia, Oei ras,
Portugal .
Instituto O swaldo Cruz, Rio de Janeiro, Brazil .
In stitut P asteu r , Paris, France .
In stituut voo r Planten ziek tenkundig O nde rzoek, W ageningen ,
The Netherl ands .
International Streptomyces Project.

lB S :
IJ FM :
IMI :
I MR U :

I M UR :
IC C:

IOC:
I P:
IPO :

ISP :
KNC

&

SF :

LC:
LCF:
L M:
LP :

LS H:

G ist-Brocades N .V ., Delft , T he Ne the rla nds.

Laboratoire de C ry ptogam ie, Paris, F rance.
Laboratorio Central de Fit op at ologa , Buen os Aires, Arge ntina .
Labor atorium voor Microbi ologie , Delft, The Ne the rla nds.
M ycotheque du L ab or atoire de Parasitologie de la Fac ult de
M d ecin e , Paris, Fra nce.
London Sch ool of H ygien e and Tropi cal M edi cin e , London ,

U K.
MN HN:
M W:

NC I B :

NCTC:
NCYC:

NHL:
NR R L:

NYSDH:
OEC D :

PC :
PO:
P RL :

QM:

Musum Nation al d 'Histoi re Nature lle, Paris, Fra nce.

In stitut f r Spezielle Bot anik , Abteilung M ycologie , W eimar,
GDR .
National Collec tion of Industri al Bacteri a , Ab er deen , Sco tla nd,
U K.
National Collec tio n of T yp es C ultu res, Lo ndo n, U K .
N ation al Collectio n of Yea st C ult ures, Brewing Industry
R esearch Foundation , Nutfield , U K .
Nat ional In stitute of H ygienic Scie nce, T oky o , J apa n .
C ulture Collec tion U ni t, Fermen tati on Sec tion, No rthe rn U til
izat ion R esearch Branch , U .S. Dep . Agric., P eor ia 5, IL , USA .
N ew York Sta te Dep. H ealth , Alba ny, NY , U SA ,
Organisati on for Econo m ic Cooperation an d Developm ent ,
Paris, France.
Lab or at oire de C ry ptogam ie, Paris , France.
Planten ziekt enkundige D ienst , W agen in gen , The Ne therlands.
Prairie R egion al Labor at ory , Sas katoon, Canada .
Quar terma ster R esearch and Engineering Com mand, Natick,
M A , U SA .

IO,A :
'I'llTC:

I IAMII :

un:

R an cho Sa u ra Ana Boruni c G a rdc ns , C larc m o n t , C A, USA .

Department 01' Botany , Uuivcrsity 01' Toronto , Toronto ,
O nta rio , Cu na da .
Univcrsity 01' Albcrta, Mould H erbarium and C ulture Collection , Edmonton , Albcrta, Canada .
Dep . Food Scicncc and T echnology , Univ . of California , Davi s,
C A, USA .

l1SSR ,
A lU :
WIl :

USSR Antibiotics R escar ch In stitute , Moscow , USSR .

U niv crsity of Wisconsin, Dep artment of Bacteriology , M adison ,
WI , USA .

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haptcr 1

Introduction

'I'III{OUGH THE COOPERATIVE EFFORT OF THE NATIONAL RESEARCH COUNCIL OF

Spain (Consejo Superior de Investigaciones Cientficas), a comprehensive

rcstudy of the genus Penicillium sensu stricto has been conducted by the author
al the Laboratory of General and Applied Mycology of the Instituto 'Jaime
Fcrrn' de Microbiologa, Spain, during the past five years . This has led to the
preparation of this volume covering this very difficult, but important genus of
moulds .
A great number of Penicillium species and strains were examined in culture,
microscopically and under the scanning electron microscope (SEM) by the
author, with the technical assistance in the last task of Drs A .T. Martnez,
Mara de los Angeles Calvo and J . Guarro.
The type species were obtained from the Centraalbureau voor Schimmelcultures, Baarn, The Netherlands, the American Type Culture Collection, the
Commonwealth Mycological Institute, Kew, UK, the Institute for Fermentation, Osaka, Japan, and from our own fungus collection (IJFM).
The members of the genus Penicillium have always been an important factor
in man's environment , but prior to the development of the microscope, they
wcre brushed aside as green, blue-green, ycllow, whitc or brown moulds
without serious attempt at interpretation. Green moulds were everywhere and
names for them appeared in evcry enumeration of the fungi of a particular
locality, or the spccies cncountered in the study of decomposng or fermenting
substanccs . Saccardo (1882) and Wehmer (1894) reported Penicillia active in
the destruction of citrus fruits. Wehmer (1893) reported certain of them to be
active producers of citric acid. Sopp (1898) related them to the che ese industry.
The development of culture laboratories during this period made possible their
isolation and examination in pure culture; subsequent physiological studies led
to increasing interest in thcir presence and significance.
By the 1940s , the Penicillia were being studied intensively in many
laboratories in the United States and other countries . Their potential for the
industrial production of certain organic substances was already well
established; they were being used more and more as research tools for investigating physiological processes and metabolic pathways in microorganisms .
Moulds belonging to this genus are among the most ubiquitous of all fungi .

' I'hcy coustitu to ih c norm al ami a n nh undu nt purt 01' (he mycollu rn 01' nll soils,
th cy play a cti ve mi es in lllallY d ccomposition a nd d ct crioruti ou pmcesscs , th cy

attack man and his dornesti c a n irna ls, and th ey are arnong th e most corn mo n
01' all labora to ry co nta m inan ts . Bcginnin g in th e lat e lBOOs so rne 01' thcm wer c
found to produce interestin g m etabolic produ ct s a nd tod a y a co ns ide ra ble
number 01' spec ies a re used in indu stry 1'01' th e produ ction 01' food specialties,
organ ic ac ids, e nzymes, and a n tibiotics . T h us, m an y peopl e train ed in
differ en t di sciplines a nd workin g in ve ry di ver se fields , mu st conce rn
th emselves w ith spec ies 01' Penicillium. For th ese peopl e , few 01' wh om a re
spec ia lists in m ycological ta xonomy, th ere mu st be av ail able a means 01'
identifying th eir specime n th at is under stable a n d easy to use , an d whi ch at th e
sa me time d oes not viola te th e basi c principI es a nd objective s 01' sound
bot anical nomenclature .

Typification of th e genus Penicillium

The ge n us was create d b y Link in 180 9 1'01' m oul ds p rodu cin g brushlike
spo ru la ti ng structures , three species being recognized a nd described in ve ry
ge ne ra l terms ; P. candidum, P. expansum a n d P. glaucum. Co rda ( 183 7- 1854),
M ontagn e ( 1856 ), a n d othe r m ycol ogists 01' th e su bse que n t 50 years d escribed
add itional spec ies but ge ne rally without definite informati on. Since th e h abitat
01' P. expansum was give n by Link primarily a s rotten fruit, this spec ies m ay be
recogni zed a n d it is no w ge nerally acce pte d as th e Penicillium spec ies ca using
th e rot 01' apples . In 1809 Link co nside re d P. glaucum to be relat ed to P.
expansum a n d lat er (Link 1824) , wh en redescribing P. glaucum, h e listed
P. expansum as a synonym 01' it. In th e me antime Gray ( 1821) validated Link ' s
P. expansum and P. candidum (but not P. glaucum) a n d thus the cor rec t cita t ion
01' the fir st sp ecies is P. expansum Link ex Gray .
An ascospo r ic species was n ot reported until 1874 wh en Br efeld published
his det ail ed report on P. glaucum Link , whe re in he described a nd illu str at ed
scleroti oi d a scocarps . A few m ycologists introduced othe r ge ne r ic names 1'01'
thi s stage . Ludwig ( 1892) wa s th e fir st to propose suc h a separa tion ,
introducing th e n ame Eupenicillium 1'01' the m ould stu d ied b y Brefeld . This was
follow ed b y Langeron ( 1922 ) , wh o, ap pa re n tly unaware 01' th e existence 01' th e
n ame Eupenicillium a nd without referring to Ludwig' s publication , prop osed
th e ge ne r ic n ame 01' Carpenteles 1'01' th e sa me mould . Benj a m in ( 1955 ) create d
the gen us Talaromyces 1'01' th ose ascospo ric sp ecies 01' Penicillium whi ch comprise
th e P. luteum se r ies 01' R ap er a n d Thom ( 1949) . M ore recentl y Stolk and
Samson (1971) cre a te d th e genus Hamigera 1'01' those species 01' Talaromyces
characte r ize d by asci develo p ing singly from croziers, a nd th ey redefined th e
genus Talaromyces by restricting it to sp ecies which produce asc i in chains.
Thom ( 19 10, 1930 ), Cl ement a nd Sh ear ( 193 1) a nd R aper a nd Thom ( 1949)

;)
t ypificd th c gcnus Pen icillium Link, liut su ch typifi cation is not in accordan cc
with the l nt crn utional C ode 01' Bolanical Nom cu clutu rc (St nllcu 1972) .
Accordin g lo Arti clc 1:1 1', thc d esi gnat ion 01' P. ctustaceum Linn acus ex Fri es as
I ype mu st be foll ow ed, becau se Fries (18 32) in th e Systema Myeologieum vol. 3 ,
aeeepted Link's ge n us Penicillium, hut design at ed P. erustaeeum Linnaeus ex
Fri es , based on Mu eor crustaccus L. (with the syno ny ms P. glaueum Link and P.
expansum Link) as the typ e spec ies of th e genus . Linnaeu s based hi s Mu eor
crustaceus on M ieh eli ' s (1 729) ' Botrytis non ramosa, alba, seminibus rotundis ' and
referred to plate 9 1, lig . 3 01' Mieheli's iUustrations . This figure possibly
illu strat es a Botrytis-like er us tose fu ngus whi eh shows sim ila rities to P.
exponsum . Hawksworth a nd Pitt ( 1976) th erefor e proposed th e eonserva tion of
Penieillium Link as th e leetotype . They d esignat ed a neotyp e IMI 39 76 1 = CRS
:125 .48 = Thorn 4852 of P. expa nsum, whi eh becomes the neot yp e of th e
conse rve d ge n us .
No perfeet state is known in P. expansum nor in relat ed spee ies. The gene rie
llame Penieillium is based on an imperfect fun gus and eonseque n tly it
represen ts a ge nus of th e H yphorn ycet es.
H owever, R ap er (19 57) con tes ted th e desirability o f applyin g dual
nom en d ature to PeniciUia , em p has izing the unity 01' all PeniciUia a nd insistin g
th at the ge n us Penicillium sho uld indude both ascogeno us and anascogenous
spec ies. Eve n tho ug h induding both perfeet a nd imperfect spec ies has its
advantages, in accordance with the Intern ation al Code 01' Bot anical
Nomendature (Stafleu 197 2) (A r t. 59) th e ge n us Penicillium sho uld be
res tric ted to th e a nascospo ric species.
The ' M a n ual o f th e Penieillia ' (Rap er a nd Thom 1949), whi ch d escribes 99
accepted sp ecies and varieti es of Penieillium sen su stricto and list s alm ost 700
spec ies, stiU remains now adays th e sta nd ard a nd indisp ens abl e guide to th e
taxon orn y of th e ge n us, d espite a pleth ora o f new species described since its
publica tion . R aper a nd Thorns grasp of relation ships with in th e ge nus was
ex traord inary a nd, by reducing more th an 700 species lo synon ym y , and
forrnin g th e remainder into se ries, th ey m ade it possibl e for erud ite workers to
be reason ably co nfide n t of m ost identification s. H ow ever, beeause 01' th e
paucity of obv ious taxonomic characte rs, and th e arbitrary nature 01' many 01'
th e M anual's gro upings, th e genus Penieillium stiU rem ains taxonom ica lly
difficul t , eve n 1'01' ex pe rts .
M an y new , add itional, species a nd varieties have been deseribed in th e
liter ature, representing sorn e 140 Peni ciUia, and various new surveys have
been published (Abe 1956 ; Kulik 1968 ; Pidoplicko 1972 ; Sam son et al. 1976 ;
Fassatiov 1977 ; Pitt 1979b) in th e yea rs sin ce the ' Man ual' was published ,
and so it is ob viou sly now ou t 01' d at e.
The acc u m ula tion of all this m at eri al seemed suffi eient to th e au thor to
su ppo r t a syste matic restud y 01' aU PeniciUia. As th e present syste m 01'

classifying th esc fun gi is hascd p rimarily on cu ltu ra l a nd morphological

charac te ristics, th e aim 01' ou r 'Manual and atlas 01' th c Peni cillia ' has bcen to
review the gcnus as presentIy dernarcated taxonomically, in orde r lo provide
a basis for th e practical c1assification of those Penic.:illia that are onl y kn own by
their imperfe ct state. *

Cultural a n d m orphologi cal cr ite r ia e m p loyed in the d elimitation and

d es cription 01' specie s
As has been outline d by R aper and Thom ( 1949), th e taxonomy ofthe gen us
Penicillium is ba sed primarily on th e cult u ra l and morphological characteristics
01' colonies grown on standard ized agar m edi a . The applica bility a nd ade quacy
01' th ese principies are reflected partly by th e fact th at th ese author s were able to
re duce th e number 01' spec ies ass igne d to thi s gen us fro m m or e th an 700 to 99
well-d efin ed and ea sily recogni zed taxa , and fu rthe r, by th e fact th at th eir
procedures have now been universally accepted 1'01' th e stu dy 01' th e taxon omy
01' th e Penicill ia .
Ab e ( 1956 ) su ggested th at the u se 01' cultu ral a nd morphological characte r
isti cs sho u ld be su pplemented by th e study 01' bioch emical prop erties 01' th e
cultures, e .g . th eir ability to pro duce antib iotics, th e solubility of pigme n ts, th e
pH changes effected in liquid m edi a , th e colour reacti on 01' cu ltu re filtrates
with ferric chloride, th e producti on 01' fluorescent su bs ta nces, a nd pectinase
ac tivity . While suc h in formati on m ay con tribute to a better kn owled ge 01' th e
physiology 01' these fungi , it is doubtful wh ether th e us e 01' such criteria will be
01' sign ifica nce in th e taxonomy 01' th ese m ould s, particul arl y as m any 01' th ese
proposed b ioche m ica l properties hav e been found to be dep endent on strain.
Accordingly , th e cult ural a nd m orphological criteria introduced by Thom
( 1930) a nd exte nde d by R aper a nd Thom ( 1949) have been ado pted with littl e
modifi cati ons and cons iste ntly applied in the present taxono mical tr eatment 01'
th e gen us Penicillium.
C U LT U RA L CHA RAC TE R ISTICS

The colony 01' a Penicillium exhib its certa in ty pical and striking characte r
istics. T hese inc1ud e colour and colour changes; texture whi ch m ay be velvety
whe n th e stipes arise from th e su bs tratum, lanose whe n th e stipes 01' the conid io
Note . Shortly a fter complet ion of the prese nt study, the book "The G enus Penicillium and its
T eleom or ph ic Sta tes, Eupenicillium and Talaromyces' by John I. P itt (1979b) was published . Th ree
spec ies of Penicillium, P. erythromellis H ockin g, P. dendriticum, and P. loliense Pitt were describ ed in
it as new in the wo rld. These have bee n inc1 uded here in an Addend urn , where they a re
described a nd illust rated applying the sta ndard method s used throug hout the presen t work. O n
th e other hand , a nd as a matte r of conve nience, no acco unt had been m ade her e of Pitt' s new
class ificat ion ofthe gen us Penicillium. For further informa tion , the book itsel f(Pi tt 1979b) should
be consu lted .

phorcs ari sc from simple nc rial hyphnc, [unlculose wh en th e stipc arisc 1'1'0111
prostrat c Iascicles 01' acri al hyphae, uvfasciculatc wh cn th e stipcs ari sc 1'1'0111 crcct
lusciclcs 01' aerial hyphae or sy u nc ma ta . 111 sev era] spccics th c stipcs 01' the con i
. liophores are both monon cmatous and loos ely synnematou s, as in P. expansum .
In P. claviforme and in P. concentricum the synnemata are differentiated into a
d istin et stalk and an enlarged head composed of th e sporulating structures .
Another typical characteristic is the habit, which may be restricted to broadly
spread in g , a nd range from plane to de eply furrowed 01' wrinkled . The
suhstratu rn may 01' may not be dis coloured . Conspicuous drops 01' transpired
lluid a re charac te r istic of sorn e spec ies but not 01' othe rs . Odours may 01' m a y
1101 be produ ced, and if present m a y be charac te ristic 01' just 'mouldy ' .
Although th e gen eral appearance 01' su ch colon ies is influen ced b y the
n-l ative ab u n d a nce 01' conidial structures and scleroti a , whi ch may vary from
s rrain to strain , th e cu ltu ral charac teristics 01' a sp ecies seem to depend on the
use 01' a particular substratum .
C olon ies 01' most sp ecies and strains grow well on th e usu al Czapek' s
solu tion agar. Sorne sp eci es, however, are ch a rac ter ized b y a ver y pOOl'
dcvelopment on this medium. In m ost ca ses su ch restrict ed gro wth m ay b e
attributed to th e in ability 01' th ese strain s to synthe size the vit amins n ecessary
or to utili ze nitrate as a source 01' nitrogen . On th e othe r hand , malt-extract
agar usu all y su ppo rts good growth .
The co m position 01' the medium has a mark ed effec t on th e texture and
su rface ch a rac te r 01' colo n ies. On C za pek ' s aga r m edium the colon ies are
usu all y clos e textured , co m pa ra tively thi ck a nd sometimes va r ious ly furrowed
01' wrinkled . On malt-extract ag ar medium th e co!onies are u su ally loose
textured , co m paratively thin and plane, a nd sometimes a p pear zanat e .
CO LOU R OF C ON ID IA L AR EAS

C olony colour other th an shades 01' blue- green as well as colou r changes a re
ver y us eful in the id entification 01' strains a nd character iza tion 01' species.
Ev ery d escription 0 1' Penicillium em phasize s sorn e colo u r 0 1' se ries 0 1' colou rs
as charac te r ist ic 0 1' con id ial areas. When using seve ral cu ltu re m edi a , it is
necessary to sp ecify th e colou rs upon each by n ame , 0 1' numerical references to
sorne al' th e recognized colou r standards. Ridgway 's 'Colo u r St andards and
Nomenclature ' (191 2) h as been us ed by Raper a n d Thom ( 1949 ) . Much of the
Europe a n lit erature referred to Klinck sieck a nd V al ette ' s 'Code des Couleurs '
( 1908) . Sgu y ' s ' Code Universel des Couleurs ' ( 193 6) was publish ed in
Fr an ce . M ore recently , R a yn er (1970) published his ' Mycolog ical C olour
Chart'. Reference to suc h m anuals will undoubtedl y beco m e necess ary in th e
future since both Ridgway 's , and Klincksi eck and Valette 's works a re out al'
print a n d consequentl y no longer eas ily availa ble .
C olou r records show co ns ide rable variations 1'0 1' particul ar strains a s

int e rp ret ed by dilTe renl individu nls , 01' eve n by th e sa me person workin g a t
diff c rcnt times . For thi s rcaso n cita tion of co lo urs is nev c r a n a bsolu to gu ide. In
gene ral, how ev er, th e colo u rs produ ced by a give n strain upon a sta ndard ize d
su bs tratu m under standard cond itions, fall co nsiste n tly within a narrow range
of shades . H ence, th e cu ltu re medium must be spec ified if the co lou rs cited a re
to hav e real value . O n the other h and , colo u r photographic records 01' colonie s
represent a n excelle n t m ean s 1'01' di scardin g personal interpret ation 0 1' colou rs
an d h a ve been used by th e a u thor throughout th e present stu dy.
The ae r ial m ycel ium 0 1' th ese fun gi is ge ne rally colo u rless. Areas 01' ye llo w
a nd red h yphae co m monly a ppe a r in th e one -stage branch ed Penicilli a 0 1'
R ap er a n d Thom' s ( 1949) Biver ticillat a -symmetrica sec tio n. In th is case th e
colou r ing substance is largel y deposited as granu les on th e surface 0 1' th e cell
rathe r th an with in th e cell itself.
Su b merged m ycellium , on th e othe r hand , m ay sh ow di stincti ve colours
which are best a pp rec ia te d from below a n d a re routinc1y desi gnated as colou r
0 1' reverse. Di rty wh ite 01' dull buffs a re sometimes produ ced , but th e
com monest colours a re var ious shades 0 1' ye llo w, orange, red 0 1' purple . In
sorn e cases th e pi gment produced m a y diffus e into th e surrou nd ing agar
medium. T he di stincti ve a nd bri ght pigments a re parti cul arly ev ide n t in fresh
isolat es.
E X UDA T E AND ODOUR

Di stin cti vc odou rs a n d transpi ration fluids in th e colon ies m ay sometimes be

0 1' va lue as a m eans of identifica tion . H ow ever , th ese characteristics a re 01'te n
va riab le a nd sorne isolat es tend to loose th eir odo u r a nd th eir ab ility to produce
exu date on con tinued cu ltiva tion.
COLONY MARGIN

In a fa vourable su bst ratu m th e colon ies a re usu all y a pp ro ximately circ u la r,

altho ugh m argin al a reas m a y b e characte r istically even, ste lla te, 01' crena ted .
Z O NATI O N

Surface growth ca n be observe d in m any species. So rne 0 1' th em a re zo nate

when gro wn upon one su bs tratu m and not upon othe rs. Ull sch eck ( 1928), in a n
a tte m pt to expla in th is p he no meno n, stated th at colon ies growing rapidl y
secre te by p ro duc ts 0 1' th eir me tabo lism in co ncentrations su fficie nt to su ppress
0 1' reduce gro wt h . The m ycelium th en advances into fresh m edium , resumi n g
rapid growth a nd p roducin g a b u n dant fruit in g structures, only to be depressed
again by th e excess ive b yp rodu cts 01' its renewcd ac tivity .

VAIUATI ON S ANI> M UTAT I ON S

T ite Peni cillia re presc nt a variabl e gro llp 01' moulds. Sectorial m utation s,
loss (JI' ahi lity lo produce spores and t he rnany variati on s whi ch ofte n acc u r,
nlles l lo th e cha ngability 01' th cse fun gi . T hese variations m ay devel ap as a
rcsu lt 01' progressive cha nges under ca ntin ued labaratary cultivatia n. When
I he cha nge cornrna nly rernains stablc thra ugh su bsc q ue nt su bc ultures it may
he rcgarde d as a true mutati on. When the change contin ues to sho w fur th er
chunges in th e same, 01' in sorne other directi on , it is usually regarded as a
vur iai io n .

Mut ants characte rized by colou r cha nges or striking nutriti on al deficien cies
sorne times isolat ed from nature . Seve ral strains have been observed to
dcvclop in a m anner typi cal of well- recognized spec ies upon a n enriche d
mcdiurn suc h as m alt extract aga r, but to gro w sparse ly and very atypically
Il!l0n Czape k's agar medium .
Areas of sterile , or esse nt ially sterile ove rg ro wth m ay a t times occur in
o rhc rw ise typi cal str a ins of almost a ny species of Penicillium. This overgr owth
ra n be isolat ed a nd cult iva ted separately and th en exhibits littl e if a ny of th e
r ha rac reristics of th e par ental strain.
;11'(:

Morphology and terminology of the conidiogenous structures

T he developmen tal seque nce and morphological features of th e fru itin g
structu res are of prime impartance in th e identificati on of fungi. D ist inction
a nd reca gnition of species within th e genus Penicillium is ba sed on th e study of
conidial st ructu res .
T he terms used for the description of th e eleme nts of the Penicillia are
ada pted to th e m odern terminology as esta blishe d at K ananaskis in Canada
( Kcndrick 1971 ) . The term conidiophore is u sed for th e en tire system a f
conid iogenou s cells, tog ether with the differentiated su ppo rting str uc tures. T he
tcrrn ca n id iophore as used by R ap er and Thom ( 1949) is repl aced by stipe.
' r~1E STIPE

Import ant fea tu res of th e st ipes include th eir len gth , septa tion, di ameter a nd
or igin in relati on to the su bs tratum and to eac h ot he r. Althou gh va ria tion in
stip e len gth u su ally occu rs in mo st cu ltures, th e majority of stipes are of a
defin ite ave rage len gth wh ich is characte ristic of th e spec ies . They are often
com paratively short, less th an 100 tut , espec ially whe n th ey arise from ae rial
hyphae a nd bea r sim ple penicilli . When they arise di rectly from th e basal felt
01' su bs tra tu m they are u su ally longer but seldom exceed 250 p.m in len gth .
When long , th ey are sometimes branch ed and ofte n beal' one- or two-stage
branched penicilli . T he wa lls m ay be smooth and th in or m ay be various ly
rou gh en ed , appea ring deli cately echinulate , gra n ula r or asperula te. In sorne

10

cases, suc h as P. roqueforti , the wall s m ay be marked b y co nspicuo us co nc re tions

or warts . D iffer ences in th e st ipe wall a re co ns iste n t in certai n spec ies. In othe r
sp ecies th is character m ay va ry m arkedl y with th e cu lture medium . Separa tion
of co ns ide rab le usefulness, how ever , ca n still be b ased upon thi s character
since ce r ta in spec ies a ppe a r to be ca pable o f develop in g rou ghness under all or
ce rta in spec ific co nd itions, w he reas th e stipes of other spec ies remain smooth
under all co n d itio ns. Stipes a re usu all y uniform in di a m et er , but in so rne
simple forms, R aper and Thom ' s ( 1949) M onoverticillat a sec tion, th e a p ices
are co nspicuous ly swo llen or infl at ed .
The stipe m ay be sim ple a nd unbranch ed or va r ious ly branc he d. It m ay
a r ise frorn vegeta ti ve hyphae withi n th e su bs tratu m or fro m aerial vegetative
hyphae va r ious ly branch ed . It m ay sta nd alo ne, or be m ore or less closely
aggregate d in to clus ter, fas cicles , or defi nit e sy n ne mata .
TH E P E NI CI L LU S

The branch ed part of th e con id iophore a bo ve th e sti pe is refer red to as th e

penicillus. An elabo rate description of all elements comprising th e penicillus has
been give n by R aper a nd Thom ( 1949). O n ly th e sali ent fea tures of th ese
structu res that a re observed in spec ies of Penicillium a re di scu ssed here .
The typ e of penicillus characte r istic to th e spec ies ca n b est b e det ermined by
studying prepa rations wh ich m ak e possibl e the rapid co m par iso n of large
numbers offruiting structu res . This ca n best be d on e by m akin g h abit ske tches
and drawings of co n id ia l structures with th e a id of th e camera lu cid a, at
standard scale, sho wing th e relative dim ensions of penicilli , th e typ e of
branching observe d, a nd the arrangement of th e co n id ia l ch a ins.
In describin g species, or in the ide n tificat ion of unknown strains, the typ e of
penicillus m ost commonly produced must be det ermin ed , since allocatio n to
m aj or sec tio ns is m ad e upon thi s b asi s .
There is for eve ry species a gene ral typ e , nu mber , a r rangement a nd size of
elements which ca n be d escribed in fairly tangibl e te rms. Camera lu cid a
dra win gs sho w in de ta il the pattern a nd cellu lar str uc tures th at are regarde d as
characte r istic of a p a rti cul a r species . The wri tten d escription , how eve r , sho uld
be broad en ough to in clude a ran ge of patterns a n d dimension s th at occ u r.
TH E PHIALIDE

T he term phialide (V u illernin 1910) ac tually replaces th e term sterigmata

in correctIy a pplied b y R aper a nd Thom ( 1949) to th e terminal cells of th e
penicillus which bear th e con id ia (see C ole a n d K endrick 1969 ). It m ay be descr ibe d as a transformed a nd highly differentiated cell with a more or Iess
cylin d r ical body of definite Iength and di amet er which is ch a rac te r ist ica lly

lO. 1.1. A. Typica l simp le

ponl clllus, os soen in P.
lnutuontnns Westling ,
nolstlng of a terminal
vurtlcll of phialides only. B,
vplcul romig enous
penlclllus, as seen in P.
hnrlosl! Smith, cons ist ing of
pro otrote or ascending
tipo s, orising from the
ub utratum or from creep ing
noriol hy phae, typically
rumif ied, w ith rami either
single celled (metula -like) or
long and septate, usually
unequal in length , simple or
rebranched, not producing
defi nite apical verticils of
met ulae or ramu li, w ith
penicilli typ ically simple,
borne te rminally on the
apices of the ram if ied stipes.
e, Typical divaricate
penicillus, as seen in P.
jenthine um Biourge,
con sisting of a terminal
group of irregularly arranged
rami and /or metulae bearing
clust ers of ph ialides (one stage branched) . O, Typical
biverticillate-asymmetric
ponicillus, as seen in P.
expsnsum Link ex Gray
emand. Thom, showing
metulae and rami below the
phialides (two-staqe

o
branched) . E, Typical
biverticillate-symmetric
penicillus, as seen in
Penici//ium variabile Sopp,
consisting of a stipe
terminating in a simple
vert icil of metulae, each 01

which bears a symmetrical

verti cil 01 closelv packed,
typically lanceolate ph ialides.
(a = con idia; b = ph ialides;
c = metulae; d = rami;
e = st ipe of the
conidiophore .)

Latin ampulla = flask) of the ty pe show n in fig: 1.1 A and B. The one -stage
bra nchcd forms, R ap er and Thom's (1949) Biverticilla ta - sym metrica section,
p roduce phi alides that are us ually acerose (from the La tin acerosus = needleshaped ) like leaves of Pinus , as shown in fig: 1. 1 E . A few species prod uce
al most cylindrica l phi alides, lackin g a definite term inal na rrowing (see R ap er
an d T hom's ' M an ual of the Pen icill ia ' (1949) , page 49 , Fig. 12 A l a nd A2) .
T he phialide neck (the con id iu m -be aring tube of R aper a nd Thom 1949)
wa s tcr med collulum by Za leski (1927) , (from the La tin collulum = littl e neck) ,
it is usually long in sorne spec ies (see R ap er a nd T hom's Manual 1949, pa ge
47, Fig. 11 B) or sho rt, and is also of taxon omical im po rta nce .

lO. 1.1. A , Typ ical simple

nlclllus, 8S 900n in P.
quontnns Westling ,
110lBtlno of a term inal
vortlcll of phialides only . B,
vplcul rarnlqenous
ponlclllus , as seen in P.
//IIrlus;; Smith , consisting of
prontrate or ascend ing
tlpos, arising from the
ubntratu m or from creeping
oriol hypha e, typically
rurnlfi od, with rami either
single celled (metu la-Iike) or
long and septate , usually
unequal in length, simple or
rebranched, not producing
definite apical vert icils of
metulae or ramuli , with
penicilli typ ically simple ,
born e terminally on the
apices of the ram ified stipes .
e, Typ ical divaricate
penicillus, as seen in P.
jenthinellum Biou rge ,
cons isting of a terminal
group of irregularly arranged
rami and /or metulae bearing
c1usters of phi alides (onestage branched) . O, Typ ical
biverticillate -asymm etric
penicillus . as seen in P.
expensum Link ex Gray
emend . Thom, showing
metulae and rami below the
phialides (two-staqe

o
branched). E, Typical
biverticillate-symm etric
penicillus, as seen in
Penicillium variabile Sopp,
cons isting of a stipe
term inating in a simple
verti cil of metulae , each 01

w hich bears a symme trical

vert icil of closelv packed,
typi cally lanceolate phialides.
(a == conidia; b == phialides;
c == metulae ; d == rami;
e == stipe of the
con idiophore.)

Latin amp ulla == flask) of the ty pe show n in fig: 1.1 A and B. T he one-s tage
bran ch ed forms, R aper and Thorns (1949) Bivert icilla ta- sym metr ica section,
p rodu ce ph iali des that are us ually acerose (from the Latin acerosus == needleshaped) like leaves of Pinus, as shown in fig: 1. 1 E. A few species produce
al most cylindrical phialides, lackin g a definite term inal narrowing (see R aper
and T ho m's ' Manual of the Pe nic illia' (1949), page 49, Fig. 12 A l and A2) .
T he phialide neck (the conidium-bearing tube of R aper a nd T ho m 1949)
was tcr med collulum by Zaleski (1927) , (from th e Lat in collulum == little neck),
it is usu ally long in sorne spe cies (see R aper a nd Thom ' s Manual 1949, page
47 , Fig. 11 B) or sho rt, an d is also of ta xono mical impor tance.

M I(TUJ.Al-:
Thc ICI'IIl metula (We sllillg 1911) is maint uincd fill' (he pcnuh imut c hrnn ch
wh ich bca rs the w ho rl 01' phialides . Thcy are charac te r ist icnlly borne in a
te rmi nal vcrticil on th e st ipc, or on it a nd cm o ne 0 1' more branch cs . Whe never
the wa lls a l' th e stipe a re smoo th, th ose of th e metul ae are also co ns iste n tly
smooth. W he n the walls 01' th e stipe are pi tted or ro ug h, th e walls 01' th e
m etulae may or may not be sim ilarly ro ughened .
BRA NCH E S

AII cells be tween m etulae a n d stipe s are refer red to as b ranc hes . 1'0 indicat e
th e ty pe 01' branching, th e ter m s pro posed by R aper a n d Thom ( 1949) a re
replaced by th e te rms simple (not branch ed = monoverticill at e), one-stage
branched (b ive rticilla te, as in P. citrinum or species 01' th e Biverticillat asym metrica sec tion), two-stage branched ( = Bive rti cill at a - symme tri ca , as in P.
italicum W chmer, var. italicum Sa mson, Stolk et H adl ok 0 1' P. expansum), or three
or more stage branched, as in P. griseoJulvum.
CON ID IA

T he co n id ia are cndogenous a n d unicellular. Eve ry co n id iu m a r ises

th eoreti call y as a cylin d rical bod y with in a n d extru d ing from th e phialide n eck ,
as in m an y other fun gi with phial ospores. The shape, size a nd scu lp turing 01'
co n idia are ve ry import an t characteristics a l' species. Con id ia m ay be globose,
ovoid or ellipsoidal, smooth or various ly ro ughe ne d a nd co m parativc ly small
or large , but usuall y not exce ed ing 4 .5- 5 .0 uit: in diamet er. Singly th ey a ppe a r
hyaline und er th e light mi croscop e , but in a m ass they show various shades al'
blue- gr een , ye llow or brown . The co n id ia a re p roduced in parall el or di vergent
chains . Bccause the phialides are m ostly a r ra nged parallel in th e sim ple
pen icilli (monovertic illate) , th e con id ia l chains a re 01't en arra nged in co m pact
to ill-d efined colu m n s. When pe n icilli a re di varicate or asy m metr ica l, th e
co n id ia l chains a re usu all y tangled or di vergent. In ce rtain spec ies colu m nar or
tangled masses al' co n id ia form co nt inuous cr usts covering large a reas al' the
colony. In a ny case th e general a r ra n ge men t al' con idial chains in th e m ass is
u su all y recognizable in successive cu ltures a l' each sp ecies under stu dy.
SCLER OTIA

Typ ically th ese cons ist 01' very h ard m a sses a l' pse u do pare nc hy mato us tissue
in which cell walls are ve ry thick. Among th e sim p le Penicilli a, R ap er a n d
Thom ' s Monoverticillata section , th e produ ction a l' ellip tica l to glob ose
sclerotia is regularly characte r istic al' m embers a l' th e P. thomii series. In P.
turbatum al' th e same series, small, soft m asses 01' thi ck-walled cells are usually
encou nte re d . These h ardly re presen t true sclerotia in the se nse al' P. thomii a n d
P. raistrickii, but ac tually th eir presence is st ill accepted as evi de nce al' a

1'('1111 ionship

lo thcse spccic, hcn cc thcir assi f.{IWI ion to t heru.

A ItIIlJIlf.{h IIIt: naturc 01' th csc stI'IICtllJ'(;S is still ohscurc, th cir cxtrao rclina ry
rmH;l lIh la llcc lo 11'111.: nscomnta , in addit ion lo siudi cs ca rricd out by Scolt
I tll)lI) , whi ch lcd lo th c discovcry 01' th e pcrf cct Iorms 01' P. lapidosum Raper el
1;'(;lIl1dl, IJ, ochrosalmoneum Udagawa, P, hyrayamae Udagawa , and P. cinnamo1)/1111/11'1'11111 Abe I!X Udagawa , prcviously known and described a s forming only
clcrotiu, indicntcd lo us t hat sclerotia may be regarded m erely a s abortive
\lll'llllla la , owinf.{ to a gendi c block. Furthermore, to con firm our personal
pedlll 01' vicw, the oppositc 01' Scott's findings , n amely, a strain of previously
dl's('J'il>cd perf cct form , producing abortive ascomata has rec ently been
111 1IH:l'vcd by the author of the present study. Hamigera avellanea Stolk et Samson,
MIl'a ill I.JFM 7239 , isolated in Barcelona by DI' A . Calvo in April 1980, and
Me' llt lo liS Ior idcntifi cation , w as unable to produce as ci, and ye t produ ced the
I ypica l nct work of yellow- encrusted hyphae which forms the en velope of the
lIHl'llllla ta 01' this sp ecics (appearing as sclerotium-like structurcs). With no
d lllll>l , th aboye strain suffcrs a genetic block at th e pathway leading to the
lill'llIalion 01' asci and as cospores. Thus, it will probably be a matler of time
uutil sonicone reports as comata for P. thomii al' P, raistrickii, to m ention only
I WII 01' thc best kn own sclerotium-producing species .

'hapt cr 2

M aterial and methods

I'i\Ri\MO UNT REQlJIREMENT OF DET ERMINATlVE TAXONOMIC METHODS IS T I-li\ T

I hcy provide the maximum a mou nt of information from the fewest possible

lelll s in the minimum time. First must come the cho ice ofmedia : Thom (1910)
1I 1led potato agar and bean agar ; W estling (1911) used prune cxtract gclatine;
lliulIr ge (1923) a nd Zal eski (1927) gr ew their cu ltures on Raulin' s solu tion plus
c ln t ine 0 1' agar. Raper and Thom (1949) em ploye d three standard media:
( :zlIpck ' s a gar , stee p agar , and malt-extract agar. In more re cent studies stccp
If.l lll' has so metimes be en omitted because it is not widely a va ila ble . Abe (195 6)
IIIH:d , in ad d it ion , a Czapek's a ga r m edium with nitrite su b st itu ted for nitratc.
111 Il llld ies on Eupenicillium , Stolk and Scott (1967) added d escriptions of gr owth
1111 co ru -m eal ag ar. Pitt (1973) recommended that th e three basic media uscd
IIY R npcr a nd Thom (1949) for prep a ring co lon ia l a nd morphologieal
dl'llcl'ipli ons be reduced to two : Czap ek ' s yeas t-au tolysate agar (CYA) and
ur ult -cxtra ct a ga r (MEA) . In th e present work we adop te d th e foIlowing threc
Imll ic media , wh ose formula a re given below: Czapek's agar (CA), CYA and
M EA. AIl species cu r re n tly d escribed have been grown on th ese three standard
cu ltu re m edia. AH th e forms gro w well at room temperature (20 -25 C) with
d illusc d a ylight. Hundreds of Penicillium strains , including th e typ e stra ins
ohruined from the most important internation al cu ltu re co Ilections, have becn
x udi cd a nd com pared. The cult u res cite d in th e text by number, bearing th e
prcfix IJ FM , a re accession ed with our own fungu s cu ltu re coIlect ion .

S T i\ ND A R D I Z E D CULTU RE MEDIA

Czapek '5 agar medium h as the foIlow ing co m posit ion (Rape r and Thom 1949) :

N aN03
K 2 H P0 4
M gS0 4 7H 20
KCl
FeS04. 7H20
Su crose
Agar (Difeo)
Di still ed wat er

3 .0 g
. 1.0 g
. 0 .5 g
. 0. 5 g
. 0 .01 g
. 30 .0 g
. 20 .0 s
. up to 1000 m

Tite pH is not udj ustcd a nd tite medium is autoc1aved la r 15 min at 121C

(15 lb ove rprcssurc).
Preparation tim e Ior media ba scd on Ozapek's solu tion muy be reduced by
preparing th e min eral ingredi ente oth cr than ph osphate as 100-fold stock
solution, and adding it to the medium at the rate of 10 mI per 890 mI of water
used .
Czapek's eoneentrate has the fol1owing com position per 1000 mI of solution:
NaN03
MgS04 7H20
KCI
FeS04 ' 7H20

.
.
.

300.0
50 .0
50 .0
1.0

g
g
g
g

This stock solution will keep ind efinite1y ; th e slight precipitate of ferric
hydroxide does not affect th e final medium.

Czapekyeast-extraet agar has the fol1owing com position :

K 2H P0 4
Czapek's conce ntrate
Yeast extract (Difeo)
Sucrose
Agar (Difeo)
Distil1ed water

.
.
.
.
.

1.0 g
10.0 mI
5 .0 g
30. 0 g
20 .0 g
up to 1000 mI

To avoid precipitation of ph osphates and carame1ization of the sugar, th e

potassium phosphate is dissolved separate1y in 100 mI of water and autoclaved
for 15 min at 121"C (15 lb overpressure). The othe r constitu ents are dissolved
in 890 mI distil1ed wat er an d 10 mI C zapek 's concentrate are added to thi s
solution . The solution is autoclaved for th e same time at the same temperature.
After sterilization the two portions are mixed aseptical1y, while stil1 hot.

Malt extract agar has th e fol1owing com position (Raper a nd Thom 1949) :
Malt extract (Difco) . . . . . . . . . . . . . . . . . . . . . . . . . . ..
Glucose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Bacto-peptone (Difco) . . . . . . . . . . . . . . . . . . . . . . . . . .
Agar (Difco) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Distilled water . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..

20 .O g
20 .0 g
1.O g
20 .0 g
up to 1000 mI

The pH is not adj us ted a nd the medium is autoclaved for 15 min a t 121C
(15 lb overpressure) .
Media other th an C zap ek ' s agar were u sed on slide cultu res in certa in
instances wh en con id ia l development on this medium wa s poor or absent:
CYA, MEA and corn-meal ag ar. In su ch instances , wh en ever th ey were u sed ,
they are indicated in the tex t.

'om-meal agar (Di fco), T o rchydrat c th c mcd iu m , su spen d 17 g 01' Bacto r urn-mca l agar (Dif co) in 1000 mi 01' co ld d isti lled wat er and heat to boiling to
d l"Plolve th e mcdium co m plc tcly. Di stribute in tubes and autoclave for 15 min
1I I ~ I oC or 15 lb over pressure. Thc final pH of the medium wil1 be 6 .0 and is
nol udj us ted .
l U I.'I' U RE GROWTH AND INOCULATION MEDIA

C ultu res were routinely grown on MEA. Erratic results occasional1y

f( ,lIowed th e use of old cu ltu res a s in ocula. It is recommended that cultures
q~ e d less than 3 we ek s be used .
G ood results were ob tain ed using sta n d ard, glass, lO-cm Petri dishes
l'o llla in ing a pp roxim a tely 20 mi of medium. Three inoculum points on each
1'IIIIe provid ed replication . As Penicillium con id ia are h ydrophobic , sm all
nu cu la were mo st sa tisfac torily applied as diluted suspensions in a 0 .01 %
iq uco us solution 01' Tween 80 b y m eans 01' a gla ss ball-pointed pipette (fig .
O , I C), wh ich has previ ously been ste r ilize d b y immersion in 95 % alcoho l and
11 11:11 l1amed . The number of conidia in the inoculum a ppa re n tly has little effect
1111 the res u lts.
I':xamination of cultures

1rcscriptions 01' colon ial ch a racte ris tics 01' th e va rio us species studied were
from th ree -spot in ocul at ed m edi a in Petri di sh es. The media rou tine ly
used were C A, CYA , a nd MEA .
For m icr omorphological studies slide preparations were m ade using a n
IlI" e OU S solu tio n 01' 0.01 % Tween 80 as th e m ounting fluid. Small fra gments
of a 7-day-old colony were torn up into ve ry small pi eces w ith ve ry fine
d issecting n eedl es (insect pins fixed on wooden h andles will do perfectly) ,
suspend ed in the abo ye m ounting fluid in test tubes a nd shake n for 2-3 min in
u vortex mi xer a t hi gh speed in orde r to wash ou t th e clou d 01' co n id ia, wh ich
othcrwise wo u ld obscu re the m icroscopi c obse rvat ion 01' th e penicilli.
I lla d e

I NC UBATI O N TI M E

1"01' colo ny m easu rement to b e 01' m ost value , a stan dard in cubation tim e is
r-sscntia l. The author h as chose n 14 d ays as stan dard for colony measu re men t

nnd colour pho tograp hs . H owever, becau se penicilli di sintegr at e with age,
rc nde ring m icroscopi c exam ination diffi cult , a sta n dard in cu b a tion period that
is as sho rt as possible is hi ghl y desirable for th at purpose. 7 da ys a t 25 C is
cons ide re d op timal for slid e cu ltu res.
S UDE CULTU RES

Con id ial structures were exami ne d gene rally on slide cu ltures al'ter 7 d a ys
incubat ion a t 25 C , while th e fu ngus is in a n ac tive state 01' gro wt h . T he

Fig. 2.1 . Equipment. A, Reichter 'Neopan' microscope (Austria) provided with a drawing
device or camera lucida Reichter, Model 21,391 . B, Low power 'Kyowa' binocular (Japan),
with zoom lens, extremely useful for making preliminary observations of colony
characteristics. C, 1 mi inoculation pipette with a closed and bent ball-pointed end.

--

Fig. 2.2 . Equipment. A, An 'exploded' view of Cole and Kendrick 's culture chamber, showing
the drilled and slotted slide, the agar medium, and the coverslips wh ich seal the upper and
lower surfaces. B, View of the Reiss microchamber, showing: (a) the slide, (b) strip of
adhesive paper , (e) side parts of the chamber, (d) the coverslip, and (e) the inoculum. C, An
'exploded' view of the drawing device: (1), Camera lucida . (2), Fixing screw to affix the
camera lucida to the microscope. (3), Ad justing ring for focusing . (4), Fixing screw, to affix
the camera lucida eye-piece. (5), Camera lucida prism's casing. (6), Screen for cutting off the
path of light through the camera lucida 's prism when the apparatus is not in use. D,
Schematic diagram showing the basis of the camera lucida theory.

medium gc nernlly cm ploycd was C A . Iowever, when couid iul dcvelopmcnt

o n thi s su bstra tum was (mor , OYA, M EA , 01 ' co rn mcal agur wcre uscd
in stcad .
C U L T URE C HAM B E R S

Other techniques may be us ed 1'01' th e study 01' org an ization and ontoge ny 01'
ae rial str uc tu res . M any 01' them have been em ploye d 1'01' thi s study . In order to
ens u re aeri al gro wth, th e fun gu s is u su all y gro wn on slides on a thin film 01' an
appropriate su bstratu m. The conid iophores will gro w off the sides 01' th e
m edium in to th e surro u nding a ir and ca n thus be exam ine d (see V ernon 1931 ;
Riddell 1950 ; Anton y and W alk es 1962 ; C ochrane 1963; Zo be ri 196 7) . Cole
and K endrick ( 1968) described a sm all cult ure cha m ber 01' glass (fig. 2.2 , A)
whi ch all ows cultiva tion 01' th e fu ngus in suc h a m anner th at th e develop m en t
01' fu ngal struc tures ca n be contin uo u sly observe d and even film ed under hi gh
m agnifi cat ion . The constr uctio n 01' suc h a chambe r, however, requires great
per sonal skil!.
Consi de ring th at m any conidiopho res are gene ra lly negati vely geo tro pic
(Coch ran 1963), an eas ily producibl e and in exp en sive cult ure cha m ber was
described by R eiss (1970) (see fig . 2.2, B) , whi ch allows the study 01' the
development 01' re pro d uct ive fungal str uc tures in an undisturbed po siti on.
Su ch a cham be r has been used in our lab oratory with success . On a clean slide,
two small str ips 01' glass (the thi ckness 01' a slide) are glue d with nontoxic
mucila ge (i.e . U.H.U from Fisher, B hl , FRG ) at a dis ta nc e 01' 14 mm from
eac h othe r. This frame is ste rilized . A ste rile coverslip (18 x 18 mm) is placed
ayer the gap bet ween th e two glass strips a nd fixed to th em with two strips 01'
adhesive tap e . A small piece 01' a Petri di sh cultu re 01' th e fungu s is removed
under ste rile cond itio ns a nd placed on th e slide in fro nt 01' the spa ce bet ween
th e cover a nd th e slide . With a small and sterile strip 01' glass thi s in oculum is
pushed between th e slide and th e cove rslip . One sid e 01' th e op en cham be r is
closed with the strip 01' glass, th e othe r remaining ope n to ens ure a su fficient
supply 01' a ir.
T o enable negative geo tro pic gr owth 01' con id iophores, th e ino cul at ed
cham be r is put in a ve rtical position into a ste rile tr ou gh 1'0 1' slides , th e bot tom
and side s 01' whi ch are lin ed with sterile wetted filter papel'. U nde r th ese
cond itions rep roducti ve structures ca n grow upright int o th e air. The tr ou gh is
in cubat ed at a n appro pria te temperature (20 - 25C) under d iffuse light. Al'ter
varyi ng periods th e cham be r is removed and exam ine d under th e m icro scop e .
When th e mi croscop ical study has been fini sh ed , alco hol is dropped into th e
chambe r , th e coverslip is removed , cleaned and sterili zed , together with th e
lower part of the cha m ber. U se ofthe stro ng illuminati on necessary 1'01' m aking
mi crophot ogr aphs ca uses fine d rops 01' conde nsation occasionally to arise on
th e inner side 01' the cove rslip, but th ese evaporate sorne time afterwards .

,' ('tlll ll imr electro n microscopy

l'h c scnn n ing elec tro n mi croscop e (SE M), as a n a id in th e idcntificntion 01'
nllo wiu g' thc obs c rva tio n 01' su rfa cc o rna m e n ta lion 01' va rious o rga ns, is
IiI': t'cIIJlillf{ iu c rca sin gly available . T he SE M stu d ies by C la rke a nd G riffiths
1\)70) a nd Lo cci a nd Q uaro n i ( 1970, 1971 ) ha ve dcmonstrat cd as co spo rc -wall
IJI'I IIlIIIC lIla tio ll in most spe cics 01' pcrfect Aspergilli i Eurotium, Emcricella,
8rll fo'Ya) , As a rcsul t , so rne hitherto unknown mi cromorphological Ica turcs 01'
IIU'. I ~ p ()rcs 01' :17 species a nd va ri e ties of th ese fun gi have b ecn rcco rd cd. T he
1'.' ",ilIS obt uin cd suggest a need for so rne tax onorn ic revi si n in o thcr
1'.'lllIclIllica lly important grou ps. This is cc rta in ly th e case for th e gc n us
l ruicilliu m, in wh ich th e con id iu m ' s morphology a n d su rfac e o rnam c ntatio n
Il'c' very iruportant ch a ra cter istics in th e identifi cation 01' th e diff ercnt species.
1"111'1hcrmorc , su r face o rname ntation is cons iste n t a nd relia ble , bu t dilf icult to
d.' lc:J'I l1 inc as th e co n id ia a re hyaline , u su all y ve ry small , a nd freq uently ca rry
r m u plicutcd scu lpt urin g. The sca n n in g e1ectro n mi cro scopc reveal s thc fin e
mu .u uc n ta tio n of co n id ia in much m ore det ail th an does th e light mi cros co pe .
Wl' rhc rcl orc , h ave b een u sin g th e SEM in ou r labo rat o ry as a n a dd itio nal
II11 'a llS or th e char acter iza tio n a nd description of co n id ia o f a ll Penicillium
IlI'l'ic:s stu d ied .
' I'hc manipul ation of hundred s of sam ples m ade it necessa ry to look Ior il
l'lll' id method of handling a nd preparin g th e se , witho ut loss in im a ge quality.
AI't I'l' m an y tri al s , th e foll ow in g procedure * was found to be m ost e ffcc tivc :
'1'111' 1'1I11gi were cu ltiva ted u pon MEA , CA 0 1' C Y A for 1- 2 weeks . A lte r
l'illl'lIillg of the co n idia , small 3% agar bl ock s (abou t 2 x 3 x 6) m m in
rlluu-ns ions we re ge ntly pre ssed a ga in st a well-sporulat ed a rea of th e co lo ny, to
llx a great number of co n id ia to th e a ga r su r face . The agar blocks were th en
iutnx luced se parately into Beerns ca psu les (Balzers Union , Li echtenstein) ,
illlo w hich sev eral sm all holes a re d r illed . A modification of the Sam son and
SI:.Ipcrs ' technique s ( 1977) for th e prep aration of fu n gal specimen s for
HI':I1111ing e1ectron mi cro scopy a re ca rr ied out as follows :
Ill ll ~ i,

Fixntion : 6 % glu ta r ald eh yde ( 12 h in th e d ark, under m oderate vacu u m,

111

ordcr to e1iminate air bubbles)

' I'wo successive washings with a n a q u eo us solu tion 01' 0 .0 1 %
T ween 80 (2 :10 min)
' I'wo suc ce ssive washings with d ist illed water (2 : 10 m in).
ltchydrauon: M eth yl cellosolve (2 -methoxyethanol) ; three su ccess ive chan ges of
t hc dehydrating a gent are n eed ed ( 1: 10 : 10 min)
T h ree suc cessive washings with 100 % acetone (1: 10 : 10 min) .
Mar tn ez a nd R am rez' s un pu blished No te, presented a t the IV Int ern at ional Conference on
Cu ltur e Co llectio ns , BR NO , Czechoslova kia, 20 - 24 J uly , 1981.

1tI'l'I CAI. I'O INT OIWI NO IN

'-' V ~

T his step is necessary to com plete th e dehyd rution proccss, lo avoid colla pse
01' fungal cells a11(1 sh ri nkugc 01' ccll wall lay ers. A Sumd ri-Put -S a uto matic
a ppa ratus, with a 15-min purgin g tim e was uscd by th e auth or , Bcfore coa ting
(the spec ime ns are shadow cas t with palladium - gold (40 - 60 ) in a vacuum
eva po rato r), th e sa m ples we re tak en from th e ca ps ules and fixed to a srnall (5
mm di am .) tin di sc with a n electroco nd uc tive silve r glue (Do lite) . The tin d iscs
may be ea sily m arked underneath for identificati on , and fin all y placed on a
sta ndard specimen stub by m eans of th e same silve r adhesive m ixture .
The con id ia were viewed a nd photogr aphed with a J EOL jSM-50A
sca nn ing e1ectron mi cr oscop e at an acc ele rating voltage of 15-20 kV.
CAMERA L U CIDA DRAWINGS

Microscopic observation s and ca m era lu cida drawings were made with a

R eichter 'Neopan ' microscop e (Austria) (fig. 2 .1 , A) , with th e followin g
objectives: 10/0 .25; 25/0.45; 40/0 .65; and 100/1.25 (oil immersion obj ecti ve),
a nd two BPK 10 x eye-pieces .
T he drawing device or ca me ra lucida was a R eichter ' Ze iche nap para t'
M odel 21 ,391 (fig . 2.1 , A , and fig. 2.2 , C), provided with a PK 16 x eye
piece . The drawing apparatus is int ercalat ed betw een th e m ain body and th e
observa tion tubes of th e mi cr oscop e (fig . 2.2 , C ).
In order to obtain a standard magnification of 1500 x with th e oil immersion
objectiv e, th e di stance between the camera lucida's prism a nd the drawing
papel' was det ermined . It sho uld be 284 mm (see fig . 2.2, D ) . This di stance
was ob ta ine d by placin g th e mi croscope upon a perspex block (250 x 250 x 45
mm) (fig. 2.1 , A) .
AH ca mera lucid a drawings of mi cro scop ic characte ristics of th e typ e strai ns
stud ied, grown on C za pe k's agar or on one of th e other sta ndard m edi a
previously m ention ed , were ca rried ou t under the oil immersion objective at
the m entioned 1500 x magni ficati ons. H abit ske tche s, on the ot he r hand , we re
drawn by eithe r exam ining 7-d ay-old spot cultures direct1y on th e stage of th e
mi cr oscop e 0 1' 7-d ay-old slide cultures, at a m a gnifi ca tion of 150 x , by m ean s
of th e 10/0 .25 low pow er objective .
OR IG IN OF CULTU RES

The m aj ori ty of typ e strains stud ied were obtaine d from th e Cent raalbu reau
voo r Sch im m elcultures (CBS), Baarn , The Ne the rla nds. O the r were obtained
from th e Ame rica n T yp e C ulture Collection (AT CC) , R ockville , M aryland ,
USA. A few stra ins were obta ine d from th e Com mo nwealth M ycological
In stitute (C M I), Kew , U K, a nd from the In stitute for Fermen tatio n (I FO),
O sak a , j apan . Additional strains of new or prev ious ly describ ed species were
secu re d fro m numerous fresh isolates. The latt er were chec ked both cult ura lly
a nd mi croscopically against th e type stra ins.

1'1 (~ "I I'V/lt.io n

0 1'

pcnicillia

t\ 11('I'IIOIlII FO il MAINTAININ l: STOCK ClIL'I'lI llI':S

\ II Y inc rho d lor muiur ainin g a stock cultu re co llcctio n mu st kecp th e
illHfl lllMIII via b le ami in pUl"e cult ure ind efinitcly . '1'0 be fully sati sfa ctory ,
ln rwuvcr . il should also maint ain th c origin al characte rs of th e strains , and
IllIuld rcq u irc a minimum 01' tim e , cq u ip rne n t, m at erial a nd space.
Srlve: I'a l ,{c ne ra l typ es o f sro rage are at present in u se for th e cu lt u re offu ngi:
1) S llIl':I gC: undcr co nd itio ns whi ch perrnit m et abolic ac tivity to go on a t a
' 1IIII'llIa l' rat e o Thc cu lt u res a re m aintained o n a nu trien t aga r m edium
11 111' ncnr th ei r o ptim u rn temperature a n d tran sferred a t frequent in te rvals .
Sllll'a ge undc r co nd itions wh ere m etabolic ac tivity is restrict ed. This is
u-hicved ci thc r by stor ing aga r sla n ts a t 5 - tOC (the c1a ssical m ethod ) , by
'IVC'l'illg slant s with ste r ile mineral oil (Buell a nd W est on 1947 ), or by
d iNpc:l's ing spo rcs in so il (B ak erspi gel 195 3, 1954 ) .
'1) S llIl'age eo nd itions wh ere m et aboli c activity is pre sumably ab se n t, is
1ypil'a lly ac hieved by th e Iyophil or free ze-dryin g process , o r other drying
'I'l 'c hlliq ues (H arris 1954) .
1) Tl : Irc czin g' method d escribed by C a rm ichael ( 1956), which co nsists of
Mla lll cu ltu res IIf th e fun gi pl aced in a de ep-freeze ches t a t - 20 C , ac h ieves
dH' d csi rcd in acti vation of m et aboli sm , but is no m ore time- consuming th an
d H' usu al pra cti ce of sto r ing sla nts in th e refrigerato r.
AII mcth od s ca n m ain tain via b ility . H ow ever , th e fir st m eth od favours
vlI l'i:11io n a nd is ti m e-con sum in g for large co llec tio ns . T he seco nd method
1"'l'llIils lon ger interval s between transfers , but is st ill ope n to cr iticism in
1'1 '111 1ion lo strain var ia tio n a nd conven ience . The third m ethod , which
1l lldlil,ilS nuclear di vi sion , is theore tically th e best for maintain ing origin al
Mll'a in ch ara ct ers , but d ryin g tec hniq ues require special eq u ip m en t a n d ca n not
111' .ip plicd lo a ll fun gi (Raper a nd Al ex ander 194 5; Fennell et a l. 1950).
( :l l NT ll O L OF I N F E ST ATI O N OF C U LT U R E S

( :u llurcs kept in a state of ac tive m et aboli sm rathe r th an under induced

c1c ll lIla ncy, as w he n m ain tained as Iyophil o r in liquid nitro gen , a re suscep tible
1II inlcstatio n by m ites . In fa ct , thi s is one of the mos t con sta nt so u rc es of
111I11I>l e in a m ycological la bo ra tory . Once h avi n g gained access to the
lnho ratory , mites make th ei r wa y in to P etri d ishes 0 1' th ro ugh cotton- wool
pill gs into th e cu lt ure tu bes . H ere th ey feed on the grow ing fu ngi a nd in
pa rl icular on th e spores a nd fru ctifications . A part from destroyin g the
fl'lJ(' 1ifi cation s th ey in tro d u ce fu ngal a nd ba ct eri al co nta m in a nts in to th e
u ltu res . The fact th at they can wan de r quite ra pi d ly from cu lture to cu lture
uu -an s that , followin g a mite infestatio n, a who le collcctio n may be
ruut arninar ed a nd a n immen se a mou nt of wo rk is required to ob ta in pure
rul tu re again .

Miles occur in n utu re on decay ing plunt nuu eri al , on gra in , flour, chcc sc,
du st a m i so il pa rticles. T hosc infectiug lun gal cu ltures most co m mo nly bclon g
to suc h gen era as Tyroglyphus and Tarsonemus. They are most cas ily co n tro lled
by high standards of hygiene in laboratories whi ch hav e a screen in g procedure
1'01' in coming cu ltures a nd whi ch do not have to handle or ga n ic m at erial. In
laborat ories whi ch h a ve to handle fre sh plant m at erial , soil, b at ch es 01' grain 01'
m anufactured st u ffs, precautions must b e taken against mites, particul a rly in
humid c1imat es . W e found that storage 01' co n tam inate d m a terial in th e deep
freeze chest 1'01' 8 h is sufficient to kili all mites.
C HEM ICAL AN D PHYSI CA L B ARRI ERS AGAI NS T MIT E I N F E ST ATI O N

Snyder a n d H an sen ' s m ethod (1946) h as been u sed routinely in our

laborat ory with exce lle n t results. It cons ists 01' th e sealing 01' cu lture tubes with
ciga re tte papel' to provide a barrier agains t mites, bu t wh ich allo ws a ir to
diffuse th rou gh 1'01' th e gro wt h 01' th e cu ltu res . This is a m ost effective m ethod
which has no effect on the fungal cu ltu res and a l't er many ye a rs 01' use , I am
able to di scount th e op in ion that it is lab ori ou s (Sm ith 1967 ). T he m eth od
requires only a copper su lphate gela ti ne as ad he sive a nd stand a rd cigare tte
p ap el'. T he ad he sive cons ists 01' a 20 % gelatine in wat er to whi ch 2 % co pper
sulphate h as been adde d . About 25 mi 01' the m elte d mi xture is p oured into a
Petri di sh a n d allowed to solid ify . T he ciga rett e papers are unfolded a n d cu t in
h all'. Al't er th e cultu re h a s been m ade , th e cotton- woo l plu g is flamed a nd
pushed d own into th e tube . The rim 01' the tube is a ga in flamed a nd wh ile still
hot is pressed in to th e gelatine -copper su lph ate mi xture , removed a n d
immediately pressed on to th e ce n tre 01' h all' a cigarette papel'. The papel'
ad he res to the end 01' th e tube an d it can be p ressed down if a ny imperfect ion s
a re observ ed . The tubes are placed with th e top ove r th e edge of the be nch , a nd
wh en a series is com plete a nd the gelati ne adhesive ha s set, the su rplus
ciga rett e papel' ma y b e folded arou nd th e tube ' s e nd and held in place b y a
strip 01' parafrlm (Amer ica n Can Company , Marathon Product s, N eenah ,
Wisconsin , USA).

Q.)

a.
o
1
a.
ro
::J

c:
ro
E
Q.)

.....

..c:

lhap ter 3

'1'111'. I'U RPOSE OF THIS V OL UM E I S TO FACILlTATE T HE ID ENTI FI C ATI ON OF ALL

l'auicillium species reco gn ized to date , presenting full descriptions, analytical

kl'Ys, colou r ph ot ographs sho wing th e cult u ral characteristics 01' th e colonies
1'1 JW II 011 three sta nda rd m edi a in th eir natu ral colo u rs, standard scale ca me ra
Illd da lin c drawin gs al' frui tin g st ruc tures showing de tails, pattern a nd
rlluu.nsio ns 01' essen tia l cells, a nd fin all y sca n n ing electro n mi crogr aph s 01'
t'ollid ia , show ing the ir fin e su rface orname n ta tion , so th at th ose interested in
dl'1I1ifyin g eas ily th ose species 01' Pen icillium not included in R aper and T ho m' s
I!H!)) 'Manu al 01' th e Penicillia ' , will not hav e to search th e lit erature for
dl's(Tiplions a l' the m .
111 co ns tr uc ting the keys, th e chief objective has been to simplify search ing
li 11' I he description 01' a give n spe cies . Conseq uently , th ey sho uld not be
('lJlIside rcd as a ph ylogen etic treatmen t.
Ce lle ra l key t o the sections

Page

1. Ponicilli consisting of a single cluster, or verticil of phialides at the tip of

Ihe stipes; stipes usually unbranched, in some forms irregularly
branched but with each branch terminating in a distinct and separate
simple penicillus.
MONOVERTICILLATA Section

28

11. Penicilli characteristically once or twice branched below the level of

phialides ; typically asymmetrical, irregular, or one sided; phialides not
acerose or needle shaped.
ASYMMETRICA Section

201

111. Penicilli characteristically biverticillate and symmetrical, but sometimes

fractional in some species and strains; phialides typically lanceolate or
acerase, with long -tapered, acum inate necks.
BIVERTICILLATE-SYMMETRICA Section

507

IV. Penicilli large, usually symmetrical, typically branched at three or more

levels below the phialides.
POLYVERTICILLATASection

611

The Monoverticillata

( 1949) WAS T HE
M onov erticillata section. They included in it both th e perfect and imperfect
sp ecies in an a ggregat e of seri es a nd species bound together by th e a rbitrary
fact th at th e penicillu s was simple, i. e . it cons isted of a single clu ster of
phialides at th e tip of th e conid iopho re.
In th e present work , th e auth or m aintains th e Monoverticillat a section, but
in order to be in accordance with th e Internation al Code of Botanical
N omenclature (Sta fleu 1972) , al! species with perfect sta te s cons idered by
R aper a nd Thom (1949) in thi s sectio n have been exclude d fro m it.
The m ain characte ristic of thi s gro u p is th at th e penicilli co ns ist of a sing le
cluster or ve rticil of phialides at th e tip of th e stipe ; u su all y unbranched ; in
sorne forms irregul arly branched but with eac h branch terminating in a di stinct
and sep arate sim ple penicillus .
Few sp ecies are strictly simple , that is , produce stipes without branch es
below th e ve rtici l of phialides. In a sin gle se ries, design ated by R aper a nd
Thorn (1949) R amigen a , the stipe is nearl y always branch ed at various sep ta,
with the branches vary ing in len gth and usuall y di verging, but eac h branch
m aintaining th e a ppe ara nc e of a strictly simple penicillus .
The Penicillium thomii series of the Monoverticillata section is ch aracte r ized
by the production of true sclerotia, which in appearance and texture may
simulate cleistothecia .
Separation of th e simple forms from th e remainder of th e Penicillia is based
on examination of th e terminal verticil at th e tip of th e stipe. If there are no
branch es a t th e a pex of th e stipe , or if each branch maintains a fairl y definite
individuality , i.e . produces a simple penicillus, th e species is assi gn ed to th e
Monoverticillata section . However, no sharp line of sep aration can be drawn
between th e M onoverticillata and other species of R aper and Thom ' s ( 1949)
As ymmetrica section . In fact, th e different sections tr eated by th em are
intimately related and represent tr ansitions and different facets of a large and
variable group of moulds . As a matter of fact , com plete bridgings ca n be
observed between th e Monoverticillata section and th e Velutina sub -section of
th e As ymmetrica sec tion throu gh th e R amigen a series and Penicillium
corylophilum in the P. citrinum series . A similar transition from the Mono
verticillata to the Divaricata sub-section of th e Asymmetrica section occurs
through P. j ensenii.
T HE FIRST GROUP O F P ENICILLI UM TREATED BY R APER AND T HOM

:.W

thc seri es (Moc1ilicd Ir orn R ap er and Thorn 1949) Page

pro duclng sclerotia, oftan suggastive of young c1eistothecia
P. thomii series ooo
not producing sclarotia .

1, nl/pOO gonorally unbranched and bearing single, strictly simple

29

prmlollll.
onloa volvety or nearly so; stipes arising mostly from the sub
trntum.
1' . Colonias ganerally spreading broadlv ,
111 . Conidia globose or subglobose
P. frequentans series o. oo 50
bb. Conidia elliptical
P. /ividum series oooo 70
' . Colo nias growing rather restrictedly, especially upon Czapek's
linar
P. imp/icatum series oooo 82
1.1 . Colonies appearing velvety or slightly lanate, but with the stipes

horne as short branches from interwoven aerial hyphae

P. decumbens series oooo 102

'. Colonies lanate or lanate-funiculose with the stipes arising
primarily from aerial hyphae.
l ' o Colonies predominantly lanate with funiculose habit lacking or
limited
P. restrictum series oooo133
2' o Colonies with funiculose habit predominant or well developed
Po adametzii series o. oo 155
. S tipes mostly irregularly branched but with each branch bearing a
wellmarked simple penicillus
The Ramigena series oooo 185

Pmicillium thornii series

)IJ TSTA N DI NG FEATURES

Sclerotia usually produced ; they are typically hard and gritty, consisting of
vCl'y hard sclerenchyrna-like rnasses of parenchyrnatous tissue with thick
wnllcd cells . In sorne species th ey consist of cornparatively soft rnasses of thi ck
wullcd ps eudoparenchyrnatous cells.
C olon ies usually spreading rapidly, developing abundant sclerotia which
o ftc n cha racter ize the colony appearance o In sorne species, howev er, colonies
~ l'Ow slowly and develop sclerotia only on certain substrates .
C on id ial structures abundant in sorne species , not in others ; with stipes
a l'ising directly frorn the substraturn, producing a velvety appearance, 01' frorn

trai ling vcgc ta tive hyph ac; va ry ing frorn short IIp to 300 - 400 J.'m in Icn gth ,
with walls smoo th 0 1' dclicatcly cchinulnto.
Pcni cill i str ic tly sim ple. H ow cv e r, so m c tim cs l!ley rlla y show a n occas io nal
bra nch , T he y a re spa rsc in sorne specics a m i stru ins . Phi alidcs a re typically
p a ralIel in co m pact clus te rs.
C onidia m ostl y elliptical to subglobose , srn ooth wall ed , usu alIy rel atively
smalI, 2 .5 -3 .5 .tm in di amete r , fo rmin g lon g cha ins often a d he re nt in loase
colu m n s.

Key to spec ies

Page

a. Sclerotia hard , brittle, crushing with difficulty, composed of thick

walled sclerenchyma-like cells
P. thomii series . . . .
1'. Sclerotia produced upon all substrata
aa. Sclerotia uncoloured or nearly so, borne in small clusters sur

rounded by conspicuous envelopes of bright orange-red hyphae

P. sclerotiorum van Beyma . . . .

bb . Sclerotia in flesh to pink shades, not in clusters and not
embedded in masses of orange-red hyphae
P. thomii Maire . . ..
cc. Sclerotia dark brown to black, often surrounded by a loose

network of dark hyphae

P. syriacum Baghdadi . . . .
b. Sclerotia soft, composed of pseudoparenchymatous thick-walled cells;
sclerotia produced only upon certain substrata
P. turbatum sub -series. . ..
1' . Colonies upon Czapek's agar developing rather restrictedly
aa. Colonies upon Czapek's agar not showing dull, dark purple
colours in reverse; growth somewhat restricted
P. turbatum Westling . . . .
bb. Colonies upon Czapek's agar showing dull, dark purple colours in

reverse ; growth very restricted

P. pusillum Smith . . . .
2'. Colonies upon Czapek's agar developing fairly rapidly
aa. Sclerotia produced upon all substrata
1". Colonies upon Czapek's agar showing brown colours in reverse;
sclerotia white, surrounded by brownish submerged hyphae
P. donkii Stolk . . ..
2". Colonies upon Czapek's agar not showing brown colours in
reverse; sclerotia white to cream coloured, not enveloped by
vegetative hyphae
P. indicum Sandhu et Sandhu . . . .

29

34

31

36

39

39

41

44

42

bb , Sclarotla davaloplng only upon cartaln substrata

P. gr8nc8n8ri8e Ramlrez, Martlnez et Ferrer . . . . 46

llricfly ch a ra ctc r ized , th e Pen icillium thom ii ser ies co m p r ises a gro u p of sim ple
Puu icilliu which regularly produ ce sclerotia . Raper a nd Thom ( 1949) sub
lvidcd lh e series, as a matter of co nve n ien ce, as follow s: (1 ) The series proper ,
in cludi ug a ll species produ cing a bu ndan t sclerot ia upon a ll su bs trata, whe rei n
1111: scle ro tia are ve ry h ard , brittle , a n d crush w ith difficulty ; (2) The P.
turbatu m sub-ser ies, in wh ich scle ro tia a re no t p rodu ced upon sorne su bs trata
( iud lld ing Czape k's agar), a n d are softer a nd re prese nt less co m pact m asses o f
ihick-wullcd ce lls .
'I'hrce spec ies ac tually co m prise the ser ies proper ; Penicillium thomii Maire is
l he oldcst, best kn own, a nd m ost a bu ndant species, appare ntly world -wide in
listribu tion. P. sclerotiorum van Beyma is less w ide ly di stributed a nd was
llI'i!{i na l1y isolated in J ava (Indonesia). It h a s been obtaine d fro m ot he r
su u rce s , rnostl y of tropical or igi no P. syriacum Ba ghdadi , isola ted from Syrian
d cscrt soi l, is characte r ized b y it s rather restricted growth upon a ll med ia, and
by thc p rod uct io n of b ro wn to blac k scle ro tia.
Fivc speci es ac tually comp rise the Penicillium turbatum sub-series, whic h are
ch uru c re r ized b y the p roduction of softer a nd easily crushed scle ro tia . P.
lurhutum W estling a n d P. pusillum Smith , bo th characterized by thei r rest ricted
g l'llw l!J upon Czapek's agar a nd P. donkii Sto lk, P. indicum Sand h u et Sandhu,
II UI! P. grancanariae R amr ez , M artn ez el Ferrer , characte r ized b y the ir fairly
I'lIpid growth upon Czapek's agar. Altho ugh the degree ofrelations h ip between
1hcsc spec ies and the members of the P. thomii series proper is q uestionable , it is
hcl icvcd , nevertheless, that th e y ca n be keyed here more sa tis factorily than
clscwhc re .
1, Pe nicillium thomii Maire : fig. 1, colou r plate 1, and SEM plate 1 (top :
right).
Penicil/ium thomi Maire - Bull. SOCo Hist. Nat. Afr. Nord. 8: 189-192. 1917. See
also Thom , U.S. Dep. Agric. Bur. Anim. Ind., Bull. 118: p. 78. 1910; also Thom,
'The Penicillia', p . 73 . 1930; also Raper and Thom, 'A Manual of the Penicillia',
p . 156. 1949.

Synonym ies
Penicil/ium crocicola Yamamo to apud Yamamoto, Maeda et Oyasu - Sci. Rep.
Hyogo Univ. Agric. Biol. Ser. 2: 28. 1956.
Penicil/ium paral/elosporum Sasaki - J. Fac. Agric. Hokkaido Univ. 49: 147. 1950.
Penicil/ium quercetorum Baghdadi - Nov. Sist. Niz. Rast. 7 : 110. 1968.
Penicil/ium thomii Maire, varo flavescens Abe - J . Gen. Appl. Microbio!. Tokyo .
2: 51. 1956.

nly (colou r piat e 1, A) , and ofte n failin g to devclop sclerotia, es pcc ially in
trui ns lon g m aint aincd in artifici al cult u res ; c xud a tc clca r . a bu ndantly
pn ulu ced in so rne strains, not in others; odour aint , sugge sting mushrooms ;
l't:V CI'SC (colour plat e 1, a) in pal e yellow to pinkish shades ; stipes 0 1' th e
r n uid iopho res a r ising primaril y frorn the substratum a n d from interlacing
1l:l'ial hyphae , seld orn branched , with walls delicately ec hi n u la te (fig. 1, A) ,
variable in length up to 300-400 J.l.m by 3.0-3 .5 J.l.m in diamet er, with a p ices
Hlightly c nla rge d in a ves icle abou t 4.0- 5. 0 J.l.m in di amet er; penicilli strictl y
ximplo, bea rin g co n id ial chains usu all y in loose colu m ns up to 150 J.l.m 0 1' more
in lcn gth (fig. 1, C); phialides m ostl y parallel , in cro wde d clu sters, 8 -12 in th e
vcrticil, usu all y 8- 13 J.l.m by 2.5- 4.0 um , with a somewhat n arrowed , sho rt
ucck (lig. 1, A) ; con id ia elliptical to subglobose (fig. 1, B, a nd SEM plate 1,
lop : right) , m ostl y 3. 0-3 .5 J.l.m in lon g axis, app arently smooth wall ed when
oh scrved under th e light mi cr oscop e but rough wall ed when observe d under
Ih e S EM.
Co lonies on CYA spreading broadly , a tt ain in g a di amet er 0 1' 45- 65 mm in
1,1 d ays at room temper ature (20- 25 OC ) (colour plat e 1, B, a nd b ), d eepl y
rudiately furrowed , ve lvety in m ost strains but with surface a ppearing loosely
l.u uu e in sorne isolat es; u su all y h eavily sp oring throu gh out , in yellow- green
shndes (S guy, pl at e X X , 299, 300), becoming dull gray in age, bu t remaining
wliite in sorne very poorly sp oring stra ins; margin irregularly dissect ed ; odo ur
l.uk in g; ex u date a bu ndan t as a m be r-colou re d drop s; a m be r-co loured pi gment
c1 iflusing into th e su r ro u nd ing agar; scle ro tia as described a boye, ab u ndan tly
produced in sorne strains , not in othe rs (colour pl at e 1, B) , commo nly obsc u re d
hy ove rly ing h yphae and co n id ia l structu res, in othe r cases dom inat ing th e
rulony ; penicilli as described aboye; reverse in pale cin n a mon shades (colou r
pla tc 1, b ) .
Colon ies on MEA spread ing b ro adly (colour pIa te 1, C a nd c), attain ing a
c1 ia met er 01' 50- 55 mm in 14 d ays a t room temperature, pl ane , zo nate, h ea vier
sporing than upon C A, th in m a r gin wi th su b merge d m yceli a exten d ing 5 mm
(JI' more beyond th e aerial growt h;
usu all y producin g abundant sclerotia,
lorrning in so rne strains a dense , con tinuous layer adjacen t to the su bstratu m,
lending th e colo ny a n orange- p in k colour interrupted only by a reas 01' h ea vy
ro nid ia l devel opment ; penicilli as described aboye but with th e stipes 01' th e
ro nid iopho re more cons p icuo us ly a nd closely ec h in u la te.
Spec ies d escription is ce n tered upon th e typ e strain CBS 38 1.48 = C M I
~(),02 7 = ATCC 10,506 = NR RL 1640 = QM 8, 002 = IJ FM 5017, isola ted fro m th e air
in the U S by K .B. R ap er , a nd fro m m any othe r strains isolat ed b y th e au thor.
T he species is ab u n dan t a nd world-wide in di stribution . It is especially
rornrnon in soil. When fir st isola te d, strains almost in va ri ably p ro duce
ab u ndan t p ink scler otia, but under con tinu ous laborat ory cul tivatio n th ey tend
to become in cr easingly co n id ial. Strains m aintained 1'0 1' long period s in

.ollcctio ns o tcn lose th c ir abilily lo produ ce thesc churac tcristic stru cturcs.
M cmbe rs of th c specics a re sul>jcct lo co nside rab le variu tin u in cu lt ure , a nd th c
sm all co lo n ics m uy co n ra in sccto rs th at a re ciihc r p rcdominnutly co n id ia l 0 1'
sclero tial. In addition to thesc vari ations , occa siona l st rn ins SllOW sclerotia less
dcliniteIy pink in coIou r , and produ ce smallcr co nid ia l st ru ctu rcs arising as
sho rt branchcs from int crl acin g ae riaI hyph ae , su ggestive 01' th e P. decumbens
seri es . But sclerotia have never be cn reportcd for an y of th e spc cies th at
comprisc th at seri es.
The sclerotia of P. thomii M aire are borne singly and are naked , 0 1' nearl y so .
The actual tenden cy is to regard as representative of P. thomii M aire all strain s
with sim ple penicilli producin g hard , brittIe , pink sclerotia .

2 . Penicillium sclerotiorum va n Beyrn a : fig . 2, coIour pIat e 2, and SEM

plat e 1 (top : left).

~ 8 0~0O
O
0 0

lw..L.1.LwJ

10 jsrn

O
00 0
0 0 0

O
00 ()

0
()0 0 O

"
,.:"1)

L--I

~~o}Jm

:'~ ['

"J.. -1

~ ..\

~J1
K.

Fig. 2. Camera lucida

drawings of Penicillium
sclerotiorum van Beyma,
CBS 287.36. A, Detailed
drawings of the penicill i. B,
Mature conidia. e, Habit
sketches. O, Sclerotium .

Penicillium sclerotiorum van Beyma - Zentbl. Bakt. Parasitenk. Inf. Hyg . Abt. 11. 96:
416-419. 1937; See also Raper and Thom, 'A Manual of the Penicillia', pp.
160-163. 1949.
Rap cr and T hom's (1949) or iginal description is included below , sin ce thi s
ndc q uate ly presents th c species in qu estion wh en recently isolat ed. The typ c
sI rain 1\TCC 10.494 has lost some of its original cu ltural charac teristics sin ce
I he tim e of its isolation in 1936 by Professor Boedijn in Buiten zorg , J av a,
particul arl y th e ca pacity of producin g sclero tia upon all substrata .
Raper a nd Thom ' s original description is a s follows:
' Colonies on Czapek's agar growing somewhat restrictedly, attaining a diameter
of 20-35 mm in 10-12 days at room temperature , variable in colour and texture
depending upon the relative abundance of sclerotia , vegetative hyphae, and
conidial structures; varying from essentially conidial presenting a velvety
appearance, through more or fess floccose from the development of abundant
vegetative mycelium, to predominantly sclerotial with limited development of
fruiting struetures or of aerial vegetative hyphae, frequently showing these different
aspects as sectors in the same colony; conidial structures Iimited or abundant, at
first arising primarily from the substratum and en masse producing areas near
artemisia green, in age often developing from aerial hyphae over the entire colony
surface; vegetative mycelium in orange-red shades; sclerotia uncoloured or nearly
so, typically borne in fairly definite c1usters surrounded by envelopes of sterile
encrust ed hyphae in bright orange-red shades near scarlet to Brazil red, usually
occurring in limited sectors but sometimes characterizing the entire colony when
young; exudate colourless or nearly so , abundantly produced in areas of heavy
sclerotial or mycelial development, lacking or Iimited 'in predominantly conidial
areas; odour slight, suggesting mushrooms; reverse in yellow to orange-red shades,
not diffusing into the surrounding agar; penicilli strictly monoverticillate, bearing
conidia in parallel chains forming loose columns up to 150-200 p.m in length by
15-20 JLm wide; conidiophores arising from the substratum or from aerial hyphae,
seldom branched, smooth-walled, up to 200-300 p.m in length by 3.0-3.5 JLm,
enlarging in terminal areas to vesicular apices 5.0-7.0 JLm in diameter; sterigmata
parallel, in crowded c1usters of 12 or more, mostly 8-10 p.m by about 2.0 p.m, with
conidium-bearing tips somewhat narrowed; conidia elliptical, about 2.5-3 .0 JLm by
2.0-2.5 p.m, smooth -walled.
Conidia on malt agar spreading, up to 55 mm in 10-12 days at room temperature,
thin, usually heavy sporing, with eonidial areas velvety and light blue-green ;
sclerotia abundantly produced, in limited c1usters projecting through the eonidial
layers and appearing as bright red islands; penieilli as deseribed above but with
conidiophore walls delieately roughened.'

Our notes follow:

C olon ies upon C A (colour plat e 2, A a nd a) sim ilar in texture a nd cult u ra l

.ha ructcristics lo th e a bo ve doscript ion , cxce pt 1'0 1' th e abs cnce 0 1' scleroti a a nd
thc produ ction ofycllow-colourc d pigm cnt d illus ing into th e surroundin g agar.
Colo n ies UpOll C YA (colour plat c 2 , B and h) attainin g a diamct er 01' 40 mm
at room tempcrature (20 - 25 OC ), co n sisting 01' a closc- tc x tu rc d basal felt,
strongly and deeply radiately furrowed, with the centre sligh tly rais ed a nd
irregularly fold ed in a cra tcrifo r m p attern ; ve getaiive m ycelium brightly
colou re d in cin nabar inous shades influencing greatly th e co lo ny a p pearance ;
hea vil y spor ing in limited sect ors in artemisia gree n shades (S guy , XXX ,
438) with co nid ia l a reas appearing velv et y ; thin m argin surro u n de d b y a fringe
01' su b merged hyphae ex ten d ing a bo u t 3 mm beyond th e ae ria l gro w th;
exudate ab u ndan t , as ye llow -o range d rops ; odou r fa in t, ' mou ldy'; ye llo w orange-co lo u re d pi gment diffusing into th e su rro u nd ing agar; co lony reverse
in deep oran ge shades (colour pl at e 2 , b) . Scl erotia n ot observe d .
C olonies upon MEA (colour pi a te 2, C and c) growing restrict edly , att a in ing
a d iamet er 01' a bou t 20 mm in 14 days a t ro om temperature , pl ane , velvety,
heavil y spo r ing as aboye ; ex u date ab u ndan t; odou r ' m o uldy'; undulating
m argin largel y su b merged, ex te nd ing abou t 5 mm b eyond th e ae r ia l gro w th;
ye llo w-co lou red p igment diffu sing into th e surrounding agar, o the rw ise, as
describ ed a b oye . Sclerotia n ot observed .
Penicillium sclerotiorum is charac te r ize d particularly by its prominent scle ro tia .
As not ed by van Beyma ( 1937), th es e a re essen tially colo urless, 01't en
somewh a t a n gu la r in p attern , ranging from 200 to 400 tm in di amet er. In
recent isolates th ey m ostl y occur in clos ely com pactc d, sm a ll clusters a nd are
co n tained within a n e nve lope 01' sterile h yphae whose walls a re studded b y a
b right orange- re d pi gmented su bs tance which was isolated and in te ns ively
studied b y C urtin a n d R eill y ( 1940) a n d desi gn at ed scle ro tio r ine, after th e
species name . They a re very h ard , 'sto ne-like ' a nd are co m po sed 01' thi ckwall ed scle re nc hy ma-like ce lls 10-1 5 tm in di amet er.
Sp ecies d escription is ce n tere d upon th e typ e strain AT CC 10,494 = C BS
287. 36 = NR R L 2,074 = C M I 40 ,569 = IFa 6,105 = QM 1,938 = IJ FM 7241 , isolated
from th e a ir b y Prof. Bo edijn , in Buitenzorg , J a va, Indone sia , in 1936 .
The species is less abundant th an Penicillium thomii M aire . It ha s been
rep orted .from soil in south Africa , from leal' litter in the Congo, and from
det eriorating military eq u ip men t in tropical and su b trop ical regions .

3 . Penicillium syriacum Baghdadi: fig. 3, colour plate 3 , and SEM plate 1,

(centre : left ).
Penicillium syriacum Baghdadi - Nov . Syst. Niz. Rast. 7: 111-112. 1968.

Colo n ies on C A (colo ur plate 3, A a nd a) grow ing rather restrict edly ,

a tta ini ng a d iamet er 01' 20-40 mm in 14 da ys al room temperature

:P

a BO 0 0

000 0

00

o o

ll..wl....J

10 pm

00 0

at.
L.--..J

100pm

Fig. 3. Camera lucida

drawings of Penicillium
syriacum Baghdad i,
CBS 418.69.
A , Detailed drawings of the
penicilli. B, Mature conidia.
e, Habit sketches . D,
Sclerotium.

('2 0 - 25 OC ), close -tex tured, to ugh , with surface appearing velve ty, dee ply
radi ately furrowed , slightly rai sed and fold ed in colony centre, at first white ,
with eve n m argin becoming dark blue-gray (Sguy, PI. X X XIII , 492) at
matu ri ty of co nid ial structu res; th in m argin wit h submerged hyp hae ext end ing
about 2 m m beyond the aerial growth ; odo ur lacking ; ex udate a bundant, in
large yellow d ro ps; bright yellow pigment di ffu sin g into the surrou nd ing agar;
colony reverse (colour plate 3, a) in ligh t brown shades . W h ile abu ndant
producti on of b row n to bl ack scle rotia was re ported in Bagh dadi ' s original
di agn osis of the spec ies, at no time , however , ha ve we obse rve d any ev ide nce of
suc h st ruc tures in th e type strain studied. This is probably du e to the long
maint en ance in artificial cultu re since the tim e of its isolation. The stipes of th e
conid iophore a rise pri marily from the substratum or from aerial hyphae (fig . 3,
C) , eithe r with walls smooth or fine ly echi n ulate (fig . 3, A), u p to 300 11m in
lcn gth by 2.0-3 .0 11m in diameter, with ap ices enla rge d in a ves icle up to
:>. 5 11m in di am eter. Con idial structures limi ted in number , not influencing the
colony appearance; pen icilli strictly sim ple , with phialides aggregated in

O
0

o
B o
o00o Oo O
o

00 0
000

0 0 00000

lwl..l.wJ.J

10

\.

\~ /

J. , } ..~"~"\..)i,

~m

pj>
ji'

, 'l i t
l .'

, J._ .

L.-.--J

100)Jm

' ." .'

ato

~
.;
~
'

'. ,

Fig. 1. Camera luc ida

drawings of Penicillium
thomii Maire, CBS 381.48.
A, Detailed drawings of the
penicilli. B, Mature conid ia.
e, Habit sketches. D,
Sclerotium .

Penicillium yezoense Hanzawa apud Sasaki et Nakane (as 'yezoensum') - J.

Agric. Chem . Soc oJpn. 18: 774. 1943.
Misapplied name
Penicillium thomii Zaleski - Bull Acad. PoI. Sci. Math. Nat . Sr. B. 492-493, PI.
56. 1927.

Colonies on Czapek's agar medium (co lour pla te 1, A a nd a) growing fa irly

rapidly in most st rains, attaining a diameter of 45 -55 mm in 14 days al room
temperature (20-25 OC ), occasionally more restricted, with a 4 mm wide,
white undulating margin, consisting of a tough basal felt wi th surface
appearing loase textured to slightIy lanate, white to pale blu e-green in colour,
spor ing weekly througho ut but more a b u n dan tIy in marginal arcas and in
localized sectors in gray-green shades (Sguy , plate XX , 298, 299) , sometimes
producing abundant, rounded to oblong, pi nk sclerotia, up to 300-350 /.I.m in
d iameter (fig. 1, D) throughout the colony area, sometimes in limited sectors

~ 1lI

compact cluste rs o f 4-8 pcr verticil, 7.0 -1 1.0 .UII by 3.0 -4.0 .o n , terrninated
in short narrowcd neck (Iig. :~, A) ; co n id ia globoso lo slightly clliptical,
3.0- 4. p.m in di amet er, deli cately echinu late to nca rly smooth wa lled when
obs erved under th e Iigh t mi cr oscope but co ns picuo us ly roughen ed wh en
observed under th e SEM (SEM plate 1, ce n tre : left); conid ial cha ins forming
loosely parallel colu m ns (fig . 3, C) .
C olonies on CYA (colour plat e 3 , B and b) spreadin g broadly , attaining a
di amet er of 40-45 mm in 14 d ays at room temperature (20- 25 OC ), sim ilar in
texture to the abo ve description ; eve n margin with a fringe of su b m erged
hyphae ex te nd ing 2-3 mm bey ond th e aerial growth , deepl y and radiately
furrowed , with ce ntre fold ed and depressed in a crate riform pattern; colon y
surface at fir st white to yellowish, with m argin becoming dark, lead gray with
age; spa rsely sporing in blui sh-violet sh ades (S guy, PI. XL , 593, 594); od ou r
lacking; ex ud a te a bu ndan tly produced in bri ght yellow drops ; bright yellow
pigm ent diffusing into th e su r ro u nd ing a ga r; colon y reverse (col ou r plat e 3, b)
in brown shades; th e stipe s of th e conid iopho re arise from the substratum and
from tr ailing ae rial hyphae as a boye; scIcro tia have not been obse rve d .
C oloni es on M EA (colou r plat e 3, C and c) spreading broadl y a ttai n ing a
diameter of 45-50 mm in 14 d ays at ro om temperature (20 -25 OC ) , plane or
with sha llow furrow s; colou rless a nd thin m argin , with a fringe of sub me rged
hyph ae ex tc nd ing a bo u t 2 mm beyond th e aerial gro wth; colony surface
gradually rais ed tow ards th e ce n tre; ce n tral and su bce ntra l a reas cove re d by an
over gr owth of whi te ae rial hyphae lendin g a sublan at e a p peara nce to th e
colony ; spo ring areas coloured in gray-green (Sguy , PI. XXIX, 430) to pale
blu e-gray shades (Sguy , PI. XXX , 449) , m ore or less zo nate, at cen tral and
subcentral a reas; odo u r lacking ; ex udate not ob served ; no colou re d pigment
diffusin g in to the surro u nd ing agar; scIerotia hav e not been obse rve d; colon y
reverse (colour plat e 3 , c) in pale yellow shades . Pen icilli as described above .
Sp ecies d escription co nce rn ing co nid ial structu res a nd colonial
characte ristics is ce nte re d upon th e typ e stra in C BS 418 .69 = BKM
F - 1,071 = lJ FM 5043 , isolated from Syri an desert soil by Ch . Baghdadi in
O ctob er 1963.
Haghd ad i' s original description of th e spec ies follow s, since thi s ade quate ly
presents th e species in question :
' Colo nies on CA atta ining a diameter of 20-25 mm in 10 days, velvety , radiately

furrowed but not deeply, irregularly folded, with margin enti re; colony surface at
first wh ite , assuming grayish colo urs, finally leading to gray shades in age ; colony
reverse in yellowish-brown shades; yellow pigment diffusing into the surrounding
agar; exudate abundant , colourless ; odour faint , suggesting m ushrooms; sclerotia
abundantly produced , variously shaped , spherical , ovoid , elliptical, brown to black
coloured , from 50 to 65 .m up to 150 .m in diameter . Conidiophores arising directly
frorn the substrat um or f rom aerial hyphae, measu ring from 30 to 300 .m in length

by 2.6-3.0 ILm in dlamet er, with wall s spinulose and aplces enlarg od in a vosicle
3.5- 6.0ILm in diam eter. Penicilli strictly simple, somet imes aggregat ed in clusters of
2-3; phlalides, bottle-shaped, 6.5-9.0 JlITI by 2.5-3.0 JlITI in clustsrs of 5-8 per
verticil; conidia globose to slightly elliptical, with walls echinulate , 3.0-3.5 11m in
diameter; conidial chains up to 65-160 11m in length .
Colonies upon MEA attaining a diameter of 25 mm in 10 days, close textured,
with surface asperulate, yellow-white in colour; exudate lacking; reverse in yellow
shades with similarly coloured pigment diffusing into the surrounding agar.
Colonies upon corn meal agar attaining a diameter of 30 mm in 10 days , buckled
and wrinkled, velvety, partially lanate, in yellow-white shades; exudate colourless ;
reverse in reddish shades. Th is fungus ca~~ot be related to any known species. It
was isolated from soil near Berza and Kabun , in the Damascus area and near Es
Suveida (Svria) in October 1963. The type culture is maintained with the Damascus
University Fungus Collection (Svrial and at the Lomonossov University Fungus
Collection, Moscow, (USSR):
(Translated from the Russian by Prof. L. Vlas).

4 . Penicillium turbatum W estling: fig. 4, colour pl at e 4, and SEM plat e 1

(centre : ri ght).

0 0 O

O 000

OC)
() B 0

...
'

. ;

00 O
O 000

1t

tIc! , ~
\

., ..

100 jsrn

Fig. 4. Camera lucida

draw ings of Penicillium
turbatum Westling , CBS
383.48. A, Detailed drawings
of the penicilli. B, Mature
conidia. C, Habit sketches.
O, Sclerotium.

Pen;c;//;um turbetum Westllng - Arklv Bot. 11: 128-130. 1911. See also Thom, 'The
Penicillia', p. 207. 1930; also Raper and Thorn, 'A Manual of the Penicillia', pp.
166-167. 1949.

C olonies on C zapek 's a gar medium (colour pIat e 4, A and a) growing

some what restrictedl y , attaining a di amet er of 30-38 mm in 14 days at room
temperature , consisting of a thin , tough , close-t extured basal felt , strongly
buckled and radiately wrinkled , zanate , velvet y in appearance , fairly heavil y
sporing in central areas in pale grayish gree n shades (Sguy, PI. XXIX, 425 ,
a nd PI. XXX , 438); undulating ma5Pin surrounded by a frin ge of submerged
hyphae exte nd in g abou t 3 mm beyohd th e ae rial growth , spo ring sparingly in
white to gray shades ; odou r faint , indefinite ; exu da te lacking or limited in
amo u nt ; reverse (colou r pIate 4, a) in cre am to light ora nge shades; pale
orange-colou red pigment diffusing into th e su rrou nd in g aga r. Sclerotia not
obse rve d upon this substratum. Stipes arising directly from the su bstratu m,
less com mo nly from cre eping hyphae , short , m ostly 40-70 /lm in len gth but
ranging from 10 to 100 /lm in length by 2.5- 3.5 /lm in diamet er, with walls
smo oth, u sually bearing strictly sim ple penicilli com pa ratively large , showing
occasionally one or two indep endent branch es (fig. 4 , A); phialides few in
number, usuall y 4-8 in th e verticil, m ostly 10-14/lm by 2 .2- 3 .0/lm,
terminating in a long tapered ne ck; conid ia elliptical, sm ooth walled or nearly
so (fig . 4, B, an d SEM plate 1, centre : ri ght), 3. 5- 5. 5 /lm by 2.5 - 4 .0 /lm in
diameter, m ostly 4.0- 4.5 /lm by 3.0-3 .5 /lm .
In ou r ex pe rience phialides ave rage a littl e longer than Raper and Thom 's
( 1949). Otherwise th e description as presented abo ye is in satisfac tory
ag ree me nt with R ap er and Thom 's description of th e spe cies (1949) .
C olonies on C YA spre ad ing m or e rapidly than upon CA (colour plat e 4, B
and b) , a tta in ing a diameter of 45-50 mm in 14 days at room temperature
(20-2 5 OC ) , deepl y and radiately furrowed and folded in th e ce ntral ar ea in a
crate riform pattern ; m argin entire; clase textured , velve ty in ap pearance,
spo ring slightly in pal e gray shade s; odour lacking or indistinct; exudate
limited to a few drops in light amber shades; reverse (colour pIa te 4, b) in
cinnamo n shade s; pale am be r pigment diffusing into th e surrounding ag ar;
sclerotia not obse rved upon this su bstra tu m. C onidial structures limited in
number, not influencin g th e colony appcarance; peni cilli as described abo ye .
Coloni es upon MEA (colour plat e 4 , C and c) spreading broadly , attaining a
diameter of 50-60 mm in 14 days at room temperature, cons iste ntly heavi er
sporing and usu ally producing abu nda nt con id ial structures ove r th e entire
colony surface , giving a powdery appeara nce to th e colony, in blue-green
shades in central areas (Sguy, P . XXVII , 400), thin and plane, slightly ra ised
in centre; thin m argin with submerged hyph ae exte nd ing largely (about
15 mm) beyond th e ae r ial growth ; producing sclerotia-like str uc tures whi ch

d ilfer mnrkedly 1'1'0111 those 0 1' Penicillium thomii M alre in th ut thcy are
.o m pnrutivcly son, crush eas ily , a nd are com poscd 0 1' pa rcnchyma-Iike cells
with walls mod crat ely thi ckcn ed. T hc sclero tia a re adj accnt to the subs tra tum
a nd parti ally obs curcd by overlay ing co nidia l str uc tu res, attaining a d iam eter
up to 150 un (fig. 4, O) , colourless to ycIlowish, com posc d 01' polyhcdral ceIls
10-1:) /lm in diameter, with waIls 1.0-2 .0/lm thi ck ; penicilli as descr ibed
ab oye.
Species description is cente red up on the typ e strain C BS 383 .48 = C M I
39,7 38 = ATC C 9 ,78 2 = NRRL 757 = I FO 7,762 = Q M 1,941 = IJ FM 5016, isolat ed by
W estling probabl y from soil, in 1911.

c,l
5. Penicillium pusillum Smith: fig. 5, colour plat e 5, and SEM plat e 1
(bottom : left ).

::=-------

,I

WWw.iJ
10 }1m

0
O

O 0
O

()
O O O
_O

0 0
O 0 .0
00 0
0
00

O
O

B O
O O

O O o

OC) O

f.f\. .
',. ,.. e

~~~l' ~

. '

/ 1 :..:;;....
. ,:.

L---l

100 }1m

"'.' .- . :

Fig. 5. Camera lucida

drawings of Penicillium
pusillum Smith, eBS 312.63.
A, Detailed drawings of the
penicilli. B, Mature conidia .
e, Habit sketches. D,
Sclerotium.

Penicillium pusillum Smith - Trans . .Brit. Mycol. Soco 22: 254-255. 1939. See also
Raper and Thom, 'A Manual of the Penicillia'. pp . 167-168. 1949

olonies on CA ;OI0111' pia le !, A and a) gl'O wing' VCI'Y resu'ictedly and

slowl y, at tuining a clia mc tc r 01' !- IO 11\111 in 11 d a ys al I'OO Ill rcmperatu rc
(20 -2 5 OC ), thin and th ough , with su r facc co lourcd in pal c pinkish shades
co nsist ing 01' tr ailin g hyphae a nd ropes (JI' hyph ue , with ce n tre usually
depressed and with th e wh ole su rface stro ng ly Iold cd and wrinkled ; colon y
rever se (co lou r plat e 5, a) in d ark purple shades bccom in g almost black with
age; purple-colourcd pigm ent diffusin g slowly int o th e surroundin g ag ar ;
od our lacking ; ex udate not observed ; sclerotia not obse rved upon thi s
substratum . Penicilli str ictly sim ple, ve ry few in number , scatte red ove r thc
colony surface; stipes 0 1' th e co nid iopho re a rising fro m tr ailing hyphae and
ropes 0 1' h yphae , mostl y sim ple bUlt with a n occasional br anch (fig. 5, A),
35 -55 IJ-m b y 1.5- 2.0 IJ-m , smooth walled , slightly enla rge d a t apex ; phialides
in number 0 1' 3- 5 per wh orl (fig. 5, A), 7-10 IJ-m b y 2. 0 -3 .0 IJ-m , terminating
in a very sho rt taper ed nec k; co nid ia smooth or nearly so (fig . 5 , H, and SEM
plat e 1 (bottom: left), ovate to subglobose, 2.5-3 .5 IJ-m in di amet cr, forming
chains in loose , di vergent co lu m ns 50 - 100 IJ-m long (fig . 5, C) .
Colo nies on C YA (co lour plat e 5, B a nd b) growing ver y restrictedly and
slow ly, but less th an upon C A , attaining a di amet er 0 1' 15 mm in 14 d ays at
room tempera ture , cons isti ng 01' a very stro ng ly folded and wri nkl ed su rface
5 m m dee p, with m argin a b rupt ; scarcely sp oring in light cream shade s; odo u r
lackin g; ex udate not obse rve d; no pi gm ent d iffus ing into th e su r ro u nd ing agar;
colony reverse (co lou r pla te 5, b) in cin namo n shades; sclerotia not obse rve d
up on this su bstratu m . Pen ieilli as described a boye.
Colo nies on MEA (co lou r plate 5 , C and c) grow ing as restrictedl y as up on
CA , atta ining a d ia me ter 01' 7- 10 m m in 14 d ays at room te m pe ratu re
(20 - 25 OC ) , ve ry stro ngly folded and wrinkled as described a boye, cream
colou re d ; odo ur lacking ; ex udate not observe d ; colony reverse (co lou r plate 5,
c) in pale cream sha des; no pigm ent diffus ing in to th e su rrou nd ing agar;
scle rot ia (fig . 5, D ) browni sh to pin kish-brown, irregul arl y globose, averaging
300 IJ-m in d ia met er , mostly co nflue nt, compo sed 0 1' m asses 0 1' irregul arl y
globose or pol yh edral cells .
M ea surements ob ta ine d in our ob servation , as recorded a boye, a re m
almost complete ag ree me nt with Sm ith's original di a gn osis 0 1' the spec ies
( 1939) .
Spec ies d esc ript ion is cen te re d up on the typ e stra in CES 312.63 = C M !
89,286 = !J F M 5046, isolat ed fro m sa mples 01' d ried blue peas by D r. e.E .
Turfitt.

6. Penicillium indicum S andhu el Sandhu : fig. 6, co lo ur pi a te 6, a n d SEM

plate 1 (bo tto m: ri ght ).
Penicillium indicum Sandhu et Sandhu - Can. J. Bot. 41: 1273-1274. 1963. See also

t\'~J-oo
~d~-~
.
oo
o 0 0 00 O
0000
O O O O

O O O B

Lu..u..Lu.u..I
10 um

;::T2

e,\.

\"

~11clrli
!

100 }Jm

Fig. 6. Camera lucida

drawings of Penicillium
indicum Sandhu et Sandhu,
CBS 115.63. A, Detailed
drawings of the penicilli. B,
Mature con idia. e, Habit
sketches. D, Sclerotium.

Kulik, 'A Compilation of Descriptions of New Penicillium Species '. Agric .

Handbook. No. 351. Agric. Res. Ser. U.S.A . Dep. Agric. p . 44, 1968.

C olon ies on C zapek's agar (co lour plate 6, A and a) gro wing fa irly rapidly ,
attaini ng a d ia me ter of 40- 50 mm in 14 days at room tem pe ratu re
(20-25 OC), plane bu t somewhat raised and flu ffy in the ce ntre, which is
cove re d by tufts of ste rile hyphae , giving a cottony appearance to th e colony;
wh ite at first , becoming grayish green in heavily sporing a reas, wi th m argin
irregul arl y di ssected from the uneven growth of the vegetative myccl iu m;
exudate abun dant , colou rless; odou r lacking or in definite ; sclerotia abu ndant ly
produced , lending a granula r appearance to the colony su rface, globose to
su bg lobose (fig . 6, D ), white to cream colou red, covering th e entire colony
su rface, soft , easily crushed, formed of thick-walled pseudop arenchyrna-like
cells, not envelope d by h yphae , m ostly 60- 150 .tm in th e long ax is. Penicilli
strictly sim ple (fig . 6, A), with the stipes of the conid iopho res a ris ing prim arily
from ae rial hyphae , short, smooth-walled, ranging fro m 17.0 to 40 .0 .tm by
1.5- 2.5 tut , with apices en la rged in a vesicle 3.5 to 6. 0 tui: in d iame ter;

phi ulides parallcl, crowded up lo 15 per vcrticil , mostly 6 .0 -9 .0 .tlll by

2.5 -3 .0 .tlll ; conid ia globoso lo subglobosc, smoo th wallcd whcn ob scrvcd
un dcr the Iight microscop e (lig. G, B) 0 1' slightly ro ug he ncd whcn obscrved
under th c SEM (SEM plat o 1, bott om : ri ght), usu all y 1.5- 2.5 .tUl ; colony
reverse (colour plat e 6, a) in pale cream rosy shades.
Colonies on C YA (colour pla te 6, B and b) spread ing broadly , atta ining a
di amet er of 50 -55 mm in 14 days a t room temper a ture , sim ila r in texture to
tho se gro wn upon CA but strongly raised up to 5 mm in th e central area ,
pulling apart and cracking th e agar underneath , with shallow radial fu rrows
and a cr ateriform dep ression at th e ce nt re; heavily sporing throughout in
arte m isia gr een colours (Sguy, Pi;. XXVII , 404), wi th su rface covered by
'"
numerous tu fts of white sterile hyphae
giving a gran ular appeara nce to th e
colony. Penicilli as described abo ye, with conid ial chains forming com pact
colu m ns. O dour lackin g 0 1' indefinite ; exudate colou rless ; ligh t yellow pi gm ent
diffusin g into th e surrou ndi ng ag ar; colony reverse (colour plat e 6, b) in pale
salmo n p ink shades with a gree n cas t sho wing throu gh in spo rulating areas;
sclerotia abu ndantly prod uce d, white to cream coloured (fig . 6, D ).
Colonies on M EA (colour plat e 6, C and e) spread ing broadl y , attain ing a
di ameter of 50- 55 mm in 14 days a t roo m temperature , heavily spo ring
throu gh out in dull gray-green shade s (S guy , PI. XXV , 373); margin with
subme rge d m ycelia exte ndi ng 5 mm beyond th e aerial growth; colony su rface
plane wi th tufts of fascicles of ae rial hyphae in ce ntral area; odo ur lacking ;
exuda te not obse rve d; colony reverse (colou r plat e 6, c) in pale pinki sh cream
shades; sclerotia abu ndan tly produced but not lendin g colour to the colony.
Pen icilli as a boye ; conid ial chains lon g, forming definitely com pact colu m ns .
Spec ies descri pti on is ce ntere d up on the typ e stra in CBS 115.63 = ATCC
18324 , isolated from a sputum specimen in N ew Delhi , India by Drs D .K. and
R .S. Sand h u , in May 1962.

7. Penicillium donkii Stolk: fig. 7, colour plate 7, and SEM plate 2 (to p:
left) .
Penicil/ium donkii Stolk - Persoonia . 7: 333-337. 1973.

Colonies on CA (colour plate 7, A and a ) spread ing fairly rapidly , a tta in in g

a diamet er of 40 -45 mm in 14 days at roo m te m pe ra ture (20 - 25 OC), deepl y
velve ty, loose textu red , appearing more 0 1' less lanat e , rai sed in cen tre by a n
ove rg ro wth of white sterile hyphae ; with shallow radial furrows; abundantly
spo ring in marginal and su bmarginal are as ; even m argin ofte n cons isting of
largely sub me rge d m ycelia exte nd ing 2-3 mm beyond th e ae rial growth ;
heav ily sporing throughout in cinnamo n shades (Sgu y , PI. XXIII , 338);
odo ur lacking ; exudate not observe d; reverse (colou r pIate 7, a) at first pinkish ,

~.

o O o

oo O O O
O

00

lwill.wJ

10 }1m

O O O

0 0 O 0

o
O o BoO

. [eil1ho~~
..

/~

?R,

Fig. 7. Camera lucida

drawings of Penicillium
donkii Stolk, CBS 188.72. A,
Detailed drawings of the
penicilli . B, Mature conidia .
C, Habit sketches. D,
Sclerotium .

Iate r sho wing dark brown shades (Sguy, PI. XLVII , 69 1); d iffus ing pigment
into the su rro u nd ing agar dark brown; soft, white, setose sclero tia- like
struc tu res usu ally produced near th e agar su rface, globose to su bg lobose (lig .
7, D), mostly 60-150 /lm in di ameter , hyaline when yo ung, but soon
bccoming brown ; pseudop aren chym at ou s walls co nsis ting of com paratively
sma ll irregu la r cells, 2-3 /lm in d iame te r, bearing n umer ous, septa te, hyaline ,
rad iating , stra ight hyphae , up to 300 /lm in len gth , completely obsc u ring th e
hrown in ne r bodies, giving a white appearance to th e struc tu re. H ea vily
spor ing in light grey -gree n shades (Sg uy, PI. XXV 375); penicilli stri ctl y
sim ple (fig . 7, A) ; stipes of th e con id iopho re s usuall y aris ing fro m th e bas al felt
a nd fro m aeri al veget at ive hyph ae, developing someti m es in marg inal a reas
lrorn subme rge d hyphae , septat e , va riable in len ght , ra ng ing from 20- 300 /lm
by 1.5- 3.0 tui: in di ameter , smooth -walled or nearl y so, with apices enla rged
in a ves icle 4.0-5.0 /lm in di ameter; phialides in cro wde d who rls up to 10-1 2
per verticil, cons isting of a cylindrical, bottle- shaped base tapering abru ptly to
a sho rt neck , 7.0- 10 /lm by 2. 0-3 .0 /lm ; conid ia su bglobose to ellipsoidal,

2.0-:U l

jllll by 1.5- 2.0 jllll , smooth when obse rved und er the light
ruicroscop e ( fi~ . 7, B) 01' dcl icat cly ro ughc nc d wh cn o1JS CI'V C<! und cr th c S EM
(SEM plat e 2 , top : lcft }; co n id ia l cha ins formin g wcll -d cfin cd , so rne timcs
sligh tly twi sted colum ns up l o 200 jlllJ lon g (lig. 7, C) .
Colo n ies on C YA (co lour plat e 7, B a nd b) sprcadi ng b road ly a nd rapidly ,
attain ing a di amet er o f 50-55 mm in 14 d ays at roo m temperature , co ns isting
of a loose-textu red felt , velvety in a ppearance, poorl y sporing throu ghout in
light pinkish shade s; strongly and deepl y radi ately furrowed , with ce n tre
depressed and irregularl y folded; thin m argin with su b merged hyph ae
exte nd ing 3- 4 m m beyond th e ae rial gro wt h; ex udate ab u ndan tly prod uced in
dark sien na shades (Sguy, PI. X.tI, 172); sim ila rly colou re d pigm ent d iffus ing
into th e su r rou nd ing agar; odo ur lacking ; reverse (colou r pl at e 7, b) in d ark
reddish b row n sha des; sclerotium-like bodi es not observed up on thi s
su bs tratu m. Penicilli as described a boye.
Colon ies up on MEA (co lou r plat e 7, C a nd c) sp read ing b ro adl y , attaining a
di amet er o f 50-60 mm in 14 d ays at ro om te m pe ra ture (20-25 OC ), cons isti ng
of a loose-textured basal felt, poorl y sporing th rou gh ou t in light gray -green
shades (Sguy, PI. XXVII , 400), with a b u ndan tly p rod uce d sclerotiu m -like,
white, setose bodi es affecting the colo n y with a gr anul ar a ppeara nce ; thin
marg in with su b me rge d h yph ae extend ing 5 0 1' more mm beyon d th e ae rial
growth; odo ur a nd ex udate lackin g; colo ny rever se (co lour pla te 7, c) in very
d ark , almost bl ack gree n shades; gree n coloured pi gm en t diffu sin g slowly into
the su rro u nding agar. Penicilli as d escribed aboye .
Species desc rip tio n is ce n tered up on th e type strain CBS 188.72 = C M !
197 ,489 = !J F M 5023, isolat ed by L. K . liver fro m ara ble Alas ka n so il.

8. Penicillium grancanariae R amirez , M artne z el Fe rrer : fig. 8 , co lo u r

pl ate 8, a n d SEM pl ate 2 (t o p: r ig h t) .
Penicillium grancanariae Ramrez, Martnez el Ferrer - Mycopathologia. 66: 77-82.
1978.

Colonies on C A (colour pl at e 8, A and a) growing fairly rapidly , atta ining a

di amet er of 40- 45 mm in 14 d ays a t ro om temperature (20-25 OC), co nsisting
of a close-tex tu re d basal felt with an irregularl y dissect ed margin , velv et y in
ap peara nce, m argin with su b me rged m ycelia ex te nd ing a bo u t 4 mm beyond
the ae rial gro wth; almost plane with sorne cen tra l fu rrows; heavily spo ring in
gray-gree n shades (Sguy, PI. XXVII , 402); colony surface covere d by an
ab u ndan t overgro wth of whi te ste r ile h yphae , influen cing th e colon y
a ppeara nce a nd obsc uring in zones th e a bu ndan tly p roduced conidial
struc tu res; ex udate lirnited in arno u n t 0 1' lackin g; od our fain t , indistinct ; pale
a mbe r-co lou red pigrnent diffusing into the su rrou nd ing agar; reverse (co lou r

10 urn

. ~f. ).\

fR.

fj

~
't1
~

Fig. 8. Camera lucida

drawings of Penicil!ium
grancanariae Hamrez.
Martnez et Ferrer ATCC
38,668. A. Detailed drawings
of the peniclli. B, Mature
conid ia. C, Habit sketches .
D. Detailed drawings of the
thick-walled sclerotial cells .

plat e 8 , a) in ligh t cream shades; scle ro tia -like struc tures have not b een
ob served upon th is su bs tratum. P enicilli strictly sim ple o r ncarly so, sho wing
scldorn a n occa sional branch (fig . 8 , A) ; stipe s of th c co ni diopho res sho rt ,
usu all y up to 100-1 50 tm in length by 3.0-4. 0 tm in di amet er , a r ising
di rectl y fro m th e substratum, with walls r anging fro m d eli cat ely to coarscly
ro ug hened but scldom smooth , with ap ices e nla rg ed in a vesicle up to 7 tm in
di amet cr; phialides formin g cro wde d clus te rs of 10-12 pcr ve rticil, m ostl y
6.0- 13 tm by 3 .0 -5 .0 iutv , co m monly producin g ch a in s of co n id ia in fairly
co rnpa ct co lu m ns up to 400 tm long (fi g . 8, C) ; co nidia clli ptica l to
su bglobose, mostl y 3.0 - 4. 0 tm by 2.6- 3 .0 tui , ve ry delicat ely rou gh ened
whe n obscrvcd under th e light mi croscope (fig . 8 , B) , but showing a
ce rebr ifo rm pattcrn whe n observed with th e S EM (SE M plate 2, top : right) .
Colo n ies on C Y A (co lour piate 8, B a nd b) sp read ing b ro adly, attai ning a
diamct er of 45- 50 mm in 14 d a ys a t room tempcrature (20 -2 5 OC ), velvety ,
stro ng ly radiatel y fur rowed a n d irregularly folded in cen tra l area, very poorly
spor in g , m ostl y in cen tral arcas in ligh t gray-gree n shades; rc mainder n onspo r ing areas re main in g white ; cv en m argin with submerged hypha e

sxtcnding about 1 mm beyond the ncriul growth; odour und exudare lacking;
dark a m be r colou rcd pigm cnt dilfusin g into thc su rro u nd ing agur: colony
reverse (colou r plat o a, b) showin g cin namo n shades. Pen icil1 i as described
aboye. Scle rot ia ha ve not been obse rvcd .
Colon ies on malt ex tract agar (co lour plat e 8, C and e) spread ing bro adl y ,
at taining a di ameter al' 65 -70 mm in 14 days a l room tem peratu re
(20- 25 OC ), consisting 01' a rather loose-textured basal feIt , za nate, plane ,
heavily spo ring throu ghou t in defin ite gree n shades (Sguy , PI. XXV II , 403);
producing few, scattered, soft sclerotia-like bo dies in ora ng e shades , with wa lls
formed 0 1' thick pseudoparenc hymatous po lyhedral cells (fig . 8, D) . Penicilli as
described aboye .
~;
Penicillium grancanariae app ears most cIosely related with species al' th e P.
thomii series, in the P. turbatum sub -series, on account 0 1' th e presen ce al' soft
sclerotia- like bodies up on sorne subst rates (MEA in the presen t cas e), a l' the
rapid growth upon most med ia and a l' the strictly simple pen icilli.
Sp ecies descr ipti on is centered upon the typ e strain IJ F M 3745 = C BS
68 7.77 = ATC C 38, 668, isola ted from the air 0 1' Las Palmas de Gran C anaria,
Canary Islands, Spain by D I' R . Ruano-T ravieso , in July 1976.

Species of dubious position

Penicillium roseolo-lilacinum Novobranova 225-234. 1974.

Nov. Syst. Nizhnikh. Rast . 11:

Penicillium roseolo-li lacinum Novobra nova is a species described in 1974 that is

bel ieved to be cIosely related to th e P. thomii series. Sinc e the author 01' the
present study was not ab le to examine the type stra in al' the spec ies, beca use all
atte m pts to obt ai n it from In tern ational Fungu s C ollection s failed , it is
presen ted here as a spe cies al' dub iou s positi on an d is not incl uded in the K ey to
Species . Novobranova ' s original descrip tion follows:
'Colo nies on CA growing rapidly, attaining a diameter of 40-60 mm in 14 davs,
plane, with centre.slightly folded , almost completely covered by a layer of pinkish
lilac sclerotia, with marginal zones in gray-green shades, heavily sporing in some
areas while very poorly in others; exudate lacking 01' limited, hyaline , colourless;
odour very strong , 'mouldy'; colon y reverse in sandy shades with centre in dark
chestnut shades. Penicilli strictly monoverticillate, forming long and tangled chains .
St ipes smooth-walled , arising as short branches from aerial hyphae , 60-160 pm by
2.2-2.5 pm up to 3.0 pm , with apices enlarged in a ves icle up to 4.0 -4.5 Jlm in
diameter ; phialides in compact c1usters of 5-8 per verticil, acuminate, 6.0-8.0 Jlm
by 1.8-2.0 Jlm ; conidia subglobose, 2.0-2.2 pm in diameter. It belongs to the
Monoverticillata section . No related species are known .
Type: isolated form Pirus malus L. skin , 28-VII-1967 by T.1. Novobranova.

The type culture N. 194 Is depositad at the Department of Cryptogamy, Univerltv M.V. Lomonossov, Moscow, USSR, and at tha Institute of Pomology and
Ampelography, Laboratory of Microbiology of Kazachstan, Alma-Ata region.'
(Translated from the Russian by Prof. L. Vilasl,

niclllium es-suveidense Baghdadi - Nov. Syst. Niz. Rast. Lehningrad. 7: 97-114

1968.

Penicillium es-suoeidense Baghdadi is another species of dubious position since

"K'a in the author was not abl e to exa m ine the type strain of the species for the
sume reasons as aboye . It was describe"H in 1968 and is believed to be closely
rcla ted to th e P. thomii series . It is not included in th e Key to Species.
Baghd adi ' s original description follows :
'Colonies on CA attaining a diameter of 35-40 mm in 10 days, with colony sur-

Iace wrinkled, zonate, velvety, margin at first white, then becoming dirty green, with
cont re grayish; exudate colourless; colony reverse brown. Sclerotia are abundantly
produced in colony centre, variable in size and shape, giving gray colours to the
colony surface , elliptical, ranging from 300 to 430 JUTl by 150JUTl, or globose, 130JUTl
in diameter, brown coloured . The conidiophores arising directly from the substrat um , smooth walled, 500 JUTl by 3.0-3.5 JUTl, usually terminating in simple
penicilli, although an occasional branch may be encountered, measuring 14-18JUTl
by 3.5 JUTl. Phialides in number of 4-6 per verticil, mostly 7.0-8.0 JUTl by 3.5 JUTl.
Conidial chains forming divergent columns. Conidia smooth walled, at first ellip tical, 2.5 JUTl by 3.0 JUTl, becoming later globose, 3.0 JUTl in diameter.
Colonies upon MEA attaining a diameter of 20 mm in 10 days, close textured,
velvety, somewhat lanate in zones, wrinkled, in pale cinnamon shades with white
spots; exudate lacking; colony reverse yellowish.
Colonies on corn mear agar attaining a diameter of 30 mm, strongly folded,
velvety, in cinnamon shades; exudate hyaline, colourless, colony reverse in
cinnamon shades. No closely related species are known.
The type species has been isolated from Syrian desert soil near Es-Suveida. Other
isolates have been recovered near Irna, Damascus region, Syria, from meadow soil,
forest soil, from gray chestnut, from Quercus forest soil, at 5 cm of depth, in
August 1964.
The type culture N 619 has been deposited with the fungus culture collection of
the M.V. Lomonossov University, Moscow, USSR, and with the fungus culture
collection of the University of Syria at Damascus.'
(Translated by Prof. L. Vilas from the Russianl.

Penicillium frequentans series

OUTSTA ND I NG FE ATU IU ;S

Colonies grow ing ra p idly, usu ally sprcad ing broad ly, with a velve ty
appearance or with colony su rface slig h tly lan at c.
The stipes of the conid iop hores, scldom branch ed , smooth wall ed to
defini tely rou gh en ed , a rising di rectly from the basal felt or from trailing
hyphae .
Peni cilli str ictly sim ple, com paratively large , with phiali des crowded in
compact who rls, a nd with con id ial chains forming u su all y loose or com pact
~i;
columns .
Conid ia globose to su bglobose , seldo m ellipt ica l, ranging fro m 2 .5 to 5.0 utt:
or m ore in di ameter, smooth wa lled to coa rsely ro ug hened, dcp end in g u pon
the species.

Key to th e species
1. Colonies velvety or nearly so, with stipes of the conidiophores arising
mostly from the substratum . Colonies usually spreading broadly on most
media. Conidia globose to subglobose.
P. frequentans series
a. Colon ies upon CA strictly velvety, close textured .
1'. Colony reverse usually in orange -brown to redd ish-purple shades.
aa. Conidia mostly 2.5-3.5 tm, with walls thin, smooth or finely
roughened.
P. frequentans Westling
bb. Conidia mostly 4.0-5.0 tm, with walls heavy, dark green, and
coarsely roughened.
P. purpurescens (Soppl Raper et Thom
2'. Colony reverse colourless or nearly so.
aa. Colonies developing conspicuous stolon-Iike hyphae at margino
P. odoratum Christensen et Backus
bb. Colonies not developing stolon-like hyphae at margin
P. abeanum Smith
3'. Reverse usually in bright orange-red to orange shades.
aa. Colonies upon CA growing rapidly and spread ing widely.
P. luteo-aurantium Smith
bb. Colonies upon CA growing rather restrictedly
P. multicolor G-M et P. (see P. implicatum series)
4'. Colony reverse in bright yellow shades .
aa. Colonies upon CA in pale cream shades, reverse in b right
sulphu r-yellow colour.
P. cremeo-griseum Chalabuda

Page

. . . . 50

. . . . 51

. . . . 54

. . .. 67
. . . . 65

. . .. 63
. . . . 84

. . . . 59

bb. Colonias upon CA in blue-gray to mouse-gray ahadss. Reversa

yellow.
P. griseo-azureum Moreau et Moreau ex Harnrez . . .. 60
b. Colonias upon CA with surface loosa textured, with stipes arising
from the substratum and from aerial hyphae; reverse colourless,
pinkish, or in purplish shades.
P. spinulosum Thom . . . . 56
. Colonies upon CA growing very poorly due to a vitamin deficiency, but
spreading broadly upon CYA and MEA. Conidia elliptical.
P. tarraconense Ramrez et Martnez . . . . 69
(o'

Briely characte rized , sorne rnernbers of this series are a rnong the rnost
widd y di stributed a nd rnost abundant of all Penicillia . Sorne of th ern ,
cspcc ially Penicillium frequentans W estling and P. sp inulosum Thorn , a re co rnrno n
in soil a nd upon organic rnat erials under going slow decornpositi on. R aper a nd
T horn ( 1949) included in this series three well-defined sp ecies aggregates
ccntcred arou n d P. frequentans W estling, P. purpurescens (Sopp) Raper el Thorn,
nnd P. spinulosum Thorn , characte rized by con side rable varia tion within each
spccies aggregate, with , not infrequently , strain s bridgin g th e lin es of
sc pa ration between th ern . Since 1949 , how ever, six Penicillium species have
hccn described as new to science which see rn to fit quite well within th e series;
n.u nely P. cremeo-gnseum C hala b uda , P. griseo-azu reum M oreau el M oreau ex
R um rez , P. luteo-aurantium Srnith , P. odoratum Christensen el Backus, and P.
tarraconense Rarnrez el M artnez.

!). Penicillium frequentans W estling : fig. 9 , colour plate 9 , and SEM

plate 2 (ce ntre : left).

Penicillium frequentans Westling - Ark. Bot. 11: 58, 133-134 . 1911. See also
Biourge's Monographie, La Cellule. 33: 292-293 . 1923; also Thorn, 'The Penicillia',
pp. 216-217. 1930; and Raper and Thom, 'A Manual of the Penicillia', pp.

172-177. 1949.
Synonymies
Citromyces albicans Sopp - Monographie der Pilzgruppe Penicillium, pp.
128-129. 1912.
Penicillium aurantio-brunneum Dierckx - Ann. Soco Sci. Bruxelles. 25: 86. 1901.
Penicillium candido-fulvum Dierckx - Biourge's Monographie, La Cellule. 33:
275-277. 1923.
Penicillium columnare Thom - The Penicillia, pp. 214-215 . 1930.
Penicillium f1avi-dorsum Biourge - Monographie, La Cellule. 33: 290-291. 1923.
Penicillium fluitans Tiegs - Deut. Bot. Gess. Berlin. 37: 499-501. 1919.

10~m

~(I

-----==---< ,

'OoPm

(9 ~M~ (~fi

'i~) t;!j

~\~ .- Jt~~;~

'"'ihO;- ,,:"~.

f j

R.

Fig. 9. Camera lucida

drawings of Penicillium
frequentans Westling, CBS
787.70. A, Detailed drawings
of the penicilli . B, Mature
canidia . C, Habit sketches .

Penicillium glabrum (Wehmer) Westling - Ark. Bot. 11: 131-132.1911.

Penicillium oledzkii Zaleski - Bull. Acad . Polonaise Sci. Math . Nat. Sr. B. pp.

499-501. 1927.
Penicillium pfefferianum (Wehmer) Pollaci - Atti 1st. Bot. Univ. Pavia. Ser. 11. 16:

121 -136. 1916.

Penicillium sinicum Shih - Trans. Sapporo Nat. Hist. Soco 14: 286-287. 1936.

Sin ce th e typ e strain of P. freq uentans W estling C BS 105.11 , maintain ed in

a rt ificial cultu re for m an y years has lost its original cu ltu ra l a ppeara nce, th e
C BS strain 787.70 was used to prep are colou r plat e 9. The cross colony
a ppe arance description is tak en from Raper and Thom (194 9) , since thi s is
ade q uate (R idgway 's Colour C od e has been repl aced here by Sguy's C od e
U niversel des Couleurs (19 36 :
'Colonies upon CA (colour plate 9, A and a) spreading rapidly and broadly,
attaining a diameter of 50-60 mm in 14 days at room temperature (20-25 OC),
broadly zonate. radiately wrinkled in most strains, with central area commonly
sulcate, thinning at margin, consisting of a closelv woven felt of coarse hyphae

IllItar obovo or below the surtace from whlch abundant crowded conidiophores
11110, volvoty, but commonly showing limited trailing hyphae, heavily sporing,
IlflU[llIy colandine to artemisia green (Sguy, PI. XXVI, 388, 389), becoming storm
Iff1 Y In oge (Sguy, PI. XXVII, 402), in some strains darker, approximating
morlcun to dark American green (Sguy, PI. XXIX, 427); exudate Iimited, c1ear to
lIuhl nrn ber: odour faint, 'mouldy'; reverse mostly in shades of yellow-orange near
1I1 ~"ol to mars brown (Sguy, PI. XIV, 1961, but in occasional strains appearing Iight
purpluh brown' (colour plate 9, a).

IllItes ollow :
I'l tipes 01' thc con idio pho res short , up to 200 tut: in length by 3.0 - 3.5 .tm in
d lrllllc!el' (lig . 9 , A) , with walls sm ooth or finel y roughen ed , enla rged at th e
'PI' X in a vesicle up to 5.0 .tm in diamet er; con id ial structures abundantly
1'1'1It1l1ced in new isolates, forming a cro wde d, fairl y contin uo u s layer over th e
whulc colony su rface ; penicilli almos t entire ly simple but showing an
11t' I'ltsillnal bran ch ; phialid es in crowde d clu sters of 10-12 or m or e in th e
\'l1 l't ir-i l, mostl y 8 - 12 .tm by 3.0 -3.5 .tm; usually producin g chains of con id ia
I1 I'lIi rly well-define d colu m ns up to 150 .tm or more in length (lig. 9, C) ,
flnll' limcs splitting in age , in othe r strains showing conid ial chains only loosely
pnrullcl; conid ia globose to su bglobo se, with wal1s com paratively thin , smooth
111' Iiu cly rou gh en ed (fig. 9 , B) and (SEM plate 2, centre : left) mostly 3.0-3 .5
1111 1 in di amet er.
( :lIll1 l1 ies upon C YA (colou r plate 9, B and b) spreading rapidly a nd
bl'l,atlly, a ttaini ng a di amet er of 50- 60 mm in 14 days at room temperature
~ () '2 ~ OC ), azonate, velve ty, radiat ely furrowed, close tex tu re d as aboye;
hl' ltvily spo r ing in va rious shade s of gree n, sometimes ob scured by ove rruw ihs ofw hite ae rial h yphae inf1uen cing th e colony appearance; thin m argin
with subme rge d hyphae exte ndi ng about 5 mm beyond th e aerial growth;
nxud.uc abu ndantly produced in sorne strains, limited or lackin g in othe rs, in
1111 1)(:1' shade s; odo ur pronounced, ' mouldy' ; dark amber-coloured pigm ent
d llTl lsing int o th e su rro u nd ing a gar in sorne strains, ab sent in othe rs; colon y
I'I: VCI'Se (colou r plat e 9, b) in pale orange , reddish brown to light cin namo n
luul cs, Penicilli as described aboye; conid ial cha ins in well-d efined colu m ns ,
uxuull y lon ger th an upon CA, up to 300 .tm long.
(:lI lonies upon MEA (colour plate 9, C and e) spreading rapidly a nd
1ll'lIadl y, attain ing a di amet er of 55-65 mm in 14 days at room temperature
tl () _ '25 OC ) , velvety, plane, loos e textured, zonate a t m argin, with submerged
iuycclia exte nd in g largely (about 5 mm) beyond the ae rial gr owth , loose
re..cturcd ; heavily sporing throughout in dark , yellow -gr een shades (Sgu y ,
1'1. XXI , 301), with centre showing artemisia gr een colou rs (Sguy, PI. XXI ,
'.110) and a n ove rg rowth of f1occose aerial hyphae ; exudate lacking in sorne
tra ius, prese nt in othe rs ; odou r pronounced , ' mo uldy' ; colony reverse (colou r
plruc 9, c) in light reddish cinna mo n shad es, with centre dark brown . P eni cilli
( ) I I I'

as describcd aboy e; conid ial chuins in well-dcfinud COIIl IIl Il S , II p lo 500 un

lo ng , o ltc n brcak iug away w hc n cu lt u re di sh is lapped .
Spccics dcscription co nce r ning fine dcta ils 01' lIlo rphology is cc ntered u pon
the type strain G13S 105.1 1 = IJFM 5026, isolatcd Irom so il by T . R . Westling.
Speeies deseript ion coneerning cultu ra l appcarance is ccn tere d u pon st rain ces
787.70 = CM I 96,659 = IJFM 7067, isolat ed from I nd ian soil, ano u pa n many
additional strains similar in details of morphology . Colour plat es 230, 231 and
232 present three representative strains of P. frequentans, isolated from an acid
brown soil in th e provi nee of Navarra, Spain, 1976, by DI' 1. Moriyn .
This species occurs in soil and upon decaying veg etable matter, representing
on e of the most abundant and widely distributed of all th e Penicillia .
Individual strains wit hin the species show appreciable variations in gross
colony appearance and in the finer morphological detail s, w ith gradat ions
reg ula rly encountered, which seem to represent d ifferent aspects of an
abundant and va riable spe cies.

10. Penicillium purpurescens" (Sop p) R aper el T hom : fig. 10, colour plat e
10, and SEM pIate 2 (centre: right).

:@~:c~
0
e-;

; B00
@
O

lulllwJ.J

tf'>.

10 )Jm

'(j)

'~~'.
'_. -l-c{,l1
L--...J

100 )Jm

Fig . 10. Camera lucida

drawings of Penicillium
purpurescens (Sopp) Raper
et Thorn , CBS 223.28. A ,
Detailed drawings of the
penicilli. B, Mature con idia.
C, Habit sketches .

*The original spell ing of P. purpurrescens has been treated as a typographical error and correc ted
here to purpurescens.

'enlcllllum p rpurescens (Sopp) Raper e t Thom - 'A Manual of the Penlclllla'. pp.
177-180. 1949.
;ynonymies

Citromyces purpurescens Sopp - Monographie der Pilzgruppe Penicillium. pp .

117-119. 1912. Also Thom, 'The Penicillia', p. 178. 1930.

Citromyces virido-albus Sopp - Monographie der Pilzgruppe Penicillium. pp. 131132. 1912.

Penicillium baiiolum Biourge - Monographie, La Cellule. 33: 305-306. 1923.

Colonies on C A (colour plate 10, A and a) gro wing rather restrictedl y ,
attainin g a di amet er of 35- 40 mm in 14 days a t roo m te m perature, cons isting
of a rather thick, tou gh, close-t extured basal feIt , conspicuously zonate in
sorn e strains, almost azona te in others , radiately wrinkled , som etim es deepl y
bu ckled at ce ntre, appe aring ve lvety or nearly so in m argin al a reas; heavily
sporing th rough out in sorne strains, only in marginal arc as in othe rs ; conidial
areas in dark blue shades (Sguy , PI. XXX , 447) in recent isolat es, in very
p ale blue-gr een colours in olde r isolat es; exudate abu ndantly produced in sorne
strains or limited in amo u nt in othe rs, in pale amber to reddish tints; odo ur
fa int , indefinite ; colony reverse (colour plat e 10, a) sho wing pale sa lmo n pink ,
ora nge -red to purple-red (Sgu y , PI. V II , 103) or brown (Sguy , PI. VIII , 112)
shades; ligh t am ber pigm ent di ffu sin g int o th e surrounding agar; peni cilli
strictly sim ple, showing a n occasional branch ; th e stipes of th e con id iophores
u su ally arisin g di rectly fro m th e su b stratu m in a close sta nd, m ostly
100- 500 tut: long by 3.0-3,5 p.m in diameter , with walls com paratively thin ,
smo oth or finel y rou gh en ed with apices enla rge d in a ves icle 4.0 -6 .0 p.m in
d iameter (fig. 10, A) ; phiali des com monly in cro wde d who rls of 8- 12 in th e
ve rticil, 7- 12 p.m by 2.5-4 .0 uui ; conid ia a t fir st el1 iptical, becoming
subglob ose to globose, consp icuou sly rou gh ened , with a cerebrifor m pattern
whe n observed under th e SEM (fig . 10, B and SE M plat e 2 (ce ntre: ri ghtj),
m ostly 3.5 -4 .5 p.m in d iamet er , sorne larger up to 5.5 tuu ; con id ial chains in
loose colu m ns up to 150- 200 p.m long (fig. 10, C ).
Colonies upon C YA spread ing mor e rapidly th an upon C A , at taining a
diameter of 45- 50 mm in 14 days at roo m temperature (20-25 OC) (colour
pIate 10, B a nd b) , appearing velvety, radi ately furrowed , slightly rai sed with
centre depressed in a cra te riform pa tte rn ; eve n margin with subme rge d
my celia exten d in g abou t 2 mm beyond th e ae r ial gro wt h; slightly sporing in
pale brownish gray shade s (Sguy, PI. XLVII , 704), with non sporing arcas in
dirty white shade s; exudate limited , in minute drop s; odo u r lacking ; colony
reverse (colour pIate 10, b) dark b rown , almos t black ; simi la rly coloured
pigment diffusing into th e surro u ndi ng agar. Penicil1i as descr ibed aboye , with
con idial chains forming well-defin ed colum ns up to 100- 200 p.m lon g .

Co lonies upon M EA (colour piate 10, C a nd e) spreading b roadly, attaining

a diam eter 01' 55 mm in 14 day s al room temperature (20 -25 OC), plane ,
slightly convex, deeply vclvety to slightly lanate, loose texturcd and covered
with an overgrowth of white sterile hyphae giving a lanate appcarance to the
colony surface ; heavily sporing in dark olive green shades (Sgu y, PI. XXIX ,
425, 435); amber pigment diffusing into the surrounding agar ; exudate and
odour lacking; colony reverse (colour plate 10, e) in pale orange-red to reddish
purple shades; penicilli as described aboye but with conidial chain s in welldefin ed columns up to 500 Lm long, breaking away in sorne strains in cru sts
when the cultu re is tapped.
Spe cies description is cen tered upon strain CBS 223.28 = IJFM 5032, isolat ed
from soil by K . Zaleski in 1928, and upon many additional strains simil ar in
cultural appearance and in details of morphology .
The species occurs in soil and in soil-contaminated material; P. purpurescens
appears to be less abundant , but, like oth er member s ofthe Pi fr equentans seri es,
is worldwide in distribution.

11. Penicillium spinulosum Thom : fig . 11, coIou r pIate 11, a nd SEM
pIate 11 , and SEM pIate 2 (bottom: Ieft).

.f
.~ ~

:;
j

,o

,:.:

':
..:
'

: :~

'o
"o,

r.0 ,

0 0

r0

'o

'.

l"':
o,
0

o'

!;~~;o't~
tJ!" @
IV

(J'
[\1
e

L--..J

100 pm

(j(,

Fig. 11. Camera lucida

drawings of Penicillium
spinulosum Thorn, CSS
374.48. A, Detailed drawings
of the penicilli . S, Mature
conidia. e, Habit sketches.

Pimlcllllum spinulosum Thom - Bul!. Bur. An lm . lnd ., U.S . Dep. Agr e. 118: 76.
1D10. See also Thom, 'The Penicillia', pp. 183-184. 1930; also Rapar and Thom,
fA Manual of the Penicillia', pp. 180-185. 1949.

yn on y m ies
nici/lium [Citromycesl brunneo-viride v. Szilviny i - Zentbl. Bakt. Parasitenk.
Inr. Hyg . 11 Abt. 103: 145. 1941.
nicillium (Citromyces) bruntzii Sartory - C. R. Soco Biol. Paris. 76 : 605-606.
1914.
nici/lium (Citromyces) citricus Maz et Perrier - Ann. Inst. Pasteur Paris, 18:

58-559. 1904.
nicillium f1avo-cinnereum Biourge - Monographie, La Celiule. 33: 293-295.

1923.
'onici/lium (Citromvcesl lacticus Maz et Perrier - Ann. Inst. Pasteur Paris, 18:

58-559. 1904.
Ponici/lium mediocre Stapp et Bortels - Z. Bakt. Parastenk. Inf . Hyg. 11 Abt. 93:

O. 1935.
Penici/lium mucosum Stapp et Bortels - Z. Bakt. Parasitenk. Inf. Hyg . 11 Abt. 93:
1. 1935.
Penicillium pfefferianum Wehmer - Beitr. Kenntn. einheim. Pilze, 1: 22-24. 1893.
Penici/lium p fefferianum (Wehmer) Westling - Ark . Bot. 11: 132-133. 1911; also
Biourge's Monographie, La Cellule. 33: 303-305 . 1923.
Penici/lium (Citromycesl oxalicum Maz et Perrier - Ann. Inst. Pasteur Paris, 18:

558-559. 1904.
Penicillium roseo-maculatum Biourge - Monographie, La Celiule. 33 : 301-303.

1923.
Penici/lium viridi-dorsum Biourge - Monographie, La Celiule. 33: 306-307. 1923.
Penici/lium tannophagum Stapp et Bortels - Z. Bakt . Parasitenk. Inf. Hyg. Abt.
11. 93: 52. 1935.
Penici/lium tannophilum Stapp et Bortels - Z. Bakt . Parasitenk. Inf. Hyg. Abt. 11.

93: 52. 1935.

Penici/lium (Citromyces) tartricus Maz et Perrier - Ann. Inst . Pasteur Paris, 18:

558-559. 1904.

Colonies u pon CA (colour plate 11, A and a) growing rather rapidly ,

au a ining a dia me ter of 55 mm in 14 days at room temperature (20-25 OC ) ,
consisting of a loose-textu red fe1t of partialIy submerged , but predominantly
ac rial hyp hae, more or less radiately furrowed, rarely zanate , u p to 1- 2 mm
dcep in central areas, heavily spo ring in sorne stra in s, slightly in others
(lending towards steri lity under long artificia l cu1tivation); spo ring areas in
dull green shades ru n ning fro m sage gre en (Sguy, PI. XXVII, 404) to deep
olive gree n (Sguy , PI. XXVII , 402); th in even margin with a fring e of

submcrgcd hyph ae cxtcnd ing 5 0 1' more mm be yond the acr ial growrh; odo ur
very Iain t 0 1' nbse nt; exudare lackin g 01' limit cd in amou nl, crystal clcar ; colony
reverse (colour plato 11, a ) almost colou rlcss lo palc gra y, occas ionally show in g
light p inkish to crearn shades. Pcnicill i usu ally strictly m on overticillat c but
with a n occa sion al branch ; stipes 01' th e con idio pho res a rising m ainly from th e
su bs tratu m 0 1' from ae rial hyphae, va ry ing from ver y long wh en arising
directl y from th e su bstratu m, 80- 100 .tm up to 300 .tm in len gth , to qu ite
short, 25 -50 .tm wh en arising as b ranch es fro m ae rial vege ta tive hyphae ,
mostly 2. 5-3 .0 .tm in di amet er, with walls smooth in sorne strains to definite1y
rou gh en ed in others (fig. 11, A), with ap ices enla rged in a ves icle up to 4.0 .tm
01' m ore in diameter ; phialides few in number, abo ut 6 - 10 in th e ve rticil,
m ostly 6-9 .tm by 2.5-3 .5 .tm; conid ia mostly globose to su bg lobose , usu all y
3.0 -3 .5 .tm in di ameter , sorne larger , up to 5.0 usu, cons picuo us ly rou gh en ed ,
spin ulose, sho wing windin g bars in so rne strains (fig . 11, B, and SE M plat e 2
(bo tto m: left ) ; conid ial chains forming loose colu m ns 100-1 50 .tm lon g (fig.
11, C) .
Colonies upon C YA sp read ing broadl y , a tta ining a di amet er 01' 60 -65 mm
in 14 days at room temperature (20- 25 OC ) (colour plate 11, B and b) , deepl y
ra d iately furrowed , with m yce1ial felt some what heavier an d more lanate than
upon CA , with centre depressed and irregul arl y fold ed; thin m argin with a
fringe 01' submerged hyphae extend ing abou t 3 mm beyond the aeri al gr owth ;
sporing well in m arginal areas in dark gr een , almost black shades, with
nonsporing areas remaining white to light tan ; exudate and odo ur lacking ; no
pi gm ent di ffusin g into the surrou nd ing agar. Peni cilli as described aboye .
Colonies on M EA (colour pIate 11, C and c) gro wing rapidly attaini ng a
diameter 01' 50- 60 mm in 14 days at room temper ature (20 -25 OC), velve ty,
plan e , thin m argin with a frin ge 01' su b me rge d hyph ae extend ing 4- 5 mm
beyond th e ae rial gro wth, usu ally showing a loose-t extured , white lanat e felt
abo u t 2-3 mm deep ; heavily spo ring throu gh out in shades 01' very dark gree n
(Sguy, PI. XXIX , 431); odo ur and exu da te lacki ng ; no colou re d pigment
diffusin g in to th e su rro u ndi ng agar. Peni cilli as described aboye but with th e
stipes 01' th e conid iopho res cons picuo us ly ech in ulate and with conid ial chains
showing a greate r tenden cy to form 100se1y paralle1 colu m ns.
Sp ecies descriptio n concern ing cult u ra l characte ris tics are based upon the
typ e strain N RRL 1750 = C BS 374.48 = C M I 24,3 16 = ATCC 10,498 = QM
7,654 = IJFM 7096 , and upon observations m ade on a large number 01' stra ins
havin g essentially the same cu ltural and m or phological characte ristics. Strain
CB S 269. 29 = IJFM 5035, isolat ed by Biou rge , no longer represents th e species
ade q uate1y with respect to cult ural ap pearance. It is now esse ntially lanate and
almost sterile but still produces penicilli and con id ia characteristic 01' P.
spi nulosum.
Penicilliu m spinulosum T ho m is world wide in di stribution , espec ially com m on

so il. Apprcciabl e variuti ons may be cncou ntcrcd within th e spccics in

Indivi d ual stra ins with resp cct lo cu ltu ral c ha rac tc ris tics , particul arly with
l'C'fi:l'c ncc lo d cpth , colony texture, and arn ount 01" sporula tion . Thc stip es of
Ihe conid ioph orcs also differ in the dc grce of roughness , dep ending upon th e
truin a nd the substraturn . A few strain s produce more 0 1' lcss elliptical co n id ia
Il ru ntrast lo the usual globose 0 1' su bg lobose typi cal forms.
I 11

I:.! . Perucilliurnn cremeo-griseum ChaIabuda: fig . 12, colou r pIate 12, and
S EM pIat e 2 (b attam : ri ght) .

"

.~'.'

B
v.:

lwJJ.w.lJ
10 pm
~

.
I'."""""",,-'t,,

-"
~

"

', '

.-.

So~\..

';ii

R.,
;Ji

~:";" -'.~ J:'

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jrm

Fig. 12. Camera lucida

drawings of Penicillium
cremeo-griseum Chalabuda,
CBS 223.66. A, Detailed
drawings of the penicilli. B,
Mature conidia. C, Habit
sketches.

Penicillium cremeo-griseum Chalabuda - Nov. Syst. Crypt . Inst. Bot . Komarovii.

Acad. Sci. USSR 6: 168. 1950. See also M.M. Kulik, 'A Compilation of Descriptions of New Penicillium Species'. Agric. Res. Ser. U.S. Dep. Agric. p. 32.1968.

Colon ies upon CA (colour plate 12, A and a) spre ad ing fairly rapidly ,
a llain in g a di amet er of 50 mm in 14 da ys at room temper ature (20- 25 OC ),
co ns istin g of a very tou gh , cla se textured basal felt, vel vet y in a p peara nce ,

rad iatcly su1cate, dcpresscd in centre ; eve nly and thin murgin with a frin gc 01'
sub me rge d hyphae extcnding abo ut 1- 2 m m beyond the acrial gro wth, a t firs t
white , th en bccom in g sulp hur ycllow to pale crca m in age; exudate ora nge
colou red ; odo ur faint , 'mouldy'; colony surface cove red by a network of
tra ilin g vege ta tive hyphae and rop es of hyphae; pale a m ber pigm en t diffusing
into the su rrounding agar; eolony reverse (eolour piate 12, a) in yellow, eream,
to reddish shades; pe nie illi strietly simple, seldo m with an oeeas ional brane h
(fig . 12, A) ; the stipes of th e eonidio pho res arisi ng from su bmerged , ereeping
01' funieulose hyphae, u su all y short, 20 -40 tut: by 1.0- 2.5 tui, sep tate, with
wa lls smooth 01' del ieately rou ghen ed at upper parts, wit h apiees en la rged in a
vesicle 2.5 -5.0 tm in diameter; phialides fewin nu mber , 2-6 per verticil,
mos tly 6.0- 11.0 tm by 3.0 tm; eonid ia globose to subglobose, eomparatively
small, 2.0 -3 .0 tm in dia me ter, delieately rou ghen ed (fig. 12, B, and SEM
plate 2 (bo ttom : rightj) ; eonidial ehains forming loose columns up to 300 tm
lon g (fig. 12, C) .
Colon ies upon CYA (eo lour plate 12, B an d b) almost d up lieating in cultural
eharaeterist ies those grown on CA , attaini ng a diameter of 50-55 m m in 14
days at room temperature (20-25 OC), deeply velvety to somewhat lanate ,
white; growing margin thin, with a fringe of submerged hyphae ext end ing
abo u t 1- 2 m m beyond the ae ria l growth ; abundantly sporing in whi te to pale
gray shades; odour faint, indefi nite; exudate in yellow droplets; eolony reverse
(eolour plate 12, b) in pale yellowis h eream shades; light amber-colou red
pigm ent diffus ing into the surrounding agar; pen ieilli as deseribed aboye .
Colon ies on malt-extraet agar (colou r plate 12, C and e) spreadi ng broadly,
attaining a dia meter 01' 50- 60 mm in 14 days at roo m te mperature
(20-25 OC), plane , deepl y velve ty to su b floccose, ow ing to a thi n overgrowth
01' wh ite aerial hyph ae lend in g th e eolony a lan at e appearanee; thi n margin
surrounded by a fringe of submerged hyphae exte nd ing 2-3 mm beyond the
aerial growth, eonsistently heavier spo ri ng throu gh ou t in mou se-gr ay shades
(Sguy, PI. XL, 598) ; odo ur pleasant, aromatie; exudate laekin g ; yellowis h
green-eolou red pigm ent diffus ing into th e surrou nding agar; eolony reverse
(eolour pla te 12, e) in yellowis h green shades . Pen ieilli as des eribed above.
Spee ies dese ription is eente red upon the type strain CBS 223 .66 = ATCC
18,320 = BKM F - 1,034 = IJFM 5011 , isolated from Russia n fores t soil in 1949
by T. V . C ha labuda, in th e env iro ns 01' Ki ev.

13 . Penicillium griseo-azureum Moreau et Moreau ex Ramrez: fig. 13,

colour pla te 13, a nd SEM plate 3, (to p: left).

Penicillium griseo-azureum Moreau et Moreau - Rev. Mycol. Pars. 6: 59-61. 1941.
See also M.M. Kulik, 'A Compilation of Descriptions of New Penicillium Species.'
Agric. Handbook No. 351. Agric. Res. Ser. U.S . Dep. Agric. p. 42. 1968.

hl

-fJL! =!
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0 O 10).lm \. \'./.,
00 8 @ .. 0
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...'Y

00

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00 ' 0
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Oo OOoOO O

00 0
v, "
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~
, <::.~
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"

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Fig. 13. Camera lucida

drawings of Penicillium
griseo-azureum Moreau et
Moreau ex Ramrez, CBS
162.42. A, Detailed drawings
of the penicilli. B, Mature
conidia. C, Habit sketches.

Penicillium griseo -azu reum was isolated from sand dunes in France by Drs F .
and V. Moreau (1941) , who prop osed th e name P. griseo-azureum and des cribed
I he spec ies without providing a L atin di agnosis . To validat e th e species a Latin
diagn osis of it is give n below ; the names of th e origin al au thors are maintained
under Recommendation 46 C of the International Code of Botanical Nomendature (Stafleu (Ed.), 1972).
Latin diagnosis of Penicillium griseo-azureum Moreau el Moreau ex Ramrez:
Coloniae in agaro Czapekii satis celeriter crescentes, post duas hebdomadas ad
20-25C, 35 mm in diametro assecutae sunt, velutinae, denso texto basali coacto
mycelio, leviter sporulantes, primum albae, deinde griseolo-coeruleae, inaequaliter
prominentes, plicatae, radiatae, cum aequabili margine, parum elevatae ad
centrum; exsudatio adest, exiguus; odor abest; reverso cremeo-albido; neque
ascomata neque sclerotia visa sunt. Stipitibus erectis, brevis, glabris, hyalinis,
plurimum e mycelio basali vel e hyphis reptabundis, vel etiam ex substrato
exoriuntur, plerumque usque 100 flm longitudine per 2.0-3.0 flm in diametro, cum
apicibus tenus 5 flm inflatibus; penicilli pleraque monoverticillati, rami rari; phialides
pauper numero, 2-6 per verticillum, plerumque 5-11 flm longitudine per 2.5-4.0
flm in diametro. Conidia globosa vel ovoidea, plerumque 2.0-3.0 flm in dimetro, in

intricatis catenis tenus 100 .tm longitudine, parlete parce esperulato .

Coloniae in agaro Czapekii cum extracto levedinis, post duas hebdomadas ad
20-25C, 55 mm in diametro attingunt, velutinea, fortiter radiatae plicatae , denso
texto basali coacto mycelio, fortiter sporulantes, griseo-coeruleae vel griseobrunneae, zonatae; exsudatio et odor absunt; pallido succineo colore in medio
diffluente; reverso cremeo pallido.
Coloniae in agaro hordeaceo celerior quam in agaro Czapekii crescunt, post duas
hebdomadas ad 20-25C, 50 mm in diametro attingunt, planae, velutinae, margine
aequaliter, plerumque a hyphis submersis formatur; copiosae sporulantes,
superficiebus conidiorum griseo-viridibus; exsudatio abest; odor nullo; reverso
flavo; colore flavo in medio diffluente.
Typus cultura CBS 162.42, e solo tumulo arenoso Galiae a Drs F. et V. Moreau
isolata est in anno 1941.

Our notes follow :

Colon ies upon CA (colour plat e 13, A and a ) growing m od erately , a ttain ing
a diamet er of 35 mm in 14 days at room te m pe rature (20 - 25 OC ), consisting of
a dense ba sal felt , velvety in ap peara nce, very slightl y rad ia tely fu rrowed ,
1 mm deep at colony ce n tre, covered by a n ove rg ro wth of wh ite ae r ial hyphae
influen cing th e gene ra l aspec t of th e colony , wh ich is at first white , assuming
gra yish tint s in age, very poorl y spo ring; uneven m argin surrou nde d by a
fringe of su bme rge d hyphae exte nd ing abo u t 1 to 2 mm beyond th e aeri al
growth; exudat e limited , in very minute drops ; odour lacking; pale 'am bercoloure d pigm ent diffusing into the surrounding agar; colony reverse (colour
plat e 13, a) in pale cr eam shades . Penicilli usually str ictly simple , with an
occasional branch ; th e stipes of th e con id iopho res are sho rt, erect, septa te,
arising eithe r from th e su bmerge d hyphae or from aerial vegetative hyphae , up
to 100 p,m in length by 2.0- 3.0 p,m in diameter, with apices enla rged in a
vesicle up to 5.0 p,m in diameter (fig . 13, A) ; phialides usually few in number ,
2-6 per verticil , mostl y 5.0- 11.0 p,m by 2.5 -4 .0 p,m ; conid ia globose to
su bglobose, smoo th wall ed wh en yo u ng, but becoming cons p icuo us ly
rou gh en ed at m aturity (fig. 13, B, and SEM plat e 3, (top : leftj ); coni d ial
chains short and tan gled, up to 100 p,m long .
C olonies upon CYA (colour plate 13, B and b) gro wing m or e rapidly th an
upon C A , att aining a d iameter of 55 mm or m or e in 14 days a t room
temper ature (20- 25 OC), velve ty, deepl y radiat ely furro wed, fold ed in colony
centre, with superfi cial cre vices, cons isting of a tough , close-textu re d ba sal felt ;
heavier spo ring in bluish gray (Sguy , PI. XL , 598) to gr ayish brown shades
(Sguy, PI. XXXV , 5 19) in conce n tric al zone s; white gro wing m argin
5- 10 mm wide, with a frin ge of su bme rged hyphae extend ing largely beyond
th e ae rial gro wt h; odo ur lacking; exudate in minute drop s; pale amber
colou re d pigment diffusin g into th e su rrou nd ing agar; colony rever se (colour

lat e 1:" b) in pal e ora ngl: lo crcarn shndcs: pcni cill i as describod abov... .
Colonies upon M EA (eolour plate 13 , a nd e) spread ing b road ly, attaini ng
d iameter of 40 - 50 mm in 14 da ys a l room tempcrature (20 -25 OC ) , planc ,
leeply velvet y , heav ily spori ng throu gh ou t in m ou sc-gray shade s (Sgu y , PI.
{XI , 307), eve nly m argin in lighter gray tin ts , with a frin ge of su bmerge d
iyphae exte nd ing largely beyond th e ae rial gr owth (about 4 mm); odo ur a nd
.xu d a te absent ; pale am be r eoloured pigm ent di ffusin g int o th e su rrou nd ing
19ar; rever se (eolour plat e 13 , e) in pale or ange shades . Penieilli as deseribed
ibove but u su all y a rising direetly from th e su bs tra tu m, and with eonid ial
ch ains forming loose to tangled eolu m ns .
Sp eeies deseription is eente re d upon th e typ e stra in CBS 162.42 = ATCC
18 ,228 = Q M 8, 150 = IJ FM 5010 , isolat ed from sa nd dunes by Drs F . and V .
M or eau, in Franee, in 1941.

14. Penicillium luteo-aurantium Smith: fig . 14 , eo lo ur pl a te 14, and

S E M pIate 3 (top : ri ght).

00 0

00 0

Gas o ooo
O
0

O O

; 0 o
lJ.J.J.il.uu.J

10

jrrn

Aj-~
L---J
)Jm

100

iR.

Fig. 14. Camera lucida

drawings of Penicillium
luteo-aurantium Smith, CSS
260.64. A , Detailed drawings
of the penicilli. S, Mature
conidia. e, Habit sketches.

Penici//ium luteo -sursntlum Smlth - Trans, Brlt, Mycol. Soco 46: 331. 1963. See
also M.M. Kulik, 'A Compilation of Deseriptions of New Penici//ium Species' .
Agrie . Handbook No . 351. Agrie . Res. Ser. U.S . Dep. Agrie. , p. 50. 1968.

Sm ith's original description of th e species is given below since this

adequately presents the species in question . T he typ e strain CBS 260 .64,
m aintained in artificial culture since its isolation h as become pred omin antly
ste rile, no lon ger show ing th e typical cultu ral characteristics originally
described by the au thor, although still showing characte ristic de tai ls of
morphology .
Sm ith's diagnosis as follows:
'Colonies on CA grow fairly rapidly, eventually spreading widely, are velvety, at
first bright yellow, then somewhat deeper yellow, slowing developing irregular
patches of gray-green, with colourless fimbriate margin and reverse broadly zoned,
bright orange and eventually almost scarlet; penicilli almost monoverticillate, but
sometimes with small cluste rs of metulae, i.e. verging towards the P. citrinum
series, sometimes more truly divaricate, with sparing but irregular branching;
eonidiophores smooth, arising from creeping aerial hyphae, short or very short ,
mostly 10-40 Ilm, seldom longer, and 2.0-2.2 Ilm in diameter, slightly swollen at
the tip if no metulae are present; metulae when present 11-13 Ilm by 2.8-3.0 Ilm ,
slightly swollen at the tip; phialides with fairly long tapering necks, but defin itely
acuminate, as the Biverticillata-symmetrica, 9-11 Ilm by 2.5-3.0 Ilm; eonidia
subglobose to broadly ovoid, delicately roughened, mostly 2.9-3.0 Ilm by 2.0-2.5
Ilm , but occasionally 4.0-5.0 Ilm by 3.0-3.8 Ilm in diameter, in chains which are
first parallel then tangled .
Colonies on malt extract agar grow rapidly, sporing more readily than on CA, but
otherwise similar .'

M easuremen ts obtained in our observations follow :

Colonies on C A (colou r plate 14, A and a) growing moderately, attaining a
diameter of 40 mm in 14 days at room te mperature (20 - 25 OC), consisting of a
to ug h, close-textured basal felt , with a strongly convex surface, snow-white ,
cove re d by a network of trailing hyphae and ropes of hy phae admixed with
large drops of an ab undant, colourless ex udate, wit h conidial structures very
lim ited in nu mber ; odo u r lacking; colony reverse (colour plate 14, a) colourless; no coloured pigment diffu sing into the surrounding agar; pen icilli mos tly
strictly sim ple (fig. 14, A) , wit h an occasional branch ; phialides taperin g to a
fairly lon g neck suggestive of those of the Biverticillat a- sym metrica section,
9- 11 p-m in length by 2.5 -3.0 p-m in di amet er; con idia globose to subglobose
2-3 p-m by 1.5- 2 .0 p-m , de licately ro ughened when observed under th e light
m icro scop e (fig . 14, B), but showi ng ro ug hness in a cerebriform patt ern whe n
observed under th e SEM (plate 3 (top: right)); con id ial chains forming short
parallel columns (fig . 14 , C).

C olon ics upon C YA (colour plat c 14, B and b) uttai ning a di a mcter 01'

:15 mm in 14 days at room tcmpcraturc , csscn tially duplica ting in cu ltural

characteristics tho se grown upon C A , but with largcr d rop s 01' exudate in
pinkish shades; margin surrounded by a fringe of sub me rged hyphae exte nding
mo re than 5 mm beyond th e aeri al gr owth ; od our lacking; colony reverse
(colour piate 14, b) in pal e yellow shades; very poorI y sporing. Penicilli as
dcsc ribed aboye.
Colonies upon MEA (colour plate 14, C and e) spreading broadly, attaining
a diamet er 01' 50 mm in 14 days at room temperature (20-25 OC ), plane,
poorIy spo rin g ; otherwise sim ilar in gen eral colony pattern to those grown on
C A and CYA . Peni cilli as described ab oye.
C olonies on CA and CYA becoming sulphur yellow in age , with margin
becoming light blue-green as sporu lation progresses in age .
Colon ies on MEA becoming al so sulphur yellow in ag e , with reverse in
ora ng e-yellow shades .
The corre ct placement 01' thi s species is still ope n to question. In view 01' th e
act that sorne penicilli a re di varicat ely branched , it could be assigne d with th e
Asymmetrica - divaricata sub-section (Raper and Thom 1949), but on th e
other hand , on accou nt 01' its predominantIy sim ple penicilli, it would need to
be keyed in th e Monoverticillat a section .
Spec ies de scription is cen te red up on th e type strain CBS 260.64 = CMI
92 ,533=ATCC 18 ,480=IFO 8 ,17 6=IJFM 505 1, isolat ed from a sa m ple 01'
' R uaka peat' from New Zealand by Mrs S . M arcu s, at th e London Sch ool 01'
Hygien e and T ropical Medicine , in 1950 .

15. Penicillium abeanum Smith: fig. 15 , co lo u r plate 15 , a n d SEM pl ate 3,

(ce n t re : left) .
Penici/lium abeanum Smith - Trans. Brit. Myeol. Soe. 46: 333. 1963. See also M.M.
Kulik, 'A Compilation of Deseriptions of New Penici/lium Species'. Agrie.
Handbook No. 351. Agrie. Res. Ser. U.S. Dep. Agrie. p. 11. 1968.
Synonym
Penici/lium trzebinskii Zaleski, varo magnum Sakaguehi et Abe - J. Gen. Appl.
Mierobiol. Tokyo. 2: 62-63. 1956.

Colonies on C A (colour plat e 15, A and a) growing fairly rapidly, att a ining a
diamet er 01' 55 mm in 14 days at room temperature , consis ting 01' a loosetextured basal felt , covered by an overgrowth 01' ae rial vegeta tive hyphae
lending a su blana te appearance to the otherwise velvety colony surface , almost
plane, heavily sporing in sorn e area s in gray-gree n shades (Sg uy , PI. XXIX,
432) ; evenly m argin with sub me rge d m ycelia exte nding 3-4 mm beyond th e

{ :,~ '.;

Fig. 15. Camera lucida

drawings of Penicillium
abeanum Smith, CBS
348.59. A, Detailed drawings
of the penicilli. B, Mature
conidia. C, Habit sketches.

ae rial growth; od our and exudate lacking ; colony rev ers e (colour plat e 15, a )
colou rless; penicilli strictly sim ple (fig . 15, A), with stipes usu ally arising fro m
trailing aerial hyphae, so rne from su bme rge d hyphae , variable in len gth , up to
170-440 p.m by 2 .0-3 .8 tui, with apices enla rge d in a vesicle up to
3 .5- 5.0 p.m in di ameter, with walls smooth or delicately rou gh en ed ; phialides
numbering 5 to 8 in th e verticil, mostly 8 .0 - 15. 0 p.u: by 2.5 -4 .0 p.m , wi th a
short, de finite ly narrowed neck ; con id ia cons picuous ly rou gh en ed (fig . 15, B),
show ing a cere briform pattern whe n obse rve d under th e sca nning elec tron
mi cros cop e (S EM plate 3 (centre; left )) , mostly 4.0 - 5.0 p.m in di amet er,
globo se ; conid ial chains forming long, loose colu m ns (fig . 15, C).
Colon ies upon CYA (co lour plat e 15 , B and b) spread ing broadl y , attaining
a di ameter of65 mm in 14 days at room temperature (20-25 OC ), cons isting of
a similarly textured felt as described aboye but with sh allow radiate furrows ,
heavier spo ring in dark gray-gree n shades (Sg uy, PI. XXX , 446) in m argin al
areas; con id ial struc tu res obsc u red in su bmarg inal and central a reas by a n
ove rg ro wth of white aerial hyphae lending a lan at e appearance to th e colony ;

odo ur and exuda re lackin g ; co lony revcrsc (colou r plute 1;, b) in vc ry palc
ycllowi sh tint s , Peni cilli as desc ribcd a boye.
Colon ies O I! M EA (co lour pla tc 15, C and e) gro wing stilJ more rapidl y th an
up on the aboye substrata, a ttaining l diarnete r 01' 70 mm in 14 days at roo m
tempcrature (20-25 OC), vclvet y in appearance but cove red by an abu nda nt
ove rgrowth of ae rial hyph ae lending a lan at e appeara nce to the colony surface;
heavily sporing in dark gray-gr een shad es (Sguy, PI. XXX , 446), bu t con idial
structu res are partly obsc ured by the mention ed ove rg rowth; exudate a nd
od our lacking; penicilli as described abo ve but with coni dial chains forming
well-defin ed com pac t colu m ns.
Penicill ium abeanum Smith was isolat ed fro mJapa nese soil by S . Ab e in 1952,
and described by K. Sa kaguchi a nd S . Ab e (Abe 1956) as P. trzebinskii Zaleski
va l'. magnum, a nd red escribed by G . Smith ( 1963) as P. abeanum.
Species descript ion is centere d up on th e type strai n CB S 348.59 = C M I
68 ,222 = I Fa 6,239 = IJ FM 5048, being in sa tisfacto ry ag ree me nt with the
original spec ies di agn osis.
16 . Penicillium odoratum C h riste nsen et Ba ckus : fig . 16, co Io ur pI ate 16 ,
a n d SEM pI ate 3, (ce n t re: ri ght) .

'~$~f~ ~'
'~I r~G
~
jrm

10

()

R.

100 ).1m

Fig. 16. Camera lucida

drawings of Penicillium
odoratum Christensen et
Sackus, CSS 294.62. A ,
Detailed drawings of the
penicilli . S, Mature con idia.
C, Habit sketches.

Penicillium odoretum Christensen et Backus - Mycologla. 63: 469462. 1961.

Colonies upon CA (colour plate 16, A and a) sprcading broadly , attaining a
diameter of 50-55 mm in 14 days at room ternperature (20-25 OC), con sisting
of a close-textu red basal felt, deeply velvety to floccose ; fimbriate, thin margin,
colou rless, with coarse submerged hyphae extending largely beyond the aerial
growth (about 4 mm); heavily sporing, mostiy in cen tral areas in deep bluegreen shades (Sguy, PI. XXX, 447) ; colony surface deeply and radiately
furrowed, and slightiy raised ; exudate lacking; odour penetrating, aromatic,
suggesting apples; colony reverse (colour piat e 16, a) in pale cream rosy
shades; penicilli strictiy simple (fig. 16, A), with stipes of the conidiophore
arising primarily from submerged hyphae, up to 100-500 J-tm long by
3.0-4.0 J-tm in diameter , septate , either smooth to coarsely roughened , with
apices enlarged in a vesicle commonly up to 7- 9 J-tm in diameter; phialides
crowded in parallel clusters of 10-18 in the verticil , typically 9.0-12.0 utt: by
3.0-4.0 J-tm; conidia elliptical at first, then becoming ovate to subglobose,
commonly 3.5-4.0 J-tm by 2.8-3 .2 tui, thick-walled, with roughness
concentrated in transverse ridges so that conidia appear faintiy banded when
observed under the light mi croscope (fig . 16, B) and with a cerebriform pattern
when observed under th e SEM (SEM plate 3 (centre: right)); conidial ch ains
forming well-defined columns up to 500 J-tm long (fig . 16, C) .
Colonies upon CYA (colour plate 16, B and b) spreading broadly , attaining
a diameter of 50-55 mm in 14 days at room temperature (20-25 OC), de epi y
velvety to floccose, strongly radiately sulcate, with centre d epressed ; heavily
sporing throughout in blue-green shades (Sguy, PI. XXX, 438); growing
margin white, 5-10 mm wide ; exudate lacking; odour penetrating, fruity; pale
yellow coloured pigment diffusing into the surrounding agar; colony reverse
(colour plate 16, b) in cream shades; penicilli as described aboye .
Colonies upon MEA (colour plate 16, C and e) growing a little less rapidly
than upon th e aboye described substrata, attaining a diameter of 35-40 mm at
room temperature (20-25 OC ) , almost plane but with few shallow furrows,
deeply velvety to floccose, with uneven margin, com m only developing
submerged stolon-like hyphae giving rise to peninsular ridges of stipes in
advance to the colony mat; heavily sporing throughout in deep blue-green
shadcs (Sguy, PI. XXX, 446); exudate lacking; odour strong, aromatic,
resembling aroma of apples ; colony reverse (colour plate 16, c) in yellow
shades; amber-yellow-coloured pigment diffusing into the surrounding agar.
Penicilli as described aboye, with stipes arising primarily from stolon-like
hyphae.
Penicillium odoratum Christensen el Backus resembles certain members of the
P. lividum series, especially P. lrzebinskianum Abe ; the latter species, however ,
develops a darker coloured reverse upon CA, a mouse-gray colour upon MEA,

a nd has a vun illa -likc odo ur instcad 01' the apple-likc aro ma so cha ractc ristic 01'
P. odorat um, has sho rte r stipcs (60- 2110 .u n lon g) with som cwh at sm aller
ves icu lar api ces, a nd does not fo r m stolon-like hyph ae up on C YA a nd M EA.
Spe cies dcscription is ce nte rcd up on the ty pe strain C BS 294.62 = ATCC
14,769 = F 7.741 = IJ FM 5005, isolat ed from the pcat y soil 01' a spruce -Iarc h
com m u nity in Taylor Cou nty, Wi scon sin, U SA, by M. C hriste nse n and M .P.
Backu s, in 1960 .

17. Penicillium tarraconense Ramrez el Martnez : fig. 17, caIaur pIate 17,
and SEM plate 3, (ba tta m : Ieft).

". '

.,

"

"
"
"

l.l..l.u..WJJJ

10 prn

o
O

00
sO O

O()G O O

0 0 0 ()

()

~-,
,~\'Ol CK

Fig. 17. Camera lucida

drawings of Penicillium
tarraconense Ramrez et
Martnez. ATCC 42,235. A,
Detailed drawings of the
penicilli . S, Mature conidia.
C, Coiled hyphae . D, Habit
sketches.

Penicillium tarraconense Ramrez et Martnez - Mycopathologia. 72: 181-191, 1980.

Coloni es upon CA (colour plate 17, A) grow in g rudimentarily, probably due

to a vita m in defi cien cy , cons isti ng mo stly 0 1' submerged hyphae fro m whi ch
arise a few eon idial struc tu res. These rudimentary colonies a ttain a di amet er 0 1'
25 mm in 14 days at room temperature (20- 25 OC ); the stipes 01' th e spa rcely
produced eon id iopho res are eree t, smooth to rough -walled, gene ra lly less th an
100 p.m in length by 2.5-4.0 p.m in di amet er, with apices slightly enl arged in a

vcsi clc IIp to 5 .0 mI in di um ct c r: phi ali d es bo ttlc-sh np ed with sho rt -poin tcd
neck , co m m o n ly 10.0 -1 5.0 mI in lcn gth b y :U l- 1. 0 LIlI in diumct cr,
di vcr gent , 6- 10 in thc vet icil (Iig. 17, A) ; co nid ia smoo th-wallcd 0 1' ncarl y so
(lig. 17, B, and SEM plate 3, (bottom : left , clliprical , usu all y 3. 0 - 1 .0 un by
2.5- 3.0 JL m in diamete r; co nid ial chains tangled and penicilli str ictly simple
(lig . 17, D ) .
Colonies upon C YA (colour plat e 17, B a nd b) gro wing luxuriantly,
attai ning a di amet er of 60 m m in 14 d ays at roo m temper ature (20-25 OC),
velvet y , almost plane , compact, close- textured , sligh tly rai sed in central a re a;
heavily spo ring th ro ug ho ut in yellowish to gray-gree n shades (Sguy, PI.
X XIX , 424) in margi nal a rcas, and in pale oc h re shades in cen tre; the colony
surface is covered by a n ove rg rowth of loose floccose hyphae , especially in
cen tra l areas; ex udate lacki ng ; odo u r faint , indelinite ; colon y reverse (co lour
plat e 17 , b) b rown in ce nt re and in oc hraceous shades in m argin al a nd
submarg inal a reas; a m be r-colou red pigm ent d iffus ing into th e surro u nding
agar. No a scomata 01' scle ro tia have been obse rved, but a few structures
sugges ting ascomata initials have been observe d (lig. 17, C). Penicilli strictly
sim ple, with lon g stipes up to 200 JLm by 2.5 - 4. 0 JLm co m monly borne fro m
submerged h yph ae , ei the r smooth 0 1' ro ugh-walled, as described aboye .
Colonies upon MEA (co lour piate 17, C and c) gro wing rathe r m od erat ely ,
atta ining a dia m et er of 40 mm in 14 da ys at roo m temper ature (20- 25 OC),
pla ne , ve lvety, loose textured and sligh tly zo nate; heavil y sporing throu ghout
in yellow- gr een sha des (S guy, PI. X X IX, 424); thin , eve n marg in su rro u nde d
by a fringe ofs ubmerged hyph ae exte nd ing 3-4 mm beyond th e aerial growth;
exudate not ob served ; odour fragrant ; revers e colou rle ss with a gree n cas t
show ing throu gh ; no co lou re d pi gm ent di ffusin g into th e su rrou nd ing agar
(co lour pla te 17, c) . Penicilli as aboye.
Penicillium terraconense R amrez el M artnez a ppears m ost closely related with
species in cluded in th e P. frequentans se ries on accou n t of its str ictIy sim ple
pen icilli , co m paratively la rge , with lon g and erect stipes a ris ing di rectl y fro m
th e su bstrat u m, of th e velve ty a ppeara nce of th e colon ies, and of th e lu xuriant
growth on most m edi a use d, but det ail ed in vesti gati on s fail ed to m at ch it with
any of th e speeies in th e series.
Species di a gn osis is based on th e type stra in IJFM 5151 = ATCC 42,235,
isolat ed from Spanish must , M ad rid , 197 8 . It is at presen t known only fro m
th e typ e strain.

Penicillium lividum se r ie s
UT ST A NDIN G FEATU RES

Colonies a ppearing deepl y velvety to lanose , co nsisting of a tough ba sal felt

producing a bu ndant co nid ia l struc tu res in delinite blue-green shade s.

r ile stipcs 0 1' the conidiophores are ercc t 01 ' asccndi ng , lon g , usua lly
unb run chcd , w ith wa lls usually delica tcly rou g hcn cd a nd with a p ices
so rncw hat enlarged in a vcsicle .
Pcni cilli strictly simple with phialides usu ally cro wde d in the vert icil, parallel
0 1' div crgen t .
C o nid ia elliptical to su bglobo se , mos tly 3.0- 4. 0 Jlm in lon g axis, with wa lls
echinu la tc, often cons picuously .

Key to species
1. Colonies deeply lanose, stipes very long, up to 400 p.m 01' more; reverse
in orange to ochraceous shades.
a. Penicilli with phialides comparatively large, up to 14 p.m in length
1' . Conidia compa ratively large, up to 6 p'm in long axis, conspicuously roughened.
P. lividum Westl ing
2'. Conidia comparatively small , up to 3.5 p'm in long axis, with walls
smooth 01' nearly so.
P. valentinum Ramrez et Martnez
b. Penicilli with phial ides comparatively small, up to 10 p'm in length.
Conid ia elliptical with pointed ends, with walls delicately roughened.
P. aurantio-violaceum Biourge
2. Colonies not deeply lanose, deeply velvety, stipes comparatively short.
a. Penici lli with phialides comparatively large, up to 10 p'm in length;
odour suggesting aroma of vanilla ; reverse in pale ochraceous shades.
P. trzebinskianum Abe ex Ramrez
b. Penicilli with phialides comparatively small , up to 8 p'm in length;
odour lacking 01' indefinite; reverse in deep violet to violet-black
shades.
P. trzebinskii Zaleski

Page

. . . . 72

. . .. 80

. . . . 74

. . . . 78

. . . . 76

The members 01' th is series represent a normal com po ne nt 01' th e fungu s flora
01' soils, but occurring less frequently tha n othe r sim ple forms. T he seri es
actually com prises five well-d efin ed spec ies:
Penicillium lividum W estling, easily recog nized by its deep velv ety , almost
lanate colon ies, its typicall y blue-green colou r, and its conspicuo u sly rough walled , elliptica l conidia . Penicillium aurantio-violaceum Biourge , although
presenting m u ch the same cultu ral cha rac te ristics, differs, however, markedl y
in producin g strongly elliptica l con id ia, usuall y wit h poi nted ends . Penicillium
trzebinskii Zaleski, characterized especially by the colony revers e in deep violet
to violet-b la ck shades. Penicillium trzebinskianum Ab e differs from P. trzebinskii by
the produ ction 01' a strong vanilla-like a rom a, its com paratively large penicilli,

its Iaster grow ing colon ies up on C YA, an d thc prod uction 01' l rcddi sh brown
pigm cnt in thc colony rcver sc, which diffu ses into thc su rro u nd ing agar, upon
MEA . And finall y, Penicillium oalentinurn Rarnfrez et M artfnez , more closely
related to P . lividum West1ing, characterized by its dceply velvety , almost
lanat e colonies, its com pa ra tively large peni cilli , and the ora nge sha des
developed in colony reverse. The seri es appears to be related to the P.
frequ entans series throu gh P . trzebinskii, P. sp inulosum and P. odoratum .

18. Penicillium lividum W estling: fig. 18, colour plate 18, and SEM plate
3, (bottom: right) .

'::-" .:';'),

6::01 '<;~I

LwJ..lw.l.I \

10

urn

-1~1/f'
~

100

jrrn

Fig. 18. Camera lucida

drawings of Penicillium
lividum Westl ing, ATCC
10,102. A, Detailed drawings
of the penicilli. B, Mature
conidia . C, Habit sketches .

Penicillium lividum Westling - Ark. Bo1. 11: 58, 134-137. 1911. See also Dale, Ann.
Mycol. 12: 52. 1914; also Thom, 'The Penicillia' , pp. 205-206. 1930; and Raper
and Thorn, 'A Manual of the Penicillia', pp. 190-192. 1949.

C olonies up on CA (colour piate 18, A and a) attaining a diame ter 01'

40-45 mm in 14 days at room temper ature (20- 25 OC), plane , consisting 01' a

iou gh basal fclt, loosely lextured al the colon y sur lace whi ch is decply velvcty
l o almos t lanate , a t Iirst white, then assurning deep blu e lo blu e-green shade s
in heavily sporing zon cs (Sguy, PI. XXXI, 461 -463), slightly zonate, with
undulating margin white , surrounded by a narrow fringe 01' submerged hyphae
cxtcnd ing about 1 mm beyond the aerial growth; exudate lacking; odour
nhsc nt or indefinitc ; colony revers e (colour pIate 18, a) in orange to peach
shndes; pale arnber-coloured pigment diffusing into th e surrounding agar;
pcni cilli mostly strictly sim ple, with stipes 01' th e conidiophores mo stly arising
Irorn the substratum , usually unbranched, comparatively very long, up to
(iOO .tm or more in len gth by 2.5-4.0 .tm in diameter, septate , with walls
xmooth or nearly so , with apices enla rged in a vesicle up to 5.0 -6 .0 .tm in
rlin m ete r (fig . 18, A) , or arising as short branch es from a network 01' aerial
hyphae; phialides in parallel clusters or somewhat divergent, mostl y 5-10 in
(he vcrticil , comparatively large , 11.0-16 .0 tut: by 2.5-4 .0 .tm, terminating
in a fairly conspicuous tapered neck; conidia at first elliptical, then becoming
usually ovate to subglobose at maturity, mostly 4.0-5.0 tut , occasionally up to
(i.O .tm by 3.0-4 .0 tuu, with walls conspicuously roughened (fig . 18, B) ,
usually in spiral bands when observed under th e SEM (plate 3, bottom : right) ;
conidial ch ains short and tangled (fig . 18, C).
Co lonies upon CYA (colour plate 18, B and b) spreading more rapidly than
upon CA, attaining a diameter 01' 55 mm in 14 days at room temperature
(~ O-25 OC ), essentially duplicating in colour , cultural ch ar acteristics and
det ails 01' morphology those described up on CA, but deepl y radi ately furrowed
with centre depressed; exu date a nd od our lacking; colony rever se (colour plat e
In, b) in darker orange or cinnamon shade s; amber coloured pigment diffu sin g
into the surrounding a ga r. Penicilli as described aboye.
Colonies upon MEA (colour plate 18, C, and c) growing fairly rapidly ,
a ttainin g a diameter 01' 55-60 mm in 14 day s at room temperature, plane,
dcepl y velvety to Ian ate , he avil y sporing throughout in deep blue-green shades
(Sguy, PI. XXX , 446) ; od our sIightly ' mou ldy'; exudate Iacking; pale am be rcoloured pigm ent diffusing into th e surrounding a ga r; revers e (colour plate 18,
c) in orange shades ; penicilli as described aboye but with the stipes 01' the
con idiophore s arising directly from the su bs tratu m, and comparatively very
lon g, up to 1000 .tm or more , with walls frequently roughen ed in terminal
a reas; penicilli av erage a little larger than upon CA; con idia more cornmonly
rem aining ellipticaI at maturity, otherwise as described aboye.
The species is centered upon the type species ATCC 10,102 = CBS 347.48 = CMI
39, 736 = IFO 6 ,102 = NRRL 754 = IJFM 5,155 , isolated from soil in Scotland by
Miss E. Dale , in 1913, Westling' s type strain being lost before 1911. It constituted the principal basis for Thom' s redescription 01' Westling species (1930).
An other cultu re received from the Centraalbureau voor Schimmelcultures ,
Baarn , The Netherl ands, labeled Penicillium lividum W estling CBS 217 .30 ,

cn n no t b e rcgurded as snt isla ctorily rc prcs c n ta tiv e 01' th e sp ccies becau sc it

dill crs Irom th c ub ov e d cscription in its c u ltu ra l ch a r nc tc ris rics ; n amcl y , th e

produ cti on of (hin , closc-tcxtu red , more rcs tri ct cd colonics up on C A, a ttain ing
a di amet er of 25-30 mm in 14 day s at roo m tempcr ature ; peni cil1i with sho rt
stipes, ra rely more th an 50-100 tm lon g. Upon M EA th e colo nies a re
spread ing b ro adl y , a tta in ing a diamet er 01' 60 mm in 14 days a t roo m
temperature , plan e , heavil y sporing th rough ou t , with th e margin very
irregul arly di ssected from th e uneven gro w th 01' th e veget ative mycelium ,
whi ch is largely su b me rged . In colony charac te ristics, thi s peculia r strain fits P.
lividum only in th e colour 01' conid ial areas (Sguy, PI. XXXI , 461- 463),
deepl y blue-gr een . The penicil!i , however , agree q uite rea son abl y with the
a bo ve description ; thus this strain ca n no t be rem oved fro m th e species because
01' the co ni dial structu res produce d . A strain dupl icating in al! eharacte ristics
th e C ES 2 17.30 strain 01' P. lividum, strain IJ FM 5157, has been isola ted by th e
a u tho r from th e a ir in Mad rid, in 1978 .
Penicillium lividum W estl in g, altho ug h less ab u ndant th an many species, is
found amo ng isolates from soil, from sto re d cereal p roducts, and from org a nic
materials su bject to a ir 01' soil conta m ination, and is ap pare n tly cos mopo litan
in di stribution.

19. Penicill ium aurantio-v iolaceum Biaurge: fig . 19 , colou r pi ate 19 , and
SEM pla te 4 (top: left) .
Penicil/ium aurantio-vio/aceum Biourge - Monographie, La Cellule. 33: 282-284.
1923. See also Thom, 'The Penicillia' , p. 208. 1930; also Raper and Thorn, 'A
Manual ot the Penicillia', pp. 192-194. 1949.

Synanym
Penicil/ium roseo-viridum Stapp et Bortels - Z. Bakt. Parasitenk. Int. Hyg. Abt. 11.
93: 51. 1935.

Colonies upon CA (co lour plate 19 , A and a) sp read ing b road ly, attain ing a
d iameter 01' 55 -60 mm in 14 d ays at room temperature (20-25 OC), plane 01'
with shal!ow furrows , azo nate to sligh tly zona te , com para tivcly deep up to
1.5 mm in cent ra l arcas , co ns isting 01' a tough m ycelial felt at th e su bs tratu m
level , with su rface loose-t extured , deepl y vclv et y to lanate , formed by lon g ,
erect stipes; evenly m argin white , 10 -1 5 mm wide , rather thin, with
su b me rged hyph ae ex te nd ing a bo ut 2 mm beyond th e aeria l gro wt h; heavily
sporing th roughout in blue-green shades (Sg uy, PI. XXX , 443), re m aining
almo st unch anged in age; ex udate lacking ; odo u r faint , indistinct ; colony
reverse (co lo ur pla te 19 , a) in light cream to pal e salmon- pin k shades ; pen icil!i
strictly sim ple (fig. 19, A), with stipes 01' th e con id iophore s arising prim arily

lWJ.1Wl.J

10 jrrn

o"

00

'Vii/X} O 0 0 0 -;:OI'

C) B

000

R.. ) ...

c' ~

OO

(.

/'/.~ ./"

~}ifl[
100

}Jm

Fig. 19. Camera lucida

drawings of Penlclllium
aurantio-violaceum Biourge,
CBS 267.35. A, Detailed
drawings of the penicilli. B,
Mature conidia. e, Habit
sketches.

from submerged hyphae or from tr ailing ae rial hyphae , ascending up to

400 /lm or more in len gth by 2 .5-3 .5 /lm in diamet er, occasionally branched ,

with walls smooth to closely and delicately echinulate , with apices somewhat
enlarge d in a vesicle up to 4.5-5 .0/lm in di amet er; phialides in crowded
clusters of 8-12 or mor e in th e verticil, slightly divergent a t th e tip s, about
8.0-10.0/lm by 2.0-2 .5/lm , terminating in a definitely narrowed neck ;
conidia strongly elliptical, almost fusiform , about 3 .0 -3 .5 /lm by 2.0-2. 5 /lm
in diameter, with ends usually pointed, with walls som etimes appearing
smooth, wh en obs erv ed under th e light mi cro scop e (fig . 19, B), but delicately
roughened wh en obse rved under th e SEM (plate 4 , top: left) ; conidial ch ains
forming eithe r parallel or div ergent colu m n s, up to 100 /lm or more in length
(fig. 19, C) .
Colonies upon CYA (colour plate 19, B and b ) growing as rapidly as upon
CA, attaining a di amet er of 55 mm in 14 days at room temper ature
(20- 25 OC ), duplicating in texture th e colonies grown upon C A , but deeply
radiately wrinkled, less heavily spo ring in artemisia green shades (Sgu y , PI.

XXVI, :i90); odour a nd exuda re lacklng: umbcr-co lourcd pigmcnt dillu sing
into the su r ro u nd ing ag ur; colo n y reve rso ( COIOlll' plat e t 9 , h) in Iight orange to
salmo n-pink shadc s ; pcni cilli as describcd aboye.
C olonies upon MEA (colour plat e 19, C a nd e) sprcad ing broadl y, attaining
a diameter 01' 60 -70 mm in 14 days al room tcmperature (20-2 5 OC ),
consisting 01' a loose-t extured felt , with wid e gro wing margin surrounded by a
fringe 01' submerged hyphae extend ing 5 mm 0 1' more beyond the ae rial
gro wth; heavily sporing throu ghout in yellow-green shade s (Sguy , PI. XV ,
222) , remaining almost unchanged with age ; odour and exudate la cking ;
penicilli as described aboye , but with the stipes 0 1' th e conidiophor es mo stly
arising from th e substratum , com mo nly more th an 500 .tm in len gth ; conid ial
chain s forming loose colum ns up to 200 .tm long ; colony rev erse (colour piate
19, e) in pale orange shades ; no colou re d pigment diffu sin g into th e
su rro und ing agar.
Sp ecies description is cente red up on th e typ e stra in CBS 267 .35 = CMI
39,740=ATCC 1,041=NRRL 760=IFO 6,089=QM 7,485=IJFM 5019, isolated
from a beech-forest soil in Berlin (G ermany) by C . Stapp an d H . Bortels in
1935 , and us ed a s th e typ e a l' th eir new spec ies, Penicillium roseo-oiridum ,
actually conside re d as syno ny m ous with P. aurant io-violaceum ea rlier described
by Biourge .

20 . Penicillium trzebinskii Zal eski: fig. 20 , colou r plate 20, a nd SEM plate
4, (top : ri ght) .
Penicil/ium trzebinskii Zaleski - Bull. Acad. Polonaise Sci. Math. Nat. Sr. B, pp.
489-499. 1927. See also Thom, 'The Penicillia', p. 189. 1930; also Raper and
Thom, 'A Manual of the Penicillia', pp. 194-196. 1949.

C olonies upon CA (colour plate 20 , A and a ) grow ing fairl y rapidly,

a ttaining a diameter 01' 45 -50 mm in 14 days at room temperature
(20-2 5 OC ), co nsisting 0 1' a tou gh , close-t extured basal fclt , cove re d by a th in
ove rg row th a l' ae rial hyphae, giving a lanat e appearance to th e colon y surface,
deepl y radiately furrowed , with cen ter some what depressed , az onate, sporing
moderately throughout in blue-gr een shades (Sguy , PI. XXIX , 425) from a n
ad m ixtu re 01' co n id ial structu res and sterilc white ae rial hyphae; exudate and
odou r lacking; colony reverse (colour plate 20, a ) in pal e brown shades; pale
brown coloured pigm ent diffusing into th e surrounding a gar; peni cilli strictly
simple (fig. 20, A) , with stipe s 0 1' th e con id iopho res arisin g sometimes from
sub me rged hyphae , m or e com monly from th e basal felt 01' from ae rial hyphae,
variable in length, up to 150-200 .tm by 1.8 -2 .5 .tm in diameter, with walls
sm ooth 01' delicately roughened; philiades in verticils 01' 4-8 , rarely more ,
usu ally 8. 0 - 10. 0 utt: by 1.8 -2.2 .tm, parall el , terminating in a definitely

(~D

Uw.l.wLJ

:r-;~~O ~m
t '::;,
~;i ~::

()
(

B ;/t

....

fR.

[ rc r~"~\
L--.J

100 pm

Fig. 20. Camera lucida

drawings of Penicillium
trzebinskii Zaleski, CSS
382.48. A, Detailed drawings
of the penicilli. S, Mature
Habit sketches.
conidia.

e,

narrowed neck; conid ia broadly elliptica l to su bg lobose m ostly 2.0-3.5 tm in

long ax is, with wall s cons picuous ly echin ulate (fig. 20, B, a nd SE M platc 4,
top : righ t) .
Colonies u pon C YA (colour plate 20, B and b) growing fairly rapidly ,
a ttaining a di amet er of 55 mm in 14 days a t room temperature , deeply velv ety
to lan a te , deepl y radi a tely fu r rowed, azo nate , poor to m cdi um sporing in th e
sarnc sha de s as upon C A; exudate and odo u r lacking ; colony rever se (colou r
plat e 20, b) in da rk brown shades, with m argin al are as in deep blue-gr een
tin ts; dark brown-coloured pi gm ent di ffusin g into th e su rrou nd ing ag ar;
pcn icilli as describe d above .
Col on ies upon M EA (colour plat e 20, C and c) spread ing m or e rapidly th an
up on C A a nd C YA , atta ining a d iame te r of 60 mm in 14 days at roo m
temperature , plane , convex, deepl y velvety to sublanate , consisting of a loosetcxtured felt , a lm ost lanate , 1-2 mm deep , hea vily spo ring throu gh out in pale
blu e-gr een shade s (Sgu y , PI. XXX, 449) ; penicilli as descr ibed above but
with thc stipes of th e con id iopho re s arising primarily from ae rial hyphae,
usu ally 200- 300 tm in len gth by 2.0 -2.5 tm in di ameter, with apices

cnlurgcd in a vcsicle IIp lo 1,,0 - 1'.;' J~1Il in dluntctcr, with walls conspicuously
cchi nu la te ; philialides crowd ed in the whorl , up lo 12 01 ' more in thc verticil ,
othe rw ise as dcscrihcd above ; con id ia l chains in loo se colu m us u p lo 150 tun 0 1'
more in lcn gth . Ex udate lack in g ; odo u r in dc fi n itc ; colony reverso (co lou r pIa te
20 , e) eolou rless 0 1' pale eream shades .
Speeies d eseription is ee ntered up on th e typ e strain CBS 382.48 = CMI
39,749=ATCC 10,50 7 = N R RL 731 =IF O 6, 110 =QM 7,678=IJFM 5008 , isolat ed
fro m fore st soil in Poland by K . Zaleski, in 1927 .
The eor ree t plaeement of thi s speeies still remains some what questi on a ble . It
was placed b y R ap er and Thom ( 1949) with the P. lividum se ries on aeeo u nt of
its lon g , rough stipes a nd eeh in ula te eo nid ia, th at are usu all y broadly ellip tieal.
The ove r-all pietu re of the spe cies, however, suggests a close relat ionsh ip to P.
spinulosum Thom.
r.':.\
..

21. Penicillium trzebinskianum Abe ex Ramrez : .{1

. J~
' 0' . 21 , colour plat e 21 ,
and SEM plat e 4, (centre : left) .
. ;;, ~. ';~S .
.'.':

lo. _

..~"i1;; ,:
.' .1<:

'&;
.. ,"i.

.('-~ '~~ i:;'; ~

- .

'1

.: ~~:::~~:~:
: ~ -.
~~
":"''1
IT
\.
,
~~1
;
~ .. ~
~

l'!r '

t
,

10 'p r:n

. ,

e t/r' '.:

...... '

,<,l

, 4;"

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L..----.J

..

"

100 JJfT!-.. -.
;

.( 1\..
" , .-too
,
.
, '

-:

"

Fig. 21. Camera lucida

drawings of Penicil/ium
trebinskianum Abe ex
Ramrez, CBS 431.65. A,
Detailed drawings of the
penicilli. B, Mature conidia.
C, Habit sketches .

Pon/el/lium trzob/nsk/onum Abe - J. Gen. Appl. Mlcro-blol. Tokyo. 2: 63-64. 1956.

Saa also M. M. Kulik, 'A Compilation of Descriptions of Naw Pentcllltum Species .'
Agric. Handbook No. 351. Agric. Res. Ser. U.S . Dep . Agric . p. 75. 1968.

Penicillium trzebinskianum Abe was isolated frorn soil in J apan by S. Ab e , in

1956 , who proposed th e name P. trzebinskianum and described it with out
providing a Latin di agn osis . A Latin diagnosis is given bel ow : th e name of th e
ori gin al author is m aintained under Recommendati on 46 C of th e Interna tional Code of Bot anical Nomenclature (Stafleu (Ed .) , 1972 ).
Latin diagn osis of Penicillium trzebinskianum Abe ex R amrez :
Coloniae in agaro Czapekii satis celeriter crescentes, post duas hebdomadas ad
20-25C, 50 mm in diametro attingunt, velutino-Ianosae, denso texto basali coacto
mycelio, radiatae plicatae, parum depressae ad centrum, griseo-viridae coloratae;
exsudatio abest; odor adest, similis vanillam; reverso cremeo; neque ascomata
neque sclerotia visa sunt. Penicilli pleraque monoverticillati; stipitibus erectis, longis,
plerumque usque 60-300 m longitudine per 2.5-4.0 m in diametro, glabris,
hyalinis, cum apicibus tenus 3.7-7.0 m inflatibus, plurimum e mycelio basali vel ex
substrato, parum ex hyphis aeris exoriuntur; phialides multae numero, 5-12 per
verticillum , plerumque 8.0-12.5 m longitudine per 2.0-3.5 m in diametro, cum
brevi et acuminato collulo; conidia ovoidea vel subovoidea, plerumque 2.5-4.0 m
in diametro maximo per 2.5-3.0 m in diametro minimo, aspera, in catenis longis
tenus 300 m.
Coloniae in agaro Czapekii cum extracto levedinis celeriter crescunt, post duas
hebdomadas ad 20-25C, 55 mm in diametro attingunt, similiter in textura quam in
agaro Czapekii, sed conidia in intricatis catenis .
'
Coloniae in agaro hordeaceo minus celerior quam in agaro Czapekii crescunt,
post duas hebdomadas ad 20-25C, 45 mm in diametro attingunt, planae, velutinae
vel lanosae; copiosae sporulantes, superficiebus conidiorum coeruleoviridibus;
exudatio exiguum; reverso flavo vel fusco; cum pigmento flavo in medio diffluente.
Typus cultura CBS 431.65 e solo Japoniae a S. Abe isolata est in anno 1956.

Colonies upon C A (colour pIa te 21 , A a nd a) grawing fairly rapidly ,

atta in ing a di ameter of 50 mm in 14 day s at room temperature , cons isting of a
some what compact basal felt , deep'v velve ty to su bla nate in appeara nce;
eve nly white m argin , 1.0-1 .5 mm wid e , su rro un de d by a fringe of su bme rged
hypha e exte nd ing abo ut 1 mm beyond the ae rial gro wt h, radiat ely fu rrowed,
with centre depressed ; sporing hea vily in dark bluc-gr een shades (Sguy, PI.
X X V II, 40 2) ; exudate lacking ; odo ur suggesting va nilla ; reverse (colour plate
2 1 a) in pale cream sha des; penicilli strictly sim ple (fig . 2 1, A), with the stipes
of th e con id iopho res usually arising in a close sta nd fro rn th e substratu m or
basal felt , occ as ionally from aerial hyphae , m ostly 60 -3 00 titt: lon g by
2.5-4. 0 /lm in di amcter, with apices enlarged in a vesicle up to 3.7 - 7.0 /lm in
d iame te r, with walls smo oth or deli cately punct at e , some times septa te;

phialidcs crowded in com pac t cluste rs 01' 5-12 in the verticil , mostly
8.0 -12 .5 usu by 2.0 -3.5 ust, terminating in a short , tapered neck; conidia
ovate to subglobose, mostly 2.5 -4.0 J.m by 2.5 -3.0 J.m, with walls
conspicuously echinulate or verruculose (fig . 21, B, and SEM pIate 4, centre:
left) ; conidial chains forming well-defined compact or loose columns (fig . 21,
C) , up to 300 J.m in length .
Colonies upon CYA (colour plate 21, B and b) spreading broadly, attaining
a diameter al' 55 mm in 14 days at room temperature (20-25 OC), consisting al'
a compact basal feIt, deeply velvety, covered by a loase overgrowth al' floccose
aerial hyphae giving a sublanate appearance to the colony surface, strongly
radiately furrowed, with evenly margin 2 mm wide; heavily sporing
throughout in the same deep blue-green shades as upon CA ; odour suggesting
vanilla aroma; exudate lacking; colony reverse (colour plate 21, b) in
cinnamon shades ; penicilli as described aboye, but with conidial chains loosely
parallel to tangled.
Colonies upon MEA (colour plate 21, C and c) growing less rapidly than
upon CA and CYA, attaining a diameter al' 45 mm in 14 days at room
temperature, plane, deeply velvety to lanate in appearance, heavily sporing
throughout in dark, mouse-gray shades (Sguy, PI. XXXVII, 547); thin
margin surrounded by submerged hyphae extending about 5 mm beyond the
aerial growth ; exudate limited, colourless , enmeshed in the mycelial feIt ; odour
suggesting vanilla; colony reverse (colour plate 21, c) in cinnamon to reddish
brown shades; bright yellow coloured pigment diffusing into the surrounding
agar. Penicilli as described aboye.
Penicillium trzebinskianum seems to have a clase relationship to the P. lividum
series on account 01' the deep velvety to floccose character al' its colonies, its
long and rough-walled stipes, and its broadly elliptical and echinulate conidia.
The specific name given by Abe reminds its similarities with P. trzebinskii
Zaleski , especially the echinulate character al' conidia and the pigment
developed upon CA.
Spe cies description is centered upon the type strain CBS 431.65 = IFa
6,038 = IJFM 5041, isolated fromJapanese soil by S . Abe, in 1956.

22. Penicillium ualentinum Ramrez el Martnez: fig. 22, colour plate 22,
and SEM plate 4 (centre : right) .
Penicillium valentinum Ramrez et Martnez - Mycopathologia. 72: 181-191, 1980.

Colonies upon CA (colour plate 22, A and a) growing rapidly , attaining a

diameter 01' 55 mm in 14 days at room temperature (20- 25 OC ), consisting 01' a
dense, close-textured basal felt, deeply velvety to sublanate in appearance,
with colony surface covered by an overgrowth 01' white aerial hyphae in central

u..wl.woJ

10 jrrn

00 0
00
00 O a C)
O o

Bo
O (J O 'f
O OC)

()

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e

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Fig. 22. Camera lucida

drawings of Penicillium
valentinum Ramrez et
Martnez, ATCC 42,227. A,
Detailed drawings of the
penicilli. B, Mature conidia.
C, Habit sketches .

areas; evenl y , white m argin surrounded by a fringe of submerged hyphae

exte nd ing about 1-2 mm beyond the aerial growth; colony surface alm ost
plan e , with exce p tio n of th e centre whi ch presents a sh allow circula r furrow
and a depression , followed b y a some wha t raised co n vex a re a ; heavil y sporing
throu ghout in blue-gr ay to blue-green sh ades (Sguy , PI. XXX , 443, 448),
ob scured in ce n tre by th e m ention ed white overgrowth of ae rial hyphae ;
exud ate and odour la ckin g; colony reverse (colour plate 22, a) in bright ora n ge
sh ades ; amber-coloured pi gment diffusing into th e surrounding agar ; penicilli
strictly simple , com parative ly large , with th e stipes of th e co nid iophore s long,
erec t, u su all y up to 300 uii: or m ore in length by 3.0- 4. 0 p.m in di amet er ,
com monly rough walled , ar ising m ostly from th e su bstratu m and from tr ailing
hyphae , with a pices enla rged in a vesicle up to 5 p.m in diamet er ; phialides
di vergin g , usu ally 8- 12 in th e verticil, bottl e shape d, with a short-poi n ted neck
(fig. 22, A) , mostly 10-1 5 p.m in length by 3.0 - 5.0 p.m in di amet er; co nid ia
ellipso idal, sm ooth walled or nearly so (fig. 22, E , a nd SEM plat e 4 , cen tre :
ri gh t) , 3.0 -3 .5 p.m by 2.3-2.9 p.m in di amet er; conid ial chains forming loo se
to well-defined com pact colu m ns.

Colo nics lIpon CY A (co lou r plat c 22, B a nd 1 sprcad ing b roadly an d
rapidly , a u a ining a di a mct er 01' G5 mm in 1'1 days a l roo m tcmpcraturc
(20-2 5 OC), vclvcty lo su b lanatc in appcaran cc , closc texturcd ; d ecpl y
radiately furro wed; m argin en tire with a narrow Iringe 01' su b me rgc d hyphae
ex te ndi ng abou t 1 mm beyond th e ae rial gro wth ; cen tra l are as depressed a nd
irregularl y fold ed ; heavily spo ring throu gh ou t in sim ila r blue-gr een shades as
up on CA; ex udate and odo ur laeking; reve rse in lighter ora nge sh ades (co lou r
plat e 22, b) ; no eo lou red pi gment diffu sin g into the surrou nd ing agar; pe nieilli
as deseribed above, with stipes 0 1' th e co nid iopho res smooth walled to defin itely
ro ughe ne d; eonidia l ehains tangled 0 1' forming we ll-de fine d eolu m ns.
Colon ies upon M EA (eo lour pl at e 22, C and e), a tta in ing a di amet er 01'
55 mm in 14 d ays at ro om temper atu re (20-25 OC), pl ane, velvety, with
centre sligh tly rai sed ; m argin eve nly wh ite , wit h su b me rge d hyphae extending
abo u t 5 mm beyond th e aerial gro wt h; heavil y spo r ing th rougho u t in bluegree n shades (Sguy, PI. XXX , 438, 449); eo lony surfaee eove red by an
ove rg row th 01' loosely interwoven floeeose hyph ae , influeneing th e eolony
a p pearanee; ex udate an d odo ur laeking ; eolony reverse (eolour pla te 22, e)
developing light ora nge shades ; a mber-eoloured pigment diffusing in to th e
surrou ndi ng agar; penieilli as deseribed a boye, with th e st ipes u suall y rou gh
wa lled, lon g , ereet, arising m ostly fro m th e su bs tratu m ; eo nid ial ehains
formi ng loose to eom paet eolu m ns .
Penicillium valentinum a ppears most close ly related to speeies included within
th e P. lividum series on aeeou n t 0 1' th e deepl y velo u tino us eharaete r 0 1' its
eolon ies, th e fast growth on a ll m edia tested , its long, rough- wall ed stipes, a nd
its ellip tiea l eoni d ia.
Speeies di agnosis is ba sed u pon th e typ e strain IJFM 50 71 = ATCC 42,22 7,
isola ted fro m th e air , M adrid , 1978. It is a t p resent kn ow n only from th e typ e
strain .

Penicillium implicatum ser ies

OUTSTAN DING FEATU RES

Colon ies growing restrie ted ly up on most substrat a , usu all y eo m paet,
velvety, heavily spo ring, m ostly in blue -green shades , with vegetative hyphae
sometimes ene rusted and p igmented; colon y reverse showi ng eharaete ristie
ora nge- re d to m aro on colo u rs.
Penieilli str ietly sim ple, usu all y bo rne upon stipes a ris ing di ree tly from the
su bst ratu m 0 1' from a d en se ba sal felt , eree t, seld om braneh ed , va ria ble in
length, eom mo nly less th a n 200 u it, but in sorne stra ins up to 300- 400 tuu ,
with a piees usu ally enla rged in a ves icle .
Con id ia l ehains lon g, loosely parallel 0 1' in eo m paet eolumns, ero wde d in
mos t spee ies and 01'ten tending to b reak away as erus ts in age.

:Illlid ill HIIIOo lh 01' delicutcly rou ghcn cd , clliptical lo Hu!>glo!>oHe 01 ' cvc n
ItlhuHl:, 1'1'0111 2.0 l o 2.5 {t lll in su me spcc ies to 4-.0 0 1' 5,0 {t ll l in oihcrs.

l' l' v
\,

lo

spccics

Page
rat her restrictedly upon most substrata, especially

P. implicatum series . . .. 82
lonias attaining a diameter at least of 20-30 mm in 14 days at room
tumperature (20-25C) .
1' , Conidial areas in pale blue-green shades, colony reverse in bright
oro nga-red or red shades, conidia globose to subglobose, in parallel
or divergent cha ins.
P. multicolor G.-M. et P. . . . . 84
' , Conidial areas commonly in deep blue-green shades, colony
rovarse in orange-brown or maroon shades, conidia broadly ellip tlcal, usually in compact columns.
P. implicatum Biourge . . .. 86
' , Conidial areas in yellow-green to gray -g reen shades.
aa. Colony reverse in orange-red (approaching brick-red) , conidia
strongly elliptical or pyriform .
P. sublateritium Biourge . . . . 88
bb. Colony reverse in greenish yellow shades .
1". Con idia small (less than 3 ~ m in diameter), smooth -walled or
nearly so.
aa. Conidia strongly elliptical or pyriform , smooth-walled.
P. ramusculum Bat ista et Maia . . . . 92
bb. Conidia globose to subglobose, with walls smooth or
nearly so.
P. aeneum Smith . . . . 90
2" . Conidia comparatively large, up to 4.5 ~ m in long axis ,
conspicuously roughened .
P. palmensis Ramrez et al. . . . . 94
4' . Conidial areas in green-black shades , colony reverse in bright
yellow shades, sulphur yellow pigment diffusing int o the surrounding agar, conid ia globose, conspicuously spinulose.
P. ardesiacum Novobranova . . . . 98
5' . Conid ial areas in gray -blue shades, colony reverse in yellow
shades, co nidia globose to subglobose, smooth-walled or nearly so.
P. ga//aicum Ramrez et al. . . . . 100
b . Colonies growing very rest rictedly, attaining a diameter not
surpassing 17 mm in 14 days upon Czapek's agar.
P. hispanicum Ramrez et al. . . .. 96

I'h e Penicillium implicalum series, according lo R uper ami T hom ( 1949),

re prese n ta a n aggrcga tc 01' SPCcil:S, nol idcntical in dciail s 01' stru cture 0 1'
reacti on , b u t having rnu ch more in com m on and gl'ad ing inro onc ano thc r
suffi ciently to justify gro u ping th ern logether. In acld itio n lo th e thrcc spec ies
ass igne d by th ese authors n am ely : P. implicatum Biourge , P. multicolor
G rigo rieva- Mano ilova et Poradiclova , and P. sublateritium Biourge , six m ore
spec ies d escribed as new to scien ce since 1949 ha ve been included in th e
present st udy within th e P. implicatum series, namely: P. ramusculum Bati sta et
Maia , P . aeneum Smith , P. palmensis R amrez , M artnez et Fe rre r, P. hispanicum
R amrez , Ferrer et Martnez , P. ardesiacum Novobranova , and P. gallaicum
R amrez , Martnez et Berenguer.
0 1' th ese spec ies, P. implicatum Biourge is b y far th e most ab u ndant and th e
m ost variable . M embers 01' this seri es occ ur regularly upon subs trata su bject to
soil 0 1' dust con tam ination.
Penicillium multicolor appears also in th e keys to species 01' th e P. fre quentans
se ries. C ultu ra l characte ristics and det ail s 01' m orphology locat e it in th e P.
implicatum series. H owever, th e tenden cy 01' sorne stra ins to grow rather rapidly
upon C A a nd othe r su bs tra ta co n ta ining vegeta ble ex tracts necessita tes keyin g
it with the P. freq uentans se ries also .
23. Penicillium multicolor Grigorieva-M anoilova et P oradi el o v a: fig . 23,

co lo ur pl ate 23 , a n d S E M pl a te 4 , (bot to m : left).

Penicillium multicolor Grigorieva-Manoilova et Poradielova - Arch. Sci. Biol.

Lenningrad . 19: 117-131. 1915. See also Ch. Thom, 'The Penicillia', pp . 212-213.
1930; also Raper and Thom, 'A Manual of the Penicillia', pp. 198-201. 1949.
Synonym
Penicillium implicatum Biourge, varo aureo-marginatum Thom - Ch. Thom, 'The
Penicillia', pp. 211-212. 1930.

C olo nies upon C A (co lou r plat e 23, A and a) u su all y growi ng rather
restrictedl y , attaining a di amet er 01'20-25 mm in 14 days at room te m peratu re
(20- 25 OC ), but in sorne strains, as it is the case 01' stra in C BS 35 1.48 stud ied
here (colour plate 23, A a nd a) , spread ing ra the r rapidly , attain ing a di amet er
01' 50 mm in 14 d ays at room temperature , co n sisting 01' a com paratively thi ck
ba sal felt, 1.0-1. 5 mm d eep , close textu red and fairly tou gh , velvety, with
shallow radial furrow s; heavily spo ring in sorne stra ins, poorly in others,
almost lanat e but presenting a velvety appearance , 0 1' showing both cond itions
in th e sa me colony ; with gro wing m argin 1- 2 mm wide , pal e yellow to bri ght
ora nge in colou r; in sorne strains co nid ia l structures developing in locali zed
cen tral 0 1' su bcen tra l areas agains t a background 01' yellow to ora nge-red

j't'r!o
B
.:~\ ~ -:.::

(}

luWJJ.u.J

10

jrrn

"lr L
l...--....J

100 jrrn

R.

Fig. 23. Camera lucida

drawings of Penicillium
multicolor GrigorievaManoilova et Parodielova,
CSS 351.48. A, Detailed
drawings of the penicilli. S,
Habit
Mature conidia.
sketches.

e,

VI'Wt;ltivc hyphae (co nsp icuous ly studde d with orange to orange-red gra n ules
wlu- n vicwe d under low magnifi cation ), in othe rs , heavily spo ri ng throu gh out
11 hlu c-gr een to deep bluish gray-gree n shades (S guy, PI. XXX , 443, 449);
I1 Xlll la l c limited to abundant, ye llow to pale ora nge; odo ur faint , suggesting
u iush ro orns ; colony revers e (colour piate 23 , a) in bri ght ora nge-red to burnt
Ie:lllla (Sguy , PI. XIII , 181 , 194) ; yellow to pa le ora nge-coloured pigm ent
t1 ifl'll sillg int o th e surraunding agar ; conid ial structu res abu ndantly produced
11 SClII 1C stra ins, sparingly in othe rs; peni cilli strictly simple (fig . 23, A), with
tilll'S 01' th e conid iophores m ostly arising directly from th e substratu m but
ol'l'as iona lly from aerial hyphae , usually unbranch ed , varying from short up to
lOO /U 11 in len gth by 2.0 -2 .5 /lm in diamet er, wi th walls smooth in so me
tra ins, gra n ula r in othe rs, or showi ng both cond itions in th e same strain, with
Ipiccs slightly enlarged or de finitely ves icula r in some stra ins, up to
i,O- :>. 5 /lm in di amcter ; phiali des in com pact wh orl s , usually 6 - 12 in the
vr-rt icil, in some strains m or e , mostly 8 - 10 /lm by 2.0- 3 .0 uiu , with apices
pnrall cl or di vergent ; conidia at first elliptical, becoming subglobose to

globosc, compurutivcly smull, nbout 2.0-2 .5

.UlI

in dium crer, with walls

d cli cat cly rou gh cn cd wh cn obsc rv ed 1I11<il:1' th c Iighl mi croscopc (lig . 2:1 , B) but
cons p icuous ly rou gh cn cd , a ppcarin g cc rc b rilorm wh cn ob scrvcd un d er th e
SEM (plate 4 , bott om : left) ; co n id ia l cha ins up to 100 ftllll on g , loosely parallcl
~ t an ~ ed (fig . 23, C ).
Colon ies upon C YA (colour plat e 23, B and b) spreadin g broadl y, a ttain in g
a di amet er of 60 m m in 14 d a ys at roo m tempcr a tu rc (20 - 25 OC ), essc nt ia lly
duplicating those grown upon C A in texture a n d gro ss colo ny appearancc , but
deepl y radiately furro wed ; ex uda te a b u nda n tly p roduced in bri ght orange
shades ; co lony reverse (co lour pia te 23, b) sim ila rly coloured in bright ora nge
shades; odo u r faint , suggesting mush rooms; bright yellow-colou rc d pi gment
di ffusing into the su rro u nd ing agar; penicilli as describcd a boye .
Colo n ies upon MEA (co lour plat e 23 , C a nd e), a ttain ing a di amet er of
30-45 mm in 14 d ays at ro om te m pe ratu re (20 - 25 OC ), a ppcaring velve ty to
subfloccose , com mo nly tufted in cen tral a re as in mi xed orange-red (fro m
hea vily e ncruste d hyphae) a nd blue- gr een sh ades (Sguy, PI. XXX , 443)
(fr om co ni d ia l struc tu res), with the former pred ominating in so rne strains, th e
latter in ot he rs, a nd cove re d by a n ove rgro wth of white vegeta tive hyphae in
sorne strains; growing m argin with hea vily e nc r usted orange-red hyphae
largely submerged , exte nd ing about 5 mm beyond th e ae r ial gro wt h; exudate
limited 01' lacking ; odo u r faint , suggesting mu shrooms; colony reverse (colour
piate 23, e) in bright orange-red colou rs (Sgu y, PI. XIII , 181); bright orangeyellow pi gment diffu sin g into th e surround ing agar; penicilli as described
a boye .
Sp ecies description is ce n te re d upon strain CES 35 1.48 = C M I 40, 574 = ATC C
10,471 = N R RL 2, 058 = QM 2,685 = IJFM 5040 , isolat ed from exposed textil es in
Florid a by Prof. W eston from H arva rd U niv ersity , Boston in 1946; also upon
stra in N R RL 2, 059 , isolat ed from a wat er pond in Brookl yn , N .Y . by Miss
Laura A. K olk fro m Brookl yn College, Brookl yn , N.Y . , in 1944; also upon
strain N R RL 2060 , isolat ed from ex posed cello phane in Florida , b y DI' L. White
in 1945.
Attempts to ob ta in th e typ e cu lture fail ed. Penicillium multic olor was isolated
or iginally from Ru ssian so il in 1915 by O .C . G r igo r ieva-Mano ilova a nd N .N .
Poradielova .
While it ca n not be ass u med with ce rta in ty th at th e stra ins assig ne d by Raper
an d T hom ( 1949) to P. multicolor G. -M et P . duplicat e in all det ail s th e strain
origin all y described by the au tho rs ofthis species, th er e is conside rable reason ,
accord ing to the forme r a u thors, for beli evin g th a t they must ha ve b as ed th eir
spec ies upon a for m similar to those co n side re d by them .

24 . Penicillium implicatum Biourge: fig. 24, colour plat e 24, and SEM

plate 4 (bott om : ri ght) .

o O
O O
00 BO 000
O
O O O O
O O
O

00
O

liliili.wJ

10 jarn

C
)
1
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/

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100pm

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Fig. 24. Camera lucida

drawings of Penicillium
implicatum Biou rge, CBS
232.38. A , Detailed drawings
of the penicilli. B, Mature
conid ia. C, Habit sketches .

Penicil/ium imp /icatum Biourge - Monographie, La Cellule. 33: 278-280. 1923. See
also Ch. Thom , 'The Penicillia ' , pp. 210-211. 1930; also Raper and Thom, 'A
Manual of the Penicillia', pp. 201-203. 1949.

Colonies upon CA (co lour plat e 24, A a nd a) growin g rather restrictedl y ,

attaining a di amet er va ry ing from 15 to 35 mm in 14 d ays at roo m temperature
(20-25 OC ), d ep ending upon the stra in , 2-3 mm deep or m ore , wit h growin g
rnargin thin , narro w and white , ofte n grow ing irregul a rly , gra d ually sp re ad ing
lo seve ra l ce nti me te rs with or wit hout traces of zo nation, close-textured , to ugh ,
vclve ty or nearly so; heavily sporin g in blu e-gr ay to blu e-gr een shades (S guy,
PI. XXX , 447) , sho wing sorn e tendency to form crusts in age in recent isolat es,
rad iately wrinkled with cen tre raised ; odo u r indefin ite , weak or ab sen t;
cx udate lacking or lim ited in amo u n t, in co lou rle ss dropl et s, yellowish or in
red -b rown shades; colony reverse (co lour plat e 24, a) va ry ing from yellowish to
orange , to deep red-brown or occasionally in purp lish shades; yellow-colou red
pigm ent diffusing into th e surrounding agar ; penicilli usu ally strictly sim ple ,
with an occasional branch , bu t always retaining th eir monoverticillat e

cha racter; stipes 01' th e co nid iophores sho rt , mostly under 100 ,un lon g by
2.0-2 .5 'LID in diaruc tcr , a rising usu ally from th e su bs tra tum in a den se sta nd,
occasionally from trail in g h yphae , with wa lls srnoo th 01' nearl y so (fig . 24, A) ;
phialides in cornpact clusters, m ostly 8 - 12 in th e ve rticil, ranging from 6.0 to
12 .tm by 2.0 -2.5 .tm; con idia com parativ ely small, elliptica l, 2.5-3 .0 .tm by
2.0- 2.5 uit) , m any subglobose, 2.5 -3 .0 utt, occasionalIy larger up to
4. 5- 5.0 tun, heavy-walIed , sm oo th 0 1' delicately rou ghened (fig . 24, B, and
SEM plate 4 , bottom : ri ght) ; con id ial chains in definitely compact to loose
colum ns up to 200 .tm 0 1' more in len gth (fig. 24, C) .
C olonies upon C YA (colour plat e 24, B and b) gro wing m or e rapidly th an
upon C A, a ttain ing a d iameter 01' 40 mm in 14 days at roo m temperature
(20-25 OC), deepl y radiat ely fur ro wed , otherwise duplicating in colony
characte ristics a nd det ails 01' m orphology, th ose gro wn upon CA ; reverse
(colou r plat e 24, b) in yellow-orange shades; sim ila rly coloured pigment
d iffusing into the surro und ing agar; penicilIi as desc ribe d aboye .
Colonies upon M EA (colour plate 24, C, a nd e) , attaining a di ameter 01'
25- 45 mm in 14 days at room temper ature (20-2 5 OC ) , plane , cons picuo us ly
zona te, with ce ntre umbon at e ; heavily sporing th rough out in bluish- gr ay
shades (Sguy, PI. XXXIV , 508); exudate a nd odo ur lacki n g; colony reverse
(colour plat e 24, c) in bright yelIow sha des; peni cilli as describcd aboye, with
conid ial chains formin g loose colum ns , with a tenden cy to form crus ts in old
colon ies; thin m argin surrou nded by a frin ge 01' su bme rge d m ycelia extend ing
abo ut 5 mm beyond th e aerial growth.
Sp ecies description is cente re d upon th e typ e strain CBS 232.38 = IJF M 5013,
strain C BS 337 .48 = IMI 40,5 78 = ATCC 10,453 = NRRL 2,054 = I Fa 6,098 = Q M
3,2 71, and numerous othe r strains possessin g esse ntially th e sa me cult ural and
m orphological characteristics .
The aboye description is drawn in broad eno ug h terms to incl ude a gro u p 01'
stra ins di ffering som ewhat in particul ar characteristics , but showing sufficient
sim ila rity in ge neral appearanc e and det ail s 01' m orphology to be cons ide red
toge the r. The species is com mo n in soil and cosmo polita n in distribution .
T ypi cal strains 01' P. implicatum show con idial areas in deep blue-g reen
colours upon alI su bstrata, with th e blue element pred om in ant. T hey
characteristically appear as rest ricted , hcavily spo ring, with reverse in ora nge brown to m aro on shades . H owever, occasional strai ns, othe rwise typical,
develop yelIow-gree n rather th an dee p blue-gr een shades in con id ial areas.
Sinc e th e colou r change fro m strain to strain is pro gressive a nd no satisfac to ry
line 01' separation ca n be drawn , recogn ition 01' a variet y covering th ese forms is
not con sidere d adv isa ble. It wilI be sufficiently to re cog nize su ch varia tio ns .

25 . P enicillium sublateritium Biou rge : fig. 25, colour plat e 25, and SEM

plate 5 , (top : left) .

...

0 '

00 O
0

iJ..w.WJ.J.J

10 ).lm

O O

sS OOOOO

00 0 0 0 O
O
O O,

lrl~
~

100 jrrn

(1{.

Fig. 25. Camera lucida

drawings of Penicillium
sublateritium Biourge , CBS
267.29. A, Detailed draw ings
of the penicilli. B, Mature
conidia. C, Habit sketches .

Penicillium sublateritium Biourge - Monographie, La Cellule. 33: 315-317. 1!:l23.

Ch. Thom, 'The Penicillia', p. 222.1930; also Raper and Thom , 'A Manual of the
Penicillia' , pp . 203-205. 1949.

Colonies upon CA (colour plate 25, A and a) growing very restrictedly ,

att aining a d iameter of 10-20 mm in 14 da ys at room temperature
(20- 25 OC ), ve lvety , 2 mm deep , convex , with centre buckled, con sisting of
slcnder hyphae developing a close-textu red felt, firm but splitting rather easi ly;
ma rgin abrupt and undulating; sporing poorly in blu c-green shades (Sguy,
PI. XXVII I , 419); odour and exudate lacking; reverse in pale to dark orangered approach ing bri ck red shades (Sguy , PI. X III , 181) in recent isolates ;
pen icilli strictly simple (fig. 25, A), with short stipes, mostly less than 100 fJ- m
in length by 2.0 fJ-m in diamet er , wit h walls smooth or nearly so ; phial ides in
parallel clusters of 5- 10 in the verticil, mos tly 10.0-1 2.0 fJ-m by 2.0 -3.5 tun,
occasiona lly 15 fJ- m long, terminating in a rather broad neck; conidia at first
long elliptical, pyriform to apiculat e , often remaining so, more com m only be-

com in g ellip ticul, with wa lls smooth or finely roughencd (Iig. 25 , B, a nd SEM
pla te 5, top: lcft ); eonid ia l cha ins up lo 100 {u n lon g, beeom in g la nglcd in age
(fig . 25 , C). Conid ial arcas showing gray shades in a ge.
Coloni es upon CYA (colo ur plate 25 , B and b ) growi ng more rapidly ,
attaining a di amet er of 35 mm in 14 da ys a t room temperature (20-25 OC ),
consisting of a c1ose-textured basal felt, velv ety in appearance, deeply rad iately
wr inkled, buckled in ce ntre in a crateriform pattern; heavily spo ring
throughout in blue-green shades (Sguy, PI. XXVIII , 412); und ula tin g thi n
margi n, white; colony rev erse (colour plate 25 , b) in cin namon shades; odou r
and exuda te lackin g; a m ber-colou red pigment diffusing into the surrounding
agar ; peni cilli as described aboye.
Colonies u pon MEA (colour plat e 25, C and e) attain ing a d iameter of
45 mm in 14 days at room temperature, loose-textured , velvety, plane , slight ly
ra ised in centre; margin evenly whi te; heavily sporing in deep artemisia gr ee n
shades (Sg uy, PI. XXXI, 462); odour arid exuda te lackin g; colony reverse
(colour plate 25 , c) in pale yellow to cinna mon shades; pale amber- coloured
pigment di ffu sing into th e surrounding agar; penicilli as described aboye, very
small, wit h short stipes a rising mostly from trailing hyph ae ; conid ial chains
loosely parallel to ta ngled .
Spe cies description is ce ntere d upon th e typ e strain C BS 267.29 = IM I
40 ,59 4 = ATCC 10 ,502 = NRRL 2,07 1 = IFO 6, 107 = QM 7652 = IJFM 5047 , isola ted
by Biourge in 1923.
This stra in appears to suffer actually from a vitamin deficien cy sinc e it gro ws
very restrictedly , sporing poorl y upon CA bu t not upo n CYA.

26 . P enicillium aeneum S m it h: fig. 26 , colour plate 26, a nd SEM p iate 5 ,

(top : r ight) .
Penicillium aeneum Smith - Trans. Brit. Mycol. Soco 46: 334. 1963. See also M.M.
Kulik , 'A Compilation of Descriptions of New Penicillium Species'. Agric . Hand book No. 351. Agric. Res. Ser. U.S. Dep. Agric. p. 13. 1968.
Synonym
Penicillium citreo-viride Biourge var o aeneum Abe - J. Gen. Appl. Microbiol.
Tokyo . 2: 58. 1956.

Colon ies upon CA (colour plat e 26, A and a ) growing slowly to moderately ,
attaining a diamet er varying from 22 to 40 mm in 14 days at room temperature
(20-2 5 OC ) , appearing lea thery , plane , slightly raised in ce ntral ar ea ; th in ,
fimbriate m a rgin, surrou nded by a fringe of submerged hyphae exte nd ing
about 2 mm beyond th e aerial growth ; sporing in blu ish gray-green shades
when isola ted , but very poorly sporing actually , espec ially in marginal a rc a s,

l.l.w.LuJ.LI
}1m

10

. ~!.

fR.,
( '~

\.rv )i.'1'~'.".<. r~; ~1J.)

x; }

l~m

Fig. 26. Camera lucida

drawings of Penicil/ium
aeneum Smith, CBS 321.59.
A, Detailed drawings of the
penicilli. B, Mature con idia.
C, Habit sketches.

rc main in g persi stently white or yellowish cream shades; exu da te a nd odo ur

lacki ng ; peni cilli strictly sim ple (fig. 26, A) with slende r stipes, only 1.5 .tm in
c1 ia meter , smooth wa lled, with apices sligh tly enla rge d u p to 2.5 .tm in
c1 iameter ; phialides in loose to com pact clu sters, nu mb er in g 5 - 12 in the
vcr ticil, m ostly 6. 0 -7 .0 .tm by 2.0-2.5 tun , terminatin g in a very sho rt neck ;
ro nid ia com paratively small, m ostly globose , con spicuo us ly roughe ne d (fig.
~6 , B, and SE M plat e 5, top: ri ght) , 1. 5- 2.5 ust: in di ameter ; co nid ial chains
in loosely pa rallel colu mns, up to 100 .tm lon g , to tangled (fig. 26 , C); colony
rcverse (colou r piate 26, a) in pale yellow shades; pale yellow-co loured pigm ent
d iffusing into th e su rround ing agar.
Smi th' s original description 01' th e species is give n below since th is
adc quately presents the spec ies in q ues tion wh en first isolated. The type strain
C BS 32 1.59, m aintained in artificial cu ltu re since its isolat ion , has become
pred ominantly sterile and leathery , not sho wing a ny longer th e typi cal cult ural
charac teristics originally described by the a utho r, altho ugh still showing
cha racteristic det ail s 01' m orphology .
Sm ith ' s di agn osis as follows:

' Colonies on Czapek agar growing slowly, very bluish gray-green, with a narrow,
slightly yellowish, fimbriate margin, velvety but somewhat heaped in the centre;
reverse dull yellow or developing a greenish east whieh might be termed bronze.'

Colon ies upon C YA (co lou r plat e 26, B a nd b) a tta in ing a diame ter of
35 mm in 14 days a t room temperature (20- 25 OC), velvety to leather y ,
deepl y fur ro wed in a radiat e pattern , buckled in the ce ntre; undulat in g
m argin , su rro unded by a narrow fringe of su bmerged hyph ae extend ing abo ut
1 m m beyond th e ae rial gro wth; poorl y spo ri ng, rc main ing persisten tly white
to pale yellow ish shade s in colou r; exudate abu ndant to limited in amo u nt, as
light am be r d roplets ; colony reverse (colou r pla te 26, b) in cinnamon shades;
simi la rly coloured pigm en t diffusin g in to th e surround ing agar; pe nici lli as
describe d aboye, with stipes short, arising fro m trail in g ae rial hyphae .
Colon ies upon M EA (colour plate 26, C a nd c) a ttaining a d iame ter of
50 mm in 14 days at room te m pe rature (20-25 OC ) , typi cally velvety, plane or
nearl y so, w ith centre slig htly raised , with tufts of whitc hyphae cove ring the
colony su rface in sector s, lendin g it a gr anular appearance; thin , irregul arl y
d issected m argin with a wide frin ge of su bmerged hyphae ex tending abo ut
4 m m beyond th e aerial growth; heavily sporing th rou gh ou t in blue-gr ay
shades (Sg uy, PI. XXXIII , 488); ex udate lacking ; odo ur lacking or
indefinite ; penicilli as described aboye, but with stipes mostly arising from the
su bs tratu m, a nd with con id ial chains loosely parallel. No coloured pi gm ent
diffusin g into th e surrou nd ing agar; colony reverse (colour plat e 26, c) in
cream shades .
Species description is centered upon th e typ e strain CBS 321.59 = CM [
68,225 = ATCC 18,308 = [F O 6,224 = Q M 7,290 , isolat ed by S. Ab e fro m J apanese
so il in 1956. The species is also represented by Ab e ' s stra ins FAT 123 a nd FAT
124, isolated from th e same source.

27 . Penicillium ramusculum Batista el M aia: fig. 27, colou r plat e 27 , and

SEM plate 5 (centre : left) .
Penici//ium ramuscu/um Batista et Maia - Ann. Soco Biol. Pernambuco. 13:24. See
also M.M. Kulik, 'A Compilation of Descriptions of New Penici//ium Species'.
Agre. Handbook No. 351. Agrie. Res. Ser. U.S. Dep. Agrie., p. 69. 1968.

Colonies upon C A (colour piate 27, A and a) growing rather restrictedl y ,

att aining a diameter of 20 mm in 14 days a t room te m pe ra ture (20-25 OC ),
white , velvety, 2.5 mm deep , umbonate , not furro wed, azo nate, cons isting of
a close-textu re d bas al felt covere d by a network of wh ite ropes of hyphac ; very
poorl y spo ri ng actually , but wh en first isolated , sho wing bright olive-gree n
colours in spo ru la ting areas; ex uda te and odo ur lacking; colony reverse (colou r
plat e 27, a) colourless; no coloured pigment diffusin g int o th e surrounding

11

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Fig. 27. Camera lucida

drawings of Penicillium
ramusculum Batista et Maia,
CBS 251.56. A, Detailed
drawings of the penicilli. B,
Mature conidia. C, Ropes of
hyphae . D, Habit sketches.

I.(ar ; penicilli strictIy simple, arising from aeri al hyphae and ropes of hyphae
(lig . 27 , A) , with the stip es of th e conidiophores smooth walled , som etimes
Hc' platc , short , 15-2 5 tut: in length by 1. 5-2 .5 flm in diamet er, slightIy
1;lIlar gcd at the ap ex up to 3 .0 flm in di amet er; phialides in cro wde d clusters,
IIp lo 12 in th e vertici l, parallel or divergent , m ostIy 8. 0 - 12. 0 flm by
" . () - :~ . O tui , terminating in a long, tapering, narrowed neck ; conid ia pyriform
lo cllip tical with walls sm ooth or fine1y roughen ed (fig. 27 , R, and SEM plat e
,i , ce n tre: left) , 2.2 - 2 .8 flm by 1.0-1. 6 flm ; conid ial chains forming short ,
loosc!y parallcl to definite1y com pact colu m ns, bec oming tangled in age (fig.
0'7 , D).
Colonies upon C YA a tta ining a di amet er of 25- 30 mm (colour plat e 27 , R)
iu 14 days at room temperature (20- 25 OC ), similar in texture and colony
uppca ra nce to th ose grown upon CA , but appe a ring m or e gr anular ; very
poorl y sporing, rem aining persistentIy white to white ros y ; exudate and odour
lackin g; colony reverse (co lou r plat e 27 , b) in cin namo n shades ; pale a m be rcoloure d pigment diffusing into th e surrounding a ga r ; penicilli as des cribed
ahoye .

Oolonics upon M EA (colour plutc '2 7, C ami e) nu nining a cli.uuc tc r 0('

:; 0 111111 in 11 da ys al room icmpcrutu rc ('2 0 -'25 OC) , co nsisting 01' a closetexturcd basal kit , pcr sistcntl y whit e and a lmos t stc rilc ac tua lly, bu t qui te
za nate, ash gree n-colou rcd whe n lir st isolat cd (Bati sta and M ai a 1955). Even
m argin su rrou nde d by a frin gc 01' su b me rge d h yph ae cxtc nd ing a bo u t 2 mm
beyond th e ae r ial gro w th; exudate a nd odo u r lacking; colo ny su rface cove red
by interl acing rapes of hyphae , lend in g a gra n ula r a ppeara nce to th e colony;
conid ia l areas sparsely produced , es pec ially in ce n tra l areas, in pale blue-gr een
shades (Sguy, PI. XXX , 440); colony reverse (colour plat e 27 , e) colou rles s.
Penicilli as described a boye .
Sp ecies description is ce n te re d upon the type strain CES 251. 56 = C M I
63,546 = ATCC 12,292 = QM 7,0 57 = IJ FM 5050, isolat ed as a cu lture co n tam ina n t
in Pernambuco , Brazil , by A .C. Batis ta in 1955 .
Penicill ium ramusculum Ba tista et M ai a , wh ose correc t placement still remains
questi onabl e , has been tentati vely assigne d to th e P. imp licatum series, m ainly
on accou n t of its restrict ed growth on m ost m edia , but th e p resen ce of fu rrieles
of vegeta tive hyphae cove ring th e colo ny su rface wauld place it a mo ng th e
species included with in th e P. adametzii se ries .

28. Penicillium palmensis Ramrez , Martnez el F errer: fig. 28 , co lo u r

pl ate 28 , a n d SEM pl a te 5, (centre : ri ght) .

Penicillium palmensis Ramrez, Martnez et Ferrer - Mycopathologia. 66: 77-82.

1978.
C olonies upon C A (co lou r plat e 28, A an d a) gro wing rather restri ctedl y ,
a tta in ing a di amet er of 30 mm in 14 days at roo m temperature (20- 25 OC),
velve ty in a ppearance , with a thin , irregularl y di ssected m argin, frorn th e
uneven gro wth ol' th e veget ative m ycelium largely submerged and brown
colou re d, ex te nd ing 3- 4 mm beyond th e ae rial grow th; m arginal are a pl ane ,
ce n tra l a rea cove re d by an ove rgrowth of ae rial h yphae , a rra nge d in tu fts
lending a granular a p pearance to th e colony su rface; heavily spo ring,
espec ia lly in m argin al area in blue-gr een shades (Sguy, PI. XXXIII , 481 ,
483); od our faint , aro m atic ; ex udate as yellow dropl ets ; colony reverse (colo u r
pIate 28, a) in redd ish b rown shades; pale brown-coloured pi gm ent diffu sin g
into th e surrounding agar; penicilli strictly simple (fig . 28 , A) , with stipes
arising primarily frorn th e substratum , frorn su b me rged hyphae , or fro rn th e
basal felt , m ostly under 75 ttm in len gth by 3. 0- 4.5 ttm in di ameter, with
a pices enla rge d in a vesicle up to 6 ttm in diamet er, with wall s co ns picuo us ly
coa rsely rough en ed ; phialides in cro wde d clusters, mo stly 6 - 12 in the ver ticil,
bottl e shape d, enlarged at th e ba se , th en ta pe ring ab ru p tly at th e a pex in a
sho rt neck , com mo nly 10.0-14 ttm by 4 .0- 5.0 tui , di vergin g; conid ia

"

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Fig. 28. Camera lucida

drawings of Penicillium
palmensis Ramrez, Martnez
et Ferrer. ATCC 38,669. A,
Detailed drawings of the
penicilli. B, Mature conidia.
C, Habit sketches .

r-ll iptical whe n for me d and usu all y remaining so, altho ugh in sorne strains
bccorning ovate, 4.0 -4 .5 /lm by 3.0-3 .5 usi, cons picuo us ly rou gh ened with
thc ru gosity di spl ayed in an helicoidal pattern (fig. 28, B), appearing
n :rebriform whe n obse rve d under th e SE M (pla te 5, centre: ri gh t) ; conid ial
chains form ing lon g , well defined compact colu mns , up to 300 /l m lon g (fig .

28, C).
Colonies upon C YA (colour plate 28, B and b) spreadi ng broadly, attaining
a d iame ter of 48-50 mm in 14 days at room temperat ure (20-25 OC),
1ypically velve ty, radiately and deeply furrowed , wit h centre irregul arl y fold ed ;
(hin, eve nly m a rgin , with su bmerge d hyphae exte nd ing abo ut 1 m m beyond
(he ae rial gro wt h ; heavily sporing throu ghout in deep blue- green shades, wi th
the bl ue pre dominant (Sguy, PI. XXX, 448), especially in marg inal areas;
submarginal and central areas cove red by an ove rgrowth of white aerial
hyphae influen cing the colony appearance by obscuring the otherwise
abundant con id ial structures; odour lackin g ; exudate limited in amo u nt , in
pa le yellow droplets; yellow -coloured pigm ent d iffus ing int o the surrou nd ing
agar; colo ny reverse (colour pla te 28, b) in shades of ochre; peni cilli as

dcscribcd aboye, with conidial chains forming loosely parallcl columna.

Colonics up on M EA (co lour plat c 2B, e a nd c) uua in ing a d ia mcter 01'
55 mm in 14 day s a l room tc m pe ra turc (20 - 25 OC ), co ns isring' o' a loosetextured basal felt , zonat e , typ icall y vc1vcty, pla nc, with m argin thin ,
irregul arly di ssected , su rrou nded by a frin ge of su bmerge d hyphae exte nd ing
abo ut 1- 2 mm beyond th e ae rial gro wth; heavily sporin g in blue-green shades
(Sgu y , PI. XXX , 441), with conid ial struc ture s tending to form cr usts in
cen tral areas; odo ur ' mo uldy'; exudate not obse rve d; no coloure d pi gm ent
di ffusin g in to th e surrounding agar; colony reverse (colour pla te 28, c)
colou rless; penicilli as descri bed aboye, with stipes m ostly arising d irec tly fro m
th e su bs tratum, a nd conidial chains forming de finite ly com pact colu m ns .
T he species diagnos is is centered upon the typ e strain IJ F M 3840 = ATCC
38,669, isolat ed from th e air in L as Palmas de G ran Canaria, Canary Islands,
Sp a in , by DI' R. Ruano-Travieso . It is als o re presented by strains IJFM 3646,
IJ F M 3747, a nd IJ F M 3847, all isolated from the sa me sou rce.

29. Penicillium hispanicum R amrez , Ferrer el M artnez : fig. 29, colou r

plat e 29 , and SE M plat e 5 . (bo tto m: left).

c' .

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Y1 (JJ1V
'-------'

100.um

.J{.

Fig. 29. Camera lucida

drawings of Penicillium
hispanicum Ramrez, Ferrer
et Martnez, ATee 38,667.
A, Detailed drawings of the
penicill i. B, Mature eonidia .
e, Habit sketehes . D,
Submerged vegetative
hyphae with eonidial
struetures arising from it.

Penicillium hispenlcum Ramlrez, Ferrer et Martlnez - Mycopathologia. 66: 7782.

1978.
Colon ies upon CA (colour plat e 29, A and a) gro wing ve ry slowly a nd
restrictedly , attainin g a d iameter of 8 - 10 mm in 14 d a ys a t ro om temperature
(20- 25 OC ), consistin g 01' a tough, close-textu re d basal felt, with a n irregul arly
di ssectcd m argin , su r ro u nde d b y a fringe of stolon -like su b merge d hyphae ,
with colony surface irregularly rais ed and buckled; h eavily spor in g throughout
in sh ades of de ep green (Sguy, PI. XXVII , 401); exu date a n d od our lacking ;
yellow-orange-co loured pigment diffusing into th e su rro u nd ing agar ; colony
reverse (colou r pIa te 29 , a) showing the deep green shades of th e ob ve r se;
penicilli str ictIy sim ple, wi th a n occasion al branch (Fig . 29, A) , with th e stipes
ol' th e con id iop hores a r isin g primarily in a d ense sta n d frorn th e bas al felt, or
occ as ionally frorn trailing h yph ae , up to 100 tm in len gth by 2. 0-2.5 tm in
d ia me te r, with a p ices enla rge d in a vesicl e up to 7 tm in di amet er , with walls
smooth ; phialides cro wde d in parallel c1usters , numberin g 4 to 10 in th e
verticil , ranging from 10.0 to 15.0 tm by 3. 0 -5 .0 tuu , a br u p tIy tapering in a
sho rt neck; con id ia globose to su b globose , 3. 0- 4.0 tm in di amet er , appcaring
smooth wall ed or nearly so (fig . 29, B , a nd SE M plate 5, bottom : left) ; con id ial
chains in fairl y lon g , well-d efined colu m ns up to 200 tm or more in length (fig .

29, C) .
Colon ies upon CYA (co lou r pl at e 29, B a n d b) gro wing a littI e less
re strictedly than upon CA , a tta in in g a di ameter of 23-25 mm in 14 days at
room temperature (20- 25 OC ), co nsisti ng of a close -tex tu re d basal felt ,
velve ty, radiatcly furro wed and rais ed in th e colony ce n tre , sho wing a
depression in a crater ifor rn pattern ; h eav ily spo r ing throu gh out in th e sa me
dcep green sh ades as upon CA , but showing zo nes in yellow shade s; od our a nd
ex udatc lacking ; dark amber to orange-coloured pi gment diffusing into th e
su r rou n d in g a ga r, becoming d ark brown with a ge; colon y reverse (colour pl at e
29, b) in d a rk brown shades; penicilli as described above.
Colon ies upon MEA (co lour pl at e 29, C a n d e) a ttain ing a di amet cr of
26 mm in 14 d a ys at ro om te m pe rature (20- 25 OC), velve ty, zo nate, pl an e ,
with white , thin m a rgin surro u nde d by a fringe of su b merge d hyphae
exte n d ing about2 mm beyond th e ae r ial grow th; heavily sporing throu ghout
in deep blue-green sh ades, with th e blue predominating (Sguy, PI. XXX ,
447) ; ex u date a nd od our lacking; yellow-o ra n ge -colou re d pi gment diffu sin g
into th e su r rau n d ing aga r ; colony rev erse in yellow- orange shades (co lou r plate
29, e); penicilli as described aboye.
The sp ecies di agnosis is ce n te re d upon th e typ e strain IJFM 3223 = ATCC
38, 667=CBS 691.77 , isolat ed b y M .S. Ferrer from th e su rface of a Citrus
limonum L. fruit in M adrid , Spain , in 1975. It is a t present kn own on ly frorn
th e typ e species .

:W. Penicillium ardcsiacutn Novobra nova : Iig. :W, colo ur plat o 30 , and
SE M plat o 5 , (bottom: right).

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L--..l

100 urn

f./? .

Fig. 30. Camera lucida

drawings of Penicillium
ardesiacum Novobranova,
CSS 497.73. A, Detailed
drawings of the penicilli. S,
Mature conidia. C, Habit
sketches.

Penicitlium ardesiacum Novobranova - Nov . Syst. Niz . Rast. 11: 225-234. 1974.

Novobranova ' s diagnosis follow s :

' Colonies upon Czapek 's agar attaining a diameter of 20-30 mm in 10-14 days;
heavily sporing in margin, in scarlet-fire shades, poorly in central areas in yellowish
gray shades; growing margin in yellow-gray shades 10-15 mm wide , undulating;
exudate lacking; colony reverse folded in central zones, in brownish yellow shades,
becoming dirty in age, and in marginal zones in yellow-green shades; sulphuryellow-coloured pigment diffusing into the surrounding agar. Conidial chains
tangled or sometimes in compact columns. Conidiophores arising as short branches
from aerial hyphae, with walls smooth, 50-100 Jl-m by 3.0 -3.5 Jl- m , with apices
enlarged in a vesicle up to 4.0-5.0 Jl-m in diameter. Penicilli str ictly monoverticillate,
with phialides acuminate, forming crowded c1usters of 7-12 in the verticil,
7.0-11 .0Jl-m by 3.5-4.0 Jl- m . Con idia globose to subglobose, delicately roughened ,

.3- 4.5 ,tm in dlameter. It is asslgned with the P. imp/icotum series. No related
pecies are known .
Type: The type strain N. 421 has been isolated from branches of Vitis vinifera L. in
ctober 1967 by T.1. Novobranova, in Kazachstan, Alma-Ata region, USSR.
(Translated from the Russian by Prof. L. Vilasl,
O ur notes follow :
C olonies upon C A (colour plate 30, A and a) gro wing rather restricted ly ,
a ua in ing a diamet er of 35 mm in 14 da ys at room temperature (20-2 5 OC),
vclvcty , with flat , thin margin, su rro u nded by a frin ge of su bme rge d hyphae
cxtcnd ing 3- 4 mm beyond th e ae rial growth; su bce ntra l and ce ntra l arca s
with shallow furrows in a radial pattern, with centre raised ; heavil y sporing
1hrou ghout in deep blue-gr een shades (Sg uy , PI. XXX, 446), espec ially in
marginal zones; showing an overgrowth ofwhite to pale N apl e yellow (Sgu y ,
1'1. XVIII , 270) ae rial hyphae obs curing th e otherwise abundantly produced
ro nid ial structu re s in central a nd su bce ntral arca s; odo u r and exudate lackin g ;
ro lony reverse in deep cad m iu m yellow shades (colour plat e 30, a); bright
yellow-co lou re d pi gm ent diffusing into th e surrou nd in g aga r ; penicilli strictly
sim ple (fig. 30, A), with stipe s of th e conid iopho res arising directly from th e
substra turn, with wall s deli cately rou ghened to coarsely granular, with apices
nlarged in a ve sicle up to 6 J-m in diamet er, u sually sho rt , less th an 75 J-m in
len gth by 3. 0- 4.5 J-m in diameter ; phialides com p aratively large , 10-14
crowde d into the verticil, 10-1 3 J-m by 3.5 - 4 .5 utu , terminating in a
narrowed short neck ; conid ia glob ose to su bglobose, cons picuo u sly spinu lose
(lig. 30, B, and SEM piate 5, bottom: right) , m ostl y 3.5- 4.0 J-m in diamet er;
conid ial ch ain s in definitely com pac t colu m ns (fig . 30, C ).
Colon ies upon CYA (colour piate 30, B and b) attaining a diameter of
40 mm in 14 days a t room temper ature (20- 25 OC) , velve ty, cons picuo us ly
zona te , deepIy sulca te radi ately with few wrink1es , with ce ntre irregul arl y
fold ed and slightly rais ed ; thin a nd undulating m argin with a frin ge of
su bmerg ed my celi a exte nd ing abo u t 1 mm beyond th e aeri al growth ; heavily
sporing throu ghout in deep blue-green (Sguy, PI. XXX , 448) to dark brown
shades, but con id ial fru ctifi cations ob scured in zone s by an ove rgrowth of
white ae rial hyphae , lending a su bla nate appe ara nce to th e colony, sho wing
light salmon-pink shad es in age ; od our faint , 'mouldy' ; exudate lacking or
limited in amount in sulph ur yellow droplets ; colony revers e (co lour plate 30,
b) in orange shades with a green cas t sho wing th rough; su lph ur yellowcoloure d pigment diffusin g into th e surro u nd ing a ga r; pen icilli as de scribed
aboye .
C olonies upon MEA (colour piate 30, C and c) atta ining a di ameter of
50 mm in 14 days at room temperature (20-25 OC) , velv et y , conspicuously
zonat e , showing some what floccose tufts of aeri al h yphae in central areas; thin
m argin, with a fringe ol' submerged h yphae exte nd ing up to 10 mm beyond th e

acria l grow th; hcavil y spo ring in ve ry dc c p blu c-grecn , nlmos t bl nck shndcs
(S gu y , PI. XXX , 446) in m argin al a m i su hcc n tmi arcas, a nd in a rtc m isia
gree n shades (Sguy, PI. XXIX , 425) in ce ntra l a rcas; ex udatc a nd odou r
lack ing; colony rever se (co lou r plat e 30, e) in o ra nge sha dcs; no colou red
pigme nt di ffusin g in to th e su rraund ing agar ; pen icilli as described aboye .
The species di agn osis is cente rcd up on th e type st rain CB S 497 .7 :3 = VKM
F-1 ,7 49 = C M I 174 ,71 9 = A T C C 24, 719 = IJ FM 7070 , isolat ed from vine -shoo ts of
Vitis vinifera L. in Kaza ch stan , Alm a-Ata region , U SS R , by T. I. Novobranova , in 1967 .

3 1. Penicillium gallaicum R amrez , M artnez

pl ate 6 , (to p: left) .

el

Berenguer: fig . 3 1, co lo u r

u..w.u.w
10jJm

O O

0 0 0 000 O
00 00
00
00 8 0 0 0

00 O O
O O

f'

tV, (J! . e

i-,;r " ~' \

~:L1: V ~
L.--...J

100jJm

iR

Fig. 31. Camera lucida

drawings of PenicJllium
gallaicum Ramrez, Martnez
et Berenguer, ATCC 42,232.
A , Detailed drawings of the
penicilii. B, Mature con idia.
C, Habit sketches.

Penicillium gallaicum Ramrez, Martnez et Berenguer - Mycopathologia. 72: 27-34.

1980.

Colonies upon C A (colour plate 3 1, A and a) growing ra th er rest rictedly,

atta inin g a di ame ter of 30 mm in 14 days at room temperature (20-25 OC ),

co nsisting 01' a tou gh , den se , closc-tc x tu red basal felt , with m argin al arca planc,
ce n tra l ami s u bce n tra l arca s ru iscd a m i irrcgul url y Iold cd ; whit c , c n tirc , thi n
m a rgin surround ed by a Iringc 01' su b ruc rgc d hyph a c ex (cnd ing a bo u t 2 mm
beyond th e a er ia l gro w th ; co lo ny su r lace a p pcar ing vel vety, with ce ntre
g-ran u la r , m ainly cluc lo a th in ovc rg rowth of w hite ae r ia l hypha e ; poorly
spo r ing in pal e brow n shade s in m argin al zo nes a nd in ye llowish colou rs in
ce n tra l an cl su bce n tral arca s ; ex ucla te a n cl odo u r la cking ; reverse (colour pl ate
:)1, a) in pal e o ra nge shades in ce n tre ancl colou rl ess in m a rgin o Penicilli
com rno nly str icty sim ple, seldom wi th a n occas ional branch which bea rs a
verticil of phialides re tain ing a lways its m onoverticillate character; th e stipes of
th e co n icliophores short , u su all y less th an 50 tm long by 2.0- 3.0 tm in
d ia mete r (fig. 31, A) , wi th wall s smooth, hyal in e, a r ising e ither from th e
su b stratu m 0 1' a s short branch es fro m aeria l trailin g h yphae , with a p ices
enla rgecl in a ves icle up to 5 tm in wiclth ; phialides closely pa ck ed in pa rall el
clu sters , m ostl y 6 - 10 per verticil , ranging fro m 7.0 to 12 tm in length by
2.5 -4 .0 tm in cliamet er , with short n arrow n eck s ; con iclia glo bose to
su bglobose, smooth-wall ecl 0 1' nearl y so when obse rvecl uncl er th e light
mi cro scope , but deli cately ro ughene d when obse rve cl u n cler th e SE M (fig . 31,
B, a ncl SEM pl ate 6, top : left) , measuring 2.0 -3 .0 tm in di amet er ; co ni clial
ch a in s forrning loose colu m ns 0 1' tanglecl, up to 150 tm long (fig . 31, C) .
Colo n ies upon C Y A (co lo ur pl a te 31, B a n d b ) growing a little m ore rapicll y
th an upon CA, a ttain ing a di ame ter of 35 mm in 14 clays a t roo m temperature
(20- 25 OC ) , tou gh , velvety, close textu recl, zo nate, cleeply ra di atel y wrinkled
with numerous furrow s , with ce nt re ra isecl a nd irregul arly folclecl in a
cra ter ifo rrn p attern ; eve n m argin su rro u n de d b y a frin ge of su b mergecl hypha e
ex te n di ng a bou t 2 mm beyoncl th e ae ria l growth ; h eavil y spo ri ng za nes in
blue- green shacles (Sguy, PI. XXX , 436), poo rly spor ing zo nes showing
su lph ur yello w co lours (Sg uy , PI. XX , 286) , m ainl y clue to th e ye llow
enc ru stecl hypha e ; ex uclate in small, yello w clrops ; oclour la cking ; colony
reverse (co lo ur pl at e 31, b) in ye llow-o range sh acles ; su lphur yello w-co lou re cl
pi gment cliffu sing into th e su rro u ncling agar. Penicilli as clescribed aboye .
Colo nies upon M EA (co lo ur pl ate 31, C a n cl c) a ttain ing a di amet er of
27 mm in 14 cla ys at ro om tempera ture (20 - 25 OC ), co ns isting of a leathe r -like
b a sal felt formecl of yello w enc rus tecl vegeta tive h yph ae, pl ane , with ce ntre
sligh tly rais ecl a n cl clepressecl, za nate ; thin m argin with subm erged yellow
encru stecl h ypha e ex te nd ing 3- 4 m m beyoncl th e ae rial grow th ; h eavil y
sporing in blue-gray shacles (S g uy, PI. XXX , 449); odou r a n d ex u clate
la cking ; colon y reverse (co lo ur pl at e 31, e) in cleep su lph u r ye llow colo u rs ;
sim ilarly co lo urecl pi gment di ffu sing into th e su r ro u ndi ng agar; pe n icilli as
clescribecl a bove .
Penicillium gallaicum a ppears to stancl b etween species incluclecl within the P.
imp licatum se r ies a n cl th e P. decumbens se ries . On accou nt of th e restri ct ecl

g ro w rh o n m ost m edi a it suc ms lo lil m ost sa lisfa <: tlll'ily withi n (he ormc r

ser ies, bu t Oll acco u n t 0 1' ih c sho rt co n id io pho rcs a ris illg" mo stl y fro rn nct wo r ks
01' tr ailing h yphae it could als o be assignablc to th e P. decumbens series.
Species diagnosis is centered on the type st rain I.f F M 5597 = A T C C 42,232 , isola ted
fro m the ai r 01' M ad rid , Sp ain , in 1979 . It is a t prcscnt kn own on ly from the
typ e st rain .

Penicillium decumbens ser ies

OUTSTAN DING FEATU RES

Colonies growing slowly, u su ally restrietedly , su rfaee appear ing ve lve ty 01'
slightly la n a te , eons isting 01' a net work 01' trai ling a nd inter woven vcgeta tive
hyphae , rangi ng fro m com pa ra tively loose textu re d in so rne species to close
textured, tou gh, dense m yceli al felt s in othe rs .
Penieilli str ictly sim ple , co ns istcntly small , usually typi call y borne on short
stipes a r ising frorn trai lin g , 0 1' interwoven ae r ial hyphae , seldom from the
su bs tratum .
Con id ia small , usu ally somewhat ellip tica l, a bo u t 2.0-3. 0 .,m in th e lon g
ax is, smooth wa llcd in m ost spcc ics, delicately raughened in others.

K ey to th e spec ies
2. Colonies appearing velvety or with surface slightly lanate ; stipes borne
prima rily as short branches from loosely trailing or compacted vegeta tive
hyphae.
P. decumbens series
a. Colon ies loose-textured , w ith margin usually thin and gene rally
consisting of a loose network of interlacing hyphae bearing short
stipes .
1' . Conidial areas in Iight gray-green shades w ith reverse in yellow
drab shades upon CA but becoming cherry red upon MEA
P. chermesinum Biourge
2' . Conidial areas in dull blue -green shades with reverse usually
colourless upon all med ia.
P. decumbens Thom
b. Colon ies c1ose-textured, tough , almost leathery, restricted , with
margin compact but showing occasionally stolon-l ike hyphae .
1'. Vegetative mycelium yellow, gene rally characterizing the colony
even in age , sporu lating slightly upon CA.
P. citreo-viride Biourge
2'. Vegetative mycelium white , often characterizing the colony when
yo ung , conidial areas deve loping blue-green shades .

Page

.... 102

. .. . 106

. . . . 105

. . . . 108

aa. Phlalidas comparatlvaly larga, up to 16 Itm in langth, reversa

uncoloured or in Iight vinacaous shades.
P. fellutanum Biourge ....
bb. Phialides comparatively small, up to 10 Itm in length.
1". Colonies growing very restrictedly, less than 20 mm in
diameter.
aaa. Colonies strongly funiculose, conidia elliptical.
P. brevissimum Rai etWadhwani . .. .
bbb. Colonies velvety.
1"'. Conidia elliptical to ovoid, reverse salmon-pink.
P. gerundense Ramrez etal. . . . .
2"'. Conidia globose to subglobose.
aaaa. Conidia smooth-walled.
P. alicantinum Ramrez et al. . . . .
bbbb. Conidia spinulose.
P. malacaense Ramrez et al. . . . .
2". Colonies growing less restrictedly, up to 25-30 mm in d ameter.
aaa. Colonies tomentose, white rosy, reverse in pale pink to
vinaceous shades.
P. kurssanovii Chalabuda . . . .
bbb. Calonies not tomentase, typically velvety .
1'", Calanies in mause gray shades, reverse white rosy,
amber yellow diffusing pigmento
P. cinereo-atrum Chalabuda . ...
2'''. Calanies in yellow shades, brown diffusing pigment.
P. toxicarium Miyake ex Ramrez .. ..
3'. Vegetative mycelium white to pale vinaceaus, poarly sporing,
reverse oranqe, bright orange-red to maroan.
aa. Colanies grawing very restrictedly, attaining up to 20 mm in
diameter upon CA; diffusing pigment in vinaceous shades.
P. roseo-purpureum Dierckx ... .
bb. Colanies growing less restrictedly, attaining 35-40 mm in
diameter upon CA .
1". Colonias conspicuously tamentase, in gray olivaceaus
shades; yellaw diffusing pigment; conidia elliptical, smooth
walled, in tangled chains.
P. glauco-Ianosum Chalabuda ....
2". Calanies not tamentase, typically velvety, in blue-green
shades; brawn diffusing pigment; canidia glabase to subglabase, smooth-walled, in well -defined short calumns.
P. bilaiae Chalabuda ....

111

117

128

129
131

120

122
124

113

115

118

M cmbcrs of P. decumbens seri es are fairly abundant in nutu re ane! worldwidc

in distributi on. Thcy are com mon in soil and r equ cntly isolat cd from milit ary
eq u ipme n t deteriorating under Iield cond itio ns .
The P. decumbens seri es, as it wa s cons ide red by Raper and Thorn (1949) ,
included a range of spe cies rather div ergent in sorne respects but uniformly
chara cte rized by th e sm all penicilli borne on short stipe s arising frorn tr ailin g
aeri al hyphae . They su bd ivide d th e seri es, on th e basi s of gen eral colony
texture, int o two fairly well-defined subseri es: The first one, typified by
Penicillium decumbens Thom , is cha rac te rized by its com pa ratively slow-gro wing
and loose-textured colonies in which th e penicilli-bearing hyphae form a loose
network . In addition to P. decumbens, this sub-seri es included P. chermesinum
Biourge , characterized by its m ore rapid gr owth and th e bright red colours
develop ed in colony rever se upon MEA . The second subseri es , typifi ed by
Penicillium f ellutanum Biourge, is cha rac terized by its restricted , tough, almo st
leather y colonies, frorn th e su rface of whi ch arise trailing peni cilli-bearing
ae rial hyphae , with short stipe s in th e m anner typi cal of th e series. Penicillium
citreo-viride Biourge , a slow- growing, tardily sporing form, producing sm all,
rai sed colon ies with obve rse a nd reverse in bright yellow colou rs upon C A, was
also included , together with Penicillium roseo-purpureum Dierckx , a very slowgrowing species , tardily sporing and alm ost lanate , producin g abundant
or ange-red to vinaceous pigm ent in th e exudatc and th e colony reverse.
Owing to th e large number of new species th at hav e been described in th e
literature since 1949 , closely related, in a broad sense, to th e P. decumbens
seri es, th e following form s have te nta tively been assigne d here : P. brevissimum
R ai et W adhwani , a very slow-gr owing species, stro ngly funi cul ose ; P.
kurssanovii C hala buda , characte rize d by its less re stricted, rather tomentose ,
white rosy colonies, showing pale pink to vinaceous shades in th e colony
reverse ; P. cinereo-atrum Chalabuda , characte rized by its mouse gra y, velvety
colonies, with revers e white rosy and am be r yellow diffu sing pigmen t ; P.
toxicarium Miyake ex Ramrez , cha rac terized by its blue-green colonies a nd the
dark brown diffusing pigm ent ; P. bilaiae Chalabuda , characterized by its less
restricted colonies, velvety , blue-green colou re d, with brown diffu sing
pigm ent; P. glauco-lanosum Chalabuda , character ized by its tomentose , gray
olivaceous colon ies, with yellow diffusin g pigment ; P. gerundense R amrez et
Martnez , ch aracte rized by its very slow-growing, white , buckled colonies
upon CA ; P. alicantinum R amrez et Martnez , characterized by its m od erately
gr owing, velv ety, de ep blue-green colonies, with bright yello w diffusing
pi gment, and th e re vers e in orange shades; a nd P. malacaense R amrez et
Martnez, a very slow-grow ing form, charac te rized by its ve lvety, d ark graygreen colon ies upon MEA .

32. Penicillium decumbens T hom : lig. :12 , co lo ur plat o :12 , a nd SE M plat c

G, ( top : right) ,

0 0 0 0
O o o 0
00 o 0 0

Bo o o
() 0 0 o o
0

w....J.LwJ

10jJm

e
l....----..J

100jJm

fR.,

Fig . 32. Camera luc ida

drawings of Penicillium
decumbens T horn, CSS
318.48. A, Detailed drawings
of the penic illi. S, Mature
con idia. e, Habit sketches.

Penicillium decumbens Thom - Bull. Bur. Anim . Ind. U.S. Dep. Agr ic. 118: 71 .
1910. See also Thom , 'The Penicillia', pp. 197-198. 1930; also Raper and Thom,
'A Manual of the Penicillia' , pp . 209-212. 1949.

Coloni es upon CA (co lour plate 32, A and a) growing slowly, attammg a
diameter of20- 30 mm in 14 days at room temperature (20-2 5 OC), velve ty in
some strains, with a tenden cy to develop a white mycelial overgro wth in
othe rs, and in stil1 othe rs almo st lanat e, deep, u p to 1- 2 mm , bu t al1
cha rac te rized by loosely interwoven a nd tr ailing hyphae from whi ch arise short
stipes, plane, very thin m argin with sub m erged hyphae ext ending largely
beyond th e ae rial growth (abou t 2-4 mm) ; heavily spo ring in grayish gr een to
olivaceous shades (Sg uy , PI. XXI, 307); exudate lackin g or lim ited in
a mount; odo u r distincti ve , fragrant , su ggesting soap perfumes; colon y revers e
(colour plat e 32, a) almost colou rle ss, with a green ca st showing through ; no
coloured pigment diffusin g into th e surro u nd ing agar; peni cil1i strictly sim ple

(fig. :~2, A), with the stip e 01' the conid iophores sho rt , :>0- 100 JJ.1T1 by
2.0 -2.5 JJ.1T1 , with apiccs slightly c nla rged in a vesiclc IIp lo :U ) JJ.IIl in
diarnct cr , with walls smooth 01' deli cat cly rough cn cd , borne al su cccssivc nodos
upon trailing hyphae, which in marginal arcas grow stolo n -like struc tu res
along th e substratum ; phialides usually in compact clusters u p to 12-15 in the
verticil , 7.0-9 .0 IJ-m by 2.0 -2 .5 IJ-m, sometimes borne at two immediately
a djacen t lev els ; co n id ia ellip tical to su b globose , 2.0-2. 5 IJ-m in long axis ,
occ asionally up to 3.0 IJ-m , smooth walled (fig. 32, 13) to delicately roughened
when observed under th e SEM (plate 6, top : right) ; conidial chain s producing
short , loo sel y parallel colu m n s, up to 100 IJ-m long (fig . 32, C).
Colon ie s upon CYA (colour plate 32, B a nd b) a tt a in in g a diamet er of
25- 30 mm in 14 days at room temperature (20-25 OC ), velvety , radiately
fur rowed with ce n tre fold ed in a somewhat crate r ifo rm pattern ; thin margin
with a fringe of su bmerged h yphae exte n d ing less th an 1 mm be yond th e aerial
growth , developing a white m ycelial ov ergrowth in cen tral a nd su bce n tra l
a rcas , othe rwise duplicating the co lon ies grown upon C A in cu ltu ra l charac te risti cs ; ex u date abundantly produced, crystal clear, as colo urless droplets;
odour fra grant a s described a boye; colo n y rev erse (col our plate 32, b) in
yellowish cream shades in m argin al a reas a nd in deep gree n co lou rs sh owing
throu gh in th e ce n tra l arca s. P enicilli as described aboy e.
C olo n ies o n MEA (colour pi a te 32, C a n d c) a ttain in g a di amet er of
35-50 mm in 14 d a ys a t ro om temperature (20- 25 OC ) , typically velve ty ,
pl ane ; heavily spo rin g in dark olive-g re en sh ades (S gu y , PI. XXIX , 433);
thin m argin irre gul arly di ssect ed , surro u n de d by a frin ge of su b merged h ypha e
ex te nd ing a bou t 3- 4 mm beyond th e aerial gro w th ; exu da te a nd odou r
la cking ; co lo ny re verse (colo ur plate 32, e) colo u rless , with a greenish cast
sho wing throu gh ; n o colou re d pigment diffu sin g into th e surro u nd ing agar .
Penicilli as described aboye ; co n id ia l chains forming loos el y p arallel co lu m n s
up to 250 IJ-m long .
Spe cies d escription is cen te red upon strain ATCC 10,436 = C BS 318.48 = NR R L
742 = IF O 6,093 = C M I 39,755 = Q M 1,920 = IJFM 7068, isol ated by Prof.
Sherbakokk , University of Tennessee , Knoxville , T enne ssee , USA, also on
strain C BS 258.33 = N R RL 2,044 = IJFM 5003, which represents th e sp eci es satisfa ctorily but tends to produ ce co lon ies somew ha t de ep er a nd of looser texture
th an th e former strain , a n d a lso upon m any other strain s th at present th e same
general m orpholo gy .
The species a ppear s to be com mon, a nd worldwide in di stribution , a nd h as
b een isolated primarily from so il a nd det eriorating military eq u ip men t in
tropical co u n tr ies .

33 . P enicillium chermesinum Biourge : fig. 33, colour plate 33 , and SEM

plat e 6, (centre: left).

000
O

0 O
00

gO OO

a o o B
O

o o

00

--------------------------

0
o
l..1wl.w!J

~
~ 1O~m
~

e~,,- ,

li~~lt
100

}-1m

Fig. 33. Camera lucida

drawings of Penicillium
chermesinum Biourge, CBS
305.49. A, Detailed drawings
of the penicilli. B, Mature
conidia . e, Habit sketches.

Penicillium chermesinum Biourge - Monographie, La Cellule. 33: 284-285. 1923.

See also Ch. Thom, 'The Penicillia', pp. 192-193. 1930; also Raper and Thom,
'A Manual of the Penicillia', pp . 206-209. 1949.

Colon ies upon C A (colour piate 33, A and a) attaini ng a d iamet er of 40 mm

in 14 days at room temperature (20 - 25 OC ) , radiat ely wrinkled a nd buckled ,
with colony su rface tufted, cons picuo us ly gr anular , cons ist ing of an int erl acin g
netw ork of h yphae a nd rop es of hyphae ; white , thin m argin , wi th a frin ge of
su bme rge d h yphae extend ing abo ut 5 mm beyond th e ae rial gro wth; heavily
sporing th rou gh out in olive green sha des (Sgu y , PI. XXI , 309), with con id ial
are as some what ob scured in places by a m yceli al ove rg rowth of white ae rial
hyphae ; exu date a nd odo u r lacking or faint , suggesting wood smoke; colony
reverse (colour plat e 33, a) in ligh t yellow to flesh colours. Penicilli strictly
sim ple (Fig. 33, A), with the stipes of the conid iopho res borne as sho rt
branch es from loosely interwoven and trailing hyphae , with wall s smoo th,
mo stly 20- 40 /lm in len gth by 2.0-2 .5 /lm in di ameter, with apices enla rge d in

a vcsid c IIp lo 4 .0 -4'.5 un in di um ct er ; phialidcs in corn pact whorl s o ' 10-1 5

in the vcrti cil , mostl y 6- B .UII, ll lo 12 un in leu grh hy 2.0-2 .5 .un in
di amct cr , oft cn incu rve d; co n id ia co tu p a ra t ivc ly sma ll, cllip tica l, sm ooth
walled 01' nearly so wh en obse rved under th e light mi cr oscop e (Iig. :3:3 , B) 01'
deli cately rou gh en ed wh en obse rved under th e S EM (pl at e 6, cen tre: left) ,
mo stl y 2.0-2 .5 11m by 1.5 -2 .0 11m in diamet er, a ppe aring green in mass ;
con id ial chains producin g loosely parallel to fairly co m pac t colu m ns (li g . 33, C).
Colonies upon CYA (colour pl at e 33, B a nd b) attaining a diamet er of
45- 50 mm in 14 d ays at ro om temperature (20-2 5 OC ), velve ty, radi ately
wrinkled with numer ous furrow s, and centre irregularly fold ed a nd rai sed ;
white margin en tire; heavily sp oring throu gh ou t in oliv e gree n shades (S guy,
PI. XXI , 309), but with co nid ia l areas some what ob scured by a m ycelial
ove rgrow th, lending a lanat e ap peara nce to th e colony su rface ; odo u r and
ex udate la cking 0 1' limited , in vinaceous shades; vinaceous -colou re d pi gm ent
diffusin g into th e surro u nd ing agar; colony reverse (colour pl at e 33, b) in
cream to deep vinaceous red sh ades . Penicilli as described aboye .
Colon ies upon MEA (co lour pl at e 33, C and c) attaining a diamet er of
50 mm in 14 d ays at room temperature (20-2 5 OC ) , velve ty, plane , a ppe a ring
slightly gra n u la r; irregul arl y di ssected m argin with a fringe of su b me rge d
hyphae ex te nd ing abo ut 2 mm beyond th e ae ria l gro wth; heavil y spo ring
th rou ghout in arte m isia green shades (Sguy, PI. XXVII , 403) ; ex u date and
odo u r lacking ; colo ny reversc (colour piate 33, c) in cream to d eep wine red
shades; deep red wine-coloured pigment diffu sing into th e su rro u nd ing aga r ;
penicilli as described aboye .
Spec ies description is cen tered upon stra in C BS 305,48 = C M I 40,5 77 = ATCC
10,424 = NR RL 2,049 = QM 5, 120 = IJ FM 5049, isolat ed from th e a ir by K.B .
R ap er , and several add itional strains sho win g similar cult ural characte ristics
and det ail s of m orphology. The species ha s been rep eat edl y isolat ed from
det eri orating fabrics and other material su bject to con tact with th e soil 0 1' to
du st co ntam ination, a nd should probabl y be regarded as esse n tially a soil
orgams m.

34 . P enicillium citreo-viride Biourge : fig . 34 , co lo u r pl at e 34, a n d SEM

pi a te 6, (centre : ri ght).
Penicillium citreo-viride Biourge - Monographie, La Cellule. 33: 299. 1923. See also
Ch. Thom, 'The Penicillia', pp. 199-200. 1930; also Raper and Thom, 'A Manual
of the Penicillia', pp. 215-218. 1949.

Synonymies
Penicillium citreo-nigrum Dierckx - Ann. Soco Sci. Bruxelles. 25: 86. 1901 : see
also Biourge's Monographie, La Cellule. 33: 273-274. 1923.

.r-:

00 O 00
O
000
O

00

00 0 00
B O

000 O
O

lJ.J.J.J..lJ..w

10 }1m
~t

fR

'\

~'q,, ~.. ,(i"'" , ' k'/;",.C, r'~" \\,

i

',"

I'~ l

')' .

'

1 JJ-l
-1

l-.-..J

100 )Jm

Fig. 34. Camera lucida

drawings of Penicillium
citreo-viride Siourge, CSS
258.29. A, Detailed draw ings
of the penici lli. S, Mature
coni dia. e, Habit sketches.

Penici/lium citreo -sulfuratum Biourge

Monographie, la Cellule. 33: 285-287.
1923.
Penici/lium necrosiferum Morotchkovsky - Bul Sci. Rec. Biol. Univ. Kiev. 2: 79.
1936.
Penici/lium tCitromvces) sormanii Carbone - Atti Inst. Bot. Univ. Pavia, Ser. 11 ,
14: 259-325. 1910.
Penici/lium subcinereum Westling - Ark. Bot. 11: 137-139. 1911; also Biourge 's
Monograph ie, La Cellule. 33: 273. 1923.

C olonies on C A (colour pIat e 34, A and a) growing more or less restrictedly ,

a ttain ing a di amet er of 25-30 mm in 14 d a ys at room temper ature '
(20- 25 OC ), co ns isting of a tou gh , close -textu red b asa l felt , deeply radiately
furrowed in m argin al a reas, umbon at e in so rne strains in th e ce ntre, depres sed
in others; velvety or su b floccose , spo r ing tardily in so rne strain s, less tardily in
others, colon y su rface powdery, in white colou r; ex u d ate a nd odour lackin g ;
thin m argin , fib rous , with su b mergc d hyphac ex te n d ing 5-8 mm bcyond th e
ae r ial growth ; colon y reverse in pale ycllo w to cream shades (colour pIa te 34 ,

11 0

a) ; pal c ycllow-co lo u rc d piglllcnl dilTu sin g slowly in to (he surroundin g uga r:

peni cilli str ictly sim ple (lig. :H , A), with stipcs 01' rhc conid iophorcs iypically
a rising from tr ailin g a nd ac rial hyphac , sho rt a nd slc ndc r, mostl y 50- 100
in len gth b y 1.0 - 2.0 ",m in d iamet cr , but so mc timcs a rising directly rom (he
su bst ratum, being th en lon ger, up to 150 tuu , with walls smooth , with api ccs
sligh tly enla rge d ; phial id es in com pact c1usters 01' 8- 1 in th e ve rticil , mostly
7.0 -1 2.0 ",m by 2.2 -2 .8 tut , with co ns picuo us , fairl y lon g, tap ered necks ;
co n id ia globose to su bg lobose, usu all y 2.0-3.0 tut , occasionally up lo
4.0- 5.0 ",m in di amet er, thin wall ed, smo oth or nearl y so (SEM plate 6 ,
ce n tre : right); co n id ia l chains formin g 100se1y parallel to sligh t di vergent
colu m ns (fig. 34, C) .
Colonies u po n C YA (colour plat e 34, B and b) grow ing m or e rapidly th an
up on CA , at ta in ing a di a met er 01' 44 mm in 14 d ays at roo m te m pe ra tu re
(20-25 OC ), ve1vety in a p peara nce, close tex tu re d, deepl y radiat e1y fu rrowcd
with cen tr e umbonat e , sh owing some zo nation ; thin undulat ing margin
su rrou nde d b y a fringe 01' submcrge d hyph ae ex te nd ing abo u t 2 mm beyoncl
the ac ria l grow th; heavily spo ring throu gh out in gray-green shades (Sg uy, PI.
XXI , 309); ex udate lacking; odo ur fa int , indefinite ; colo ny reverse (co lou r
pla te 34, b) in yellow shade s; pal e am be r-co lou re d pi gm ent slowly diffusing
in to th e su rroun d ing agar; peni cill i as describe d aboye .
Colon ies up on MEA (colour pl at e 34, C and e) a ttain ing a d ia me te r 01'
47 mm in 14 d ays a t roo m temperature (20-25 OC), typi call y velvet y to
su bflocco se , almos t pl ane , with some shallow fu rrows; vegetative hyphae
de1ica te and yellow in colour ; unevenl y d issected or fine1y denticul ate m argin
surro u nde d by a frin ge 01' deepl y brown-coloured sub me rge d h yphae exte nd ing
a bou t 5 mm beyond th e ae rial grow th, th e deep b rown colo u r being due to th e
for mation 01' d ark purple-brown , needl e-sh ap ed crys ta ls into th e subs tra tu m,
near th e margin ; spori ng ab u ndan tly throu gh ou t in blue- green shades (Sguy,
PI. XXIX , 425); exud ate la cking; od our faint , aromatic; am be r-coloured
pi gment diffusing into th e su rrou nd ing agar , but partially obscu re d in
m arginal areas by th e purple-brown crystals em be dde d in th e su bs tratu m ;
colo ny reverse (co lour plat e 34, c) in d ark brown shades , showing orange to
ochraceous shades near th e margin o Penicilli as descri bed aboye .
The spe cies d escrip tion is m a inl y cen te re d upon stra in C HS 258 .29
(describe d by Di erckx as P. citreonigrum) (SE M pla te 34), upon strain C BS
308 .48 = NR RL 2,046 = C M I 40,575 = ATCC 10,425 = IJ FM 5030, isolat ed fro m
m ilita ry eq uipmen t by K .B. R aper, and on seve ral strains having similar
cultu ral characte ristics and det ails 01' m orphology . (Colou r plat es 234 a nd 235
pr esent two representative stra ins 01' P. citreo-viride, isolat ed from the a ir and
from pepper , resp ectively , by th e au tho r.)
T he spec ies sho uld be regarded as essent ia lly a soil organism th at is
cos mopo lita n in di stribu tion . In P. citreo-viride, as in other wide ly di stributed

"'Ill

' - .- 1 - -

IY appcarancc, in th e rclativc abundance 01' sporulatlon ami in lesser

ce in thc fi ne r details 01' morphology. T h is is probably thc reaso n why
1

el' work ers may hav e ba sed species descrip tion s upon va riants 01' st rai ns
kn own to re prcscnt differcnt aspects of the sarne spec ies, which is th e case
IC fivc spec ies listed above as syno nyms of P. citreo -viride.

Penicillium Jellutanum Biourge : fig. 35, colour plate 35 , and SEM

late 6, (bottom: left) .

~O . , Cl ~

(~p

O () .:\:::l

00 O O
O

O0 ,':'"':: 0

O .',

. O ()O

.'

00 <
!J.w.lullJ
10 jrrn

, <~,
R
. : .-: '// i
( ;.~
\) W
~\
.
'
~
i\:'
1"
v.., ;'.~'. " .~ . [ 1.1,.'
11

lrl1~"L

100}Jm

Fig. 35. Camera lucida

drawings of Penicillium
fellu tanum Biou rge, CBS
326.48. A, Detailed drawings
of th e penicilli. B, Mat ure
con idia. e, Habit sket ches.

enic lium fe/lutanum Biourge - Monographie, La Cellule. 33: 262-264. 1923. See
also Ch. Thom , 'The Penicillia', pp. 198-199. 1930; also Raper and Thom, ' A
Manual of the Penicillia', pp . 212-215. 1949.

ynonymies
Penici/lium phaeo-janthine/lum Biourge - Monographie, La Cellule. 33: 289-290.
1923.

Ponlcillium dlorckxli Blourg o - Monogrophlo,' La Collulo. 33: 313315. 1923.

Penicillium cinerescens Biourge - Monographle, La Cellule. 33: 30S-309. 1923.
Penicillium fellutanum Biourge, varo nigrocesten eum Abe - J. Gen. Appl.
Microbiol. Tokyo. 2: 71-72. 1956. See also Smith, Trans. Brit. Mycol. Soco 44:
332. 1963; also M.M. Kulik, 'A Compilation of Descriptions of New Penicillium
Species'. Agric . Handbook No. 351. Agric. Res. Ser. U.S. Dep. Agric . p. 40.
1968.

Colon ies upon CA (co lour plate 35, A and a) growing rather res tricted ly,
a ttaining a diameter of 20- 30 mm in 14 days at ro om te mpera tu re
(20- 25 OC ), co ns isting of a tough , close- te xtu red basal fel t, 100- 200 utt: deep ,
wi th m argin ofte n surrou nde d by a na rrow zon e ofpredo m inan tly su bmerged,
fu nicu lose , sto lon-like h yphae , radiately furrowed , with concen trical wrinkles ,
n ar rowly za nate, cave red by a mycelia l overgrowth of snow-white ae rial
h yphae in flue ncing greatly th e colony a ppeara nce; poorl y spo r ing in sorne
st rains, only in m arginal areas, with centre rem aining persistently white,
conid ial a reas in light blue- gr een shades (Sguy, PI. XX , 449); ex udate as
colou rless d ropl ets em be d de d in th e myceIial feIt ; odour not pronounced ,
rath er pleasant; colon y reverse (co lour pl at e 35, a) in pal e cream to flesh
colou rs or in light vinaceous shades , in sorne strains greenish co loured; penicilli
stric tly sim ple, wi th a n occas ional b ranch (fig . 35, A), co m parativeIy large ,
wi th sho r t stipes, usu all y 50- 100 utt: in length by 2.0-2 .5 /l m in diamete r,
with wall s smooth , ar ising from interlacing, trailing or creep ing h yphae ,
co m mo nly enla rge d at th e a pe x in a vesicle up to 4.0- 5. 0 /lm in dia me te r;
ph ialides in compact clusters 8 - 12 in th e ve rt icil, compara tiveIy la rge , m ostly
10 -1 5/lm lon g b y 2.0- 4.0 tut: in di amet er, with fairly long tapered neck ;
co nid ia elliptica l to subglobos e, mos tly a bo ut 2.5 - 3 .O /lm in long axis, smooth
or cons picuo us ly ro ug he ne d (fig. 35, B, a nd SEM pla te 6, botto m : left);
conid ia l chains form ing loosely paralleI or d iverge nt, poorl y defin ed colu m ns
up to 100 /lm in length (fig . 35, C).
Colo nies u pon C YA (colour plate 35, B and b) attaining a diameter of
35 mm in 14 days at room temperature (20 -25 OC ), simi lar in tex ture and
cultu ral cha racte ristics to those grown u pon CA, bu t m ore deep ly ra d iateIy
fu rrowed, with centre raised a nd depressed in a craterifor m pattern, an d very
poorl y sporing, re maining persisten tly white in colou r ; undul ating m argin
su rrou nded by a narrow fri ng e of su bme rg ed hyph ae ex te nding abou t 1 mm
beyond the aeria l growth ; colony su rface con spic uo u siy zanat e ; exudate
a b u ndan tly p roduced in co lou rless dro ps; odour lacking; colony reverse
(colo u r plate 35, b) in cream shades; pal e amber-colou red pigmen t d iffus ing
into the surrou nd ing agar ; penicilli esse ntia lly as described a boye .
Colon ies u po n MEA (colo ur plat e 35, C a nd c) attain ing a d ia meter of
40 mm in 14 days at room temperat u re (20-25 OC ), ty pically veIvety ,

sp ccics 01' Penicillium, individu al strains sho w a p prec inb lc vuriati on in gross
co lo n y a ppea rancc , in th e re lat ivo a bu nd n ncc 01' spo r ulutio n , a m i in lcsser
d cgrcc in the Iincr d etails 01' m orphology . T h is is proba bl y th e rcaso n w hy
ea rlier workers m a y hav e b a sed spe cies d c scripti on s up on va ria nts or str a ins
now kn own to represen : diff erent as pe cts 01' th e sa m c species, wh ich is thc ca se
of th e Ii ve sp ecies listed above as syn on ym s 01' P. citreo -oiride.

35 . Penicillium f ellutanum Biourge : fig. 35 , colour plat e 35 , a nd SE M

pi ate 6 , (bo ttom : left).

0
O ... ()(::)

( ~O a . . .>J
00 o o B o

o 00 ,0
'0 e>.....
'. O 00
0 .0
lJ..t..u..lwJ.J
10 jrrn

"
r;;>-'\

~l

(,~ .

1 1 ;:p

l'
lr1~L '

~ \- \~~1"

,::"''"

(:;;~

' . 't~ i'~'

(f.

100).lm

Fig . 35. Camera lucida

drawings of Penicillium
fellutanum Biourge, eBS
326.48. A , Detailed drawings
of the penicilli. B, Mature
conidia . e, Habit sketches.

Peniciium fellutanum Biourge - Monographie, La Cellule. 33: 262-264. 1923. See

also Ch. Thom, 'The Penicillia', pp. 198-199. 1930; also Raper and Thom, 'A
Manual of the Penicillia' , pp . 212-215. 1949.

Synonymies
Penicillium phaeo-janthinellum Biourge - Monographie, La Cellule . 33: 289-290.
1923.

PenicI11ium dierckx;; Biourg e - Monographle,' La Cellule. 33: 313-315. 1923.

Penicl11ium cinersscens Biourge - Monographie, La Cellule. 33: 30S-309. 1923.
Penicillium fellutanum Biourge , varo nigrocsstsneum Abe - J . Gen. Appl.
Microbiol. Tokyo. 2: 71 -72. 1956. See also Smith , Trans. Brit. Mycol. SOCo 44:
332. 1963; also M.M. Kulik, 'A Compilat ion of Descriptions of New Penicillium
Species'. Agric . Handbook No. 351. Agric. Res. Ser. U.S. Dep. Agric . p. 40.
1968.

Colonies u pon CA (co lour plate 35, A and a) grow ing ra th er restricte dly,
attaining a d ia meter of 20-30 mm in 14 days at room temperatu re
(20- 25 OC ), consisting of a tough, close-textured basal felt, 100- 200 /l m de ep,
with margin oft en surrounded by a narrow zone of pred om ina ntl y submerged,
fu niculose , sto lon- like h yphae , rad iately furrowed , wit h conce n tr ical wrinkles ,
narrowly zonate, covered by a mycelia l overgrowth of snow-white aerial
h yphae in flue ncing greatly th e co lony appearance ; poorl y sporing in sorne
strains, only in marginal areas , wit h centre remaining persistently white ,
conidial areas in light blue-gree n shades (Sguy , Pi. XX , 449) ; ex udate as
colou rless droplets embedded in the m ycelial felt ; odour no t pronounced,
rat he r pleasant; colon y reverse (co lou r plate 35 , a) in pale cr eam to flesh
colou rs or in ligh t vinaceou s shades , in sorne strains greenis h colou red; penicilli
strictly simple, with an occasional branch (fig . 35 , A), com pa ratively large ,
wi th short stipes, usu all y 50-100 /lm in length by 2.0-2.5 /l m in diameter,
wit h wa lls smooth , arising from interlac ing , traili n g or creep ing h yph ae ,
com m only enla rged at the apex in a vesicle up to 4.0-5 .0 p.u: in d iameter;
phiali des in compact clus ters 8- 12 in the verticil, comparative ly large , mostly
10-1 5 /l m long by 2.0- 4 .0 /lm in d ia m eter, with fairly lon g tapered neck ;
conid ia elliptical to subglobose, mos tly abo ut 2.5-3 .0 /l m in long axis , smooth
or conspicuously roughened (fig . 35, B, and SEM plate 6, bottom : left);
con idial chains form ing loosely parallel or divergent , poorl y defined co lumns
u p to 100 /l m in length (fig . 35, C).
Colon ies upon CYA (colour plat e 35, B a nd b) attaini ng a di amet er of
35 mm in 14 days at room temperature (20-25 OC), sim ila r in texture and
cult u ral ch aracte ristics to those grown upon CA, but more deep ly radi a tely
furrowed, with cen tre raised and depress ed in a crate riform pattern , and very
poorly sporing , remaining persistently white in colou r; undulating margin
surrou nde d by a narrow fringe of submerged hyphae extend ing about 1 mm
be yond the aerial growth; colony surface conspicuously zanate; ex ud ate
ab u ndantly p ro d uced in co lourless drops; odo ur lacking ; colon y reverse
(co lour plate 35, b) in cream shades ; pale amber-coloured pigment diffusing
in to the surrou nding agar ; penicill i essen tia lly as described aboye .
C olonies u pon M EA (colour plat e 35, C and c) attaining a d iamet er of
40 mm in 14 d ays at room temperature (20-2 5 OC ), typicall y velv ety,

co ns picu o u sly zo na tc , planc, slightly rai sed in th e ce n tre ; usually hcuvil y

sporin g in blue -g rc cn sh adcs (S guy, PI. XXX , 147); exud a re luckin g; 01111111
faint , indefinite ; ambcr-colourcd pi gmcnt diffusin g into th e su rro u nd ing llJ.(lIr;
co lo n y rev erso (colour plat e :1 5, c) colo u rlcss ; pcnicilli ess e n tia lly as d e~ll' r i l )C' d
aboye, but with lon ger stipes, up to 150 -200 .tm in lcngth , a nd with couirllnl
cha in s formin g lon ger , looscly parall cl colu mns up to 200 .tm lon g .
The spccies d escription is ccntcr ed upon strain C BS 326.48 - NIHU. 71() .. I 1MI
39,734 = ATCC 10 ,443 = [FO 5,761 = QM 7,554 = [JFM 5031, isol utcd in HJ :'I ~ l.
DI' O .W . Richards, and upon Thom's notes (1930) made on BioUlW:' /I tYII(. ,
now lost.
Strains includcd within th e species present th e sarn e lIlorpiloloJ.\h 'tll
ch a racte r ist ics 0 1' conidial structures, but m ay show co nside rab le vnrill tio llll 111
rat e 0 1' growth a nd colo ny a spect , suc h as the co lo n y textu re , murgiu, (' 0 10 111',
cte .

36 , Penicillium roseo-purpureum Dierckx: fig. 36, coIour plate :i6, nud ~'mM

pIate 6 (bottom : right) .

:::::..-----

0 0

o ~ oB oo9
00008

O 00 0
O 00

1~

O
O
O

e ~,- ~
.JJ: ~
~ .S- ...y
~

100 jrm

~
f{.

11'1

Penicillium roseo-purpureum Dierckx - Ann. Soco Sci. Bruxelles. 25: 86. 1901. See
also Biourge 's Monographie, La Cellule. 33: 317-319. 1923; ibid. 36: 452. 1925;
also Ch. Thom, 'The Penicillia' , p. 181 . 1930; also Raper and Thom, 'A Manual of
the Penicillia' , pp. 218-220. 1949.

Synonymies
Penici/lium carmino-violaceum Dierckx - Ann. Soco Sci. Bruxelles. 25: 86. 1901 .
Biourge's Monographie. La Cellule. 33: 281-282. 1923.
Penicillium internum Morotkovsky - Bul. Sci. Rec. Biol. Univ. Kiev. 2: 78-79.
1936.
Penicillium [Citromvcesl sanguifluus Sopp - Monographie, pp. 115-117. 1912.

Colo nies upon CA (co lour plat e 36, A and a) grow ing restrict edl y , att ai n ing
a d iameter of 25- 30 mm in 14 days a t room temperature (20-25 OC ),
gradually raised towa rds th e cen tre , velvety, with shallow wrinkl es in m argin al
zones, wh ich are ab ru p t; close-t extured basal felt , formed of slende r
interwoven h yphae ; very poorl y sporing, pe rs iste n tly white, grayish wh ite to
flesh or pal e pink shades in areas whc re eonidial structu res are m ost ab u ndan t;
exud a te lacking or limited in sorne strains, whi le a bu ndan t in others , in pin k to
vin aceou s red shades ; odo u r faint , indefinite or lacking; eolony rcver se (eolou r
pla te 36, a) white , rosy, orange-red, ora nge or te rra co tta -colo ured, dep ending
on th e strain; pale vinaceous red -colou red pi gm ent d iffus ing slow ly into the
surro u nd ing agar; penicilli strictly sim ple (fig . 36, A) , with sho rt stipes arising
usuall y as sh ort branch es from tr ailing a nd in terl acing ae ria l h yphae , or
occasionally fro m submerged h yphae , u su ally less th an 50 /l m in len gth by
1.5 - 2.0/lm in d iamete r , wit h wa lls smooth ; ph ialides, few in numbcr or
crowdcd , up to 8- 10 in th e verticil, mostl y 6-8 /lm in len gth by 1.5-2 .0 /lm in
di am eter; co n id ia at first ellip tica l, th en su bg lobo se to globose, co m paratively
small , 2.0-2 .2 /lm in di amet er, wit h wall s del ica tely rou gh en ed whe n observed
under the light microscop e (fig. 36, B) , bu t co ns picuo usly roughen ed in a
cere b rifor m pattern wh en observed under th e SEM (pla te 6, bottorn: ri ght) ;
conid ia l chains usu ally sho rt , bu t a litt le lon ger , up to 100/lm , in heavier
spo ru la ting strains (fig. 36, C) .
Colon ies upon CYA (co lou r pIate 36, B a nd b) a tt a in ing a d ia me te r of
30 m m in 14 d ays at room temper ature (20 - 25 OC ) , dupl icating in tex ture and
colony ap pearance th ose grow n up on C Y A, but m ore deepl y rad ia tely
furro wed a nd more int en sely colou red in pinkish sha des; ex udate and odo ur
lackin g ; poo rly spo ring; vinac eo us red-colou red pigm ent di ffusing into the
surro u ndi ng m ediu m ; colony rever se (co lour pla te 36, b) in salmo n pink to
b rown sien na shades; penicilli essentia lly as described a boye .
Colonies on MEA (co lour pla te 36, C a nd c) grow ing res tr icte d ly , at tai ni ng
a di a m et er of 35 m m in 14 d ays at room temperature (20- 25 OC ), velve ty,

now whit c in colour , with thin murgin surrou ndcd by a fringc 01' ycllowish
1I11 lllllc l'ged hyphae cxt c nd iug a ho ut !i IIIln bcyond th c ac ria l gro wth; colony
111'l :\Ce co vc rcd by a m yccli al ovc rgrow th , lendin g a lan nt c a ppeara nce (o th e
('olo ny; poorl y spo ring , but hcavi cr th an on th c abo ve subs tratc ; exu dare a nd
ndou r lncking or ind efi nit e ; colony reverso (colour plate :'36, e) in yellow, dull
III:I nf.(c-brown lo deep brown shades, dep ending on th c strain ; pal e ambc rI'o lolll'e <l pigment diffu sing into the su rrou nd ing agar, becorning hrown in
th c colo nics in terspace. Pen icilli esse ntia lly as described aboye .
'I'hc spec ies description is ce ntcred on the typ e strain c ns 266.29 = NR RI.
~, ()( 'l = \TC C 10,492 = C M I 40,573 = IJFM 5029, isolated by Prof. Ph . Biourge in
1~ ' '2 ~ I , a nd other stra ins having sim ilar m orphology, com b ine d with R aper ami
'l'llllln's original not es (1949) .
ludividual strains are kn own to vary a ppre ciably in pattern such as
vuria tions in colony margin , often showing aggregation of aerial hyphae int o
I'II\'CS or prostrate fas cicles, dep ending on th e su bs tra tu m a nd other
r-uvi ro n me n t a l factors.
'l'he species a ppears to be world-wide in d istribution , but not abu nda nt in
II II III I'C . R epresentative stra in s of P. roseo -purpureum hav e occa sionally bccn
slIlatcd from soil and from det eri or ating military eq u ipment in tr opi cal a nd
Id ,1 ro pical cou ntrie s.
:17 . Penicillium glauco-lanosum Chalabuda: fig. 37, colou r platc 37, and

SEM plate 7, (top: left).

Ponicillium glauco-Ianosum Chalabuda - Nov. Syst. Herb . Inst . Bol. Komarovii ,
Aead. Sei. USSR . 6: 165. 1950. See also M .M . Kulik , 'A Comp ilation of
Deseript ions of New Penicil/ium Species' , Agrie. Handbook No. 351. Agrie. Res.
Ser. U.S. Dep. Agrie. p. 41. 1968.

C olonies upon CA (colour piate 37, A and a) growing moderately , attaining

.-1 diamet cr of 45 mm in 14 days at room temperature (20-25 OC), cons isting 01'
:1 lou gh , close-t extured basal felt , deeply velv et y to lanate, with shallow
lurrows in a radiate pattern ; thin , undulating margin surrounded by a fringe of
xu b m er ged h yphae exte nd ing about 5 mm beyond th e aerial growth; poorly
sporing, espec ially in marginal are as in light gray shades (Sguy , XXXIII,
'\!J5 ); exudate and colour lacking; colony revers e (col our piat e 37, a) in pale
crcum shades ; pal e a mbe r-colou re d pigment diffusing into th e su rrou nd ing
a~a r; penicilli strictly sim ple (fig. 37, A) , with short stipes, 19-50 p.m in length
by 2.5-3 .0 p.m in diameter, with walls delicately rough ened or smooth , comuionly arising a s short branches from prostrate cree pin g h yphae and ropes of
hyph ac, with apices enla rged in a vesicle up to 3-5 tuu in diameter; phialides
kw in number, 2-8 in th e verticil , mo stly 6.5-10 p.m by 2.0-2 .5 usu,

00 cP OO
o
O O

0 0 o

;)7
"-../

Oo OO O
B O0 0 O
00
O
O O O 0
O

lu.u1J.wJ

10 jim

11~~1}
~

100}lm

Fig. 37. Camera lucida

draw ings of Penicillium
glauco-Ianosum Chalabuda,
CBS 224.66. A, Detailed
draw ings of the penicill i. B,
Mature conidia. C, Habit
sketches.

terminating in a conspicuous ly tapercd , narrowcd, long neck ; conid ia elliptical

to subglobose, smooth walled or ne a rly so (fig . 37, B, a nd SEM platc 7, top:
left) , mostly 2.5-3 .5 .tm in long ax is, occas ionally up to 4.5 utt; conid ial
chains forming loosely parallel colu m ns, or tangled (fig . 37, C) .
Colonies u pon CYA (colour plate 37 , B and b) att ain in g a diamet er of
42 mm in 14 days at room tempcrature (20-25 OC ), duplicating in texture a nd
pattern tho se grown upon CA , but deeply radiately wrinkled; heavi er sporing ,
espe cially in marginal areas in pale gr ay shades (Sguy, PI. XXXIII, 495) ;
evenly margin wit h a very narrow fringe of sub me rged hyp hae exte nd in g no
more than 1 mm beyond th e aerial growth ; colony surface in wh ite to white
ros y shades in poorly sporulated a re as ; exudatc and odour lacki ng ; colony
reverse (colour pia te 37, b) in cre a m shades to light yellow colours ; ambercoloured pi gment diffu sing into the su rro u nd ing aga r ; penicilli esse ntially as
des cr ibed above .
Col oni es upon M EA (colour plat e 37, C and c) spreading broadly, a tta inin g
a diamet er of 55 mm in 14 days at room temperature (20 -25 OC ), loose
text urcd , vclvety , with a tendency to produce densely lanate sectors of white

Icl'i1l: h yph uc , pl anc , a ppcaring gra nula r; thin m nrgin su rro u ndc d by a Irin gc
suln ucrged h yph ac extc nd ing ahou t 4 11I11l bcyond th e ac ria l grow th ; hcavily
i1H'inl-{ throu ghout in gr ay sh adcs (Sgu y , PI. XXXIII , 49 3); exud ate a nd
udu u r lnckin g ; colony revcrse (colour plate 3 7, e) in very pale cre a m shade s;
u ub cr-co lou red pigm ent difTusin g into th e su rro u nd ing agar ; penicilli
llNNl ' lI 1 ially as described aboy e .
SllC :cics di agnosis is cen tc re d upon th e typ e strain CB S 224 .66 = BK M Fj - \ T CC 18 ,47 8 =IJ FM 5027, isolated from Ukranian soil , in th e enviro n s
lit' Kic w, U SSR , by T.V . Chalabuda in 194 9 .
A d ose rel ationship between P. glauco-lanosum Chalabuda and th e P.
1t'/'/II/1!Jm .l' se ries is not presumed here , but th ey are con sidered toge the r
1Il'IIl arily as a m atter 01' conve n ience. They do , how ever, have ce rta in features
11 1'11111mon , particularly th e tough , close ly wo ven felt offine h yphae , th e com prm uively slow-gro wing colonies , espec ially on CA, and th e sho rt stipes,
uu rs t ly arisin g from tr ailing 01' pros trat e cree ping hyphae.

111'

~I Il ,

Penicillium brevissimum Rai el Wadhwan : fig. 38, colour plate 38,

:11 1<1 SEM plat e 7, (top: right).

f~
---

(~

o
O O
00
B O O
O O O O 00

0 0

0 0

00

00

0
CJ 0

R.

~~
e

100

jrrn

Fig. 38. Camera lucida

drawings of Penicillium
brevissimum Rai et
Wadhwani, CSS 763.68. A ,
Detailed drawings of the
penieilli . B, Mature eanidia .
e, Habit sketehes . O, Rapes
of hyphae with eanidial
struetures arising frorn them .

terilc hyphu c , plane , nppca ri ng gra nular ; (hin ma rgin SlIITO U IH lc d by a Iringu
JI' suhmcrged hyph ac extcnd ing abo ut '1' II1l1l bcyond thc aerial g-rowlh; hcavily
p"l'ing" rhrou gh out in gray sha dcs (Sguy, PI. X XXIII , 493); exudate a nd
HlI lIll' lacki ng ; colony reverse (colou r platc 37 , c) in very pale cream shades;
uubcr -coloure d pigm cnt di ffusin g int o th e surrou ncling agar ; pen icilli
(' HMI ' 1I1ially as dcscrib ed aboye .
Spccics di a gn osis is cen tere d u pon the type strain CBS 224 .66 = BKM FH;I I -ATCC 18 ,47 8=IJFM 5027, isola ted frorn U krania n soil, in the environs
I JI' Kic w, U SSR , by T .V . C halabuda in 1949.
\ close relat ion sh ip bet ween P. glauco-lanosum C hala buda and the P.
ircnmbcns series is not presu me d here , but they are considered togethe r
primuri ly as a m atter 01' convenie nce . They do, however , h ave certain fea tures
111 co tum on , particul arl y the tou gh , closely woven felt 01' fine h yphae , th e cornpnrurively slow-gro wing colonies, especially on CA, and th e short sti pes,
nll lslly arising from trailing 01' prostr ate cree ping hyphae .

:-111 . Penicillium brevissimum R ai el W adhwani : fig. 38, colour plate 38,

.md SEM plate 7, (top : right ).

(~

o
O

O O 10
O O O 00

0 0

00

0 0 0

0 0

O ()

R.,

Fig. 38. Camera lucida

drawings of Penicillium
brevissimum Rai et
Wadhwani, CSS 763.68. A,
Detailed drawings of the
penicilli. S, Mature conidia.
C, Habit sketches. D, Ropes
of hyphae with eonidial
struetures arising from them .

Penicillium brevissimum Rai ot W adhwani - Curr o Sci. 45: 192-193. 1976.

Colonies up on C A (colour plat c :1 B, A and l) growi ug' rcstri ctcdl y, a l tainin g
a di amet er 01' 23 mm in 14 d a ys a t roo m tempcrature (:2 0- :25 OC ), consisting 01'
a tou gh , almost, leathery basal felt , close -tc xtu red , velvc ty in a p peara ncc ,
deepl y radia tely wrinkl ed in recent isolat es, but with few sha llow furrows in
strains lon g m aintained under lab orat ory arti ficial co nd itio ns; ve ry po orly
sporing, remaining persistently white with cent re in pale yellow shade s,
pr oducing very minute co ni d ia l struc tures; exudate and odo ur lacking ; colony
reverse (co lou r plat e 38, a) in pale yellow shade s; coloured d iffusin g pi gm ent
not observed; penicilli strictly sim ple (fig . 38, A) with very sho rt stipes, usu all y
11 - 80 11m in len gth by 2.5 -3. 0 I1min di amet er , with a pices slightl y enla rged
in a ves icle, m ostly arising from tr ailing a nd in terwoven aeria l hyphae 0 1' rop es
01' hyph ae ; phial ides few in nu mber , 3-6 in the verticil, loosely a rra nged ,
6-9 11m by 2.5-3 .5 11m ; co nid ia elliptica l to cy lindrical, smooth walled 0 1'
nearl y so (fig . 38, E , and SE M plat e 7, top : ri ght), 2.8-4. 0 11m by 2.0-3. 0 11m
in d ia meter; conid ial chains forming loosely to well-defin ed colu m ns (fig. 38,

C) .
Colo nies upon C YA (co lou r plat e 38, R and b) gro wing res tr icted ly, a ttain ing a di amet er 01' 21 mm in 14 d ays at ro om temper ature (20-25 OC), deepl y
ra d iately fur row ed, with ce n tre rai sed in a cra te r iform pattern ; eve nly m argin
su r ro u nde d b y a narrow fringe 01' su bmerged hyph ae exte nd ing abo u t 1 mm
beyond th e aeri al gro wth; colony surface poorl y spo ring, remaining
pe rsiste n tly whi te to wh ite ros y , velvet y in a ppeara nce ; odo ur faint , indefinite ;
exudate lackin g; colony reverse (colo ur plat e 38, b) in cin namo n shades; pale
am be r-co loured pi gm ent diffusin g into th e surrounding ag ar; penicilli
esse ntially as described a bo ve .
Colon ies upon M EA (co lo u r plat e 38, C a nd c) growing restrict edl y ,
at ta ining a di amet er 01' 25 mm in 14 d ays a t room temperature (20- 25 OC),
velve ty, plane, sligh tly co nvex ; white , wide, evenly m argin ; heavily sporing
th rou gh out in blue-gr een shades (Sguy, PI. XXXI , 462); ex ud ate lackin g;
odour fragra nt, aromatic; colon y rever se (colou r piate 38, e) in light yellow
shades; pal e a m be r-co lou re d pi gm ent di ffusin g into the surrou nd ing aga r ;
penicilli esse nt ia lly as described a boye, but with th e sho rt stipes arising m ostl y
from trailing rop es 01' hyphae ( Fig . 38, C ).
Sp ecies description is ce nte re d up on th e type strain C BS 763.68 = IJ FM 5036,
isolat ed from rizosphere soil 01' Brassica campestris, var. toria, in Lu cknow , Indi a ,
by j .N . Rai and K. Wadhwani , in 1968.

39. Penicillium bilaiae Chalabuda : fig. 39, colour plat e 39 , and SEM
plate 7, (cen tre: left) .

7~~
00000

000 O
O

000
O

BO
0

O
O
O

i.J.J..ulillJ.J

10

}1m

R.
>V

Ce ".(

"t\[ '111
L--J

100

jrm

Fig. 39. Camera lucida

drawings of Penicillium
bilaiae Chalabuda, CBS
221.66. A, Detailed drawings
of the penicilli. B, Mature
conidia . C, Habit sketches.

Penicil/ium bilaiae Chalabuda - Nov . Syst. Herb . Inst. Bot. Komarovii, Aead. Sei.
USSR. 6: 165. 1950. See also M.M. Kulik, 'A Compilation of Deseriptions of New
Penicil/ium Species'. Agre. Handbook No. 351. Agrie. Res. Ser. U.S. Dep.
Agrie. p. 20. 1968.

Colonies upon CA (colour plate 39, A and a) growing rather restrictedly,

altaining a diameter of 35 mm in 14 days at room temperature (20-25 OC),
consisting of a tough, close-textured basal felt, velvety, sulcated ratiately by
sh allow furrows; undulating, thin margin, surrounded by a fringe of brown
colou red submerged hyphae extending about 1- 2 mm beyond the acrial
growth : colony surface covered by a mycelial white overgrowth, and hencc
influencing the colony appearance; poorIy sporing in whitish gray shades;
odour lacking; exudate in amber-coloured drops; a m b er-colou re d pigment
diffusing into the surrounding agar ; colony rcverse (colour plate 39, a) in dark
reddish brown shades; penicilli strictly simple (fig . 39 , A), with short stipes,
10-100 .,m in length by 1.5-3 .0 .,m in diamet er, smooth walled, mostly
ari sing from trailing interwoven or prostrate creeping hyphae, slightly

cnla rged al. api ces in a veside up lo :L5 P.III in di um ctcr; phi alid es usually few
in number, 2-6 in the verticil, 01' crowded in clusters , 9-14 in thc vcrt icil,
7-9 p.m by 2.5- :L5 J.tm, terrninating in a co nsp icu ously tapercd , sho rt,
narrowed neck ; con id ia usually g!obose to subglobose , apparcntly smooth
when obs erved under the light microscope (fig . 39, B) , but de!icatc1y
roughened when obs ervabl e under th e SEM (plate 7, centre: left), 2.0 - 3.0 J.tm
in diameter; conid ial chains forming more 01' less loosely parallel colu m ns, up
to 150 J.tm long (fig. 39, C).
Colonies upon CYA (colour plate 39 , B and b), attaining a diameter al'
35 mm in 14 days at room temperature (20-25 OC ), consisting al' a closetextured bas al felt , velv ety , deeply radiately sulc ate , with numerous furrows ,
with centre raised and irregularIy folded in a crateriform pattern; sporing
abundantIy in pale blue-green shades (Sguy , PI. XXX , 444) ; exudate
abundant, in amber colou red drops; odour lacking; amber coloured pigment
diffusing into th e surrounding ag ar ; colony revers e (colour piate 39, b) in dark
brown shades; undulating margin surrounded by a narrow frin ge al'
subm erged hyphae exte nd ing 2-3 mm beyond th e aeri al growth; peni cilli
essentially as described aboye.
Colonies on MEA (colour piat e 39, C and c) attaining a diameter al' 30 mm
in 14 days at room temperature (20-25 OC ), velvety in appearance, plan e,
slightly raised in central areas; colony surface cave red by a my celial
overgrowth a l' int erwoven white ae rial hyphae , and rapes a l' hyphae en crusted
with dark purple-red crystals , lending a pinkish appearance to the colony;
poorIy sporing in pal e blue-green shad es (Sguy , PI. XXX , 444); thin ,
undulating margin , surrounded by a fringe al' purple enc ru sted subme rged
hyphae ext ending about 5 mm beyond th e ae rial growth; odour and exudate
lacking; colony revers e (colour pIat e 39, e) in dark purple shad es; arnbe rcoloured to vinace ous red pigment diffusing into the surrounding a ga r;
peni cilli essentia lly as described aboye .
The spec ies description is centere d upon th e type spec ics C BS 221. 66 = BKM
F -854 = IJFM 5025, isolated from Ukranian soil in the enviro ns a l' Ki ev, USSR ,
by T .V. Chalabuda, in 1949.
A clase relationship between P. bilaiae Chalabuda and th e P. decumbens seri es
is not presumed here . However , th ey are con sidere d tog ether, primarily as a
m atter al' conve nience . They do , how ever, show certa in characte ristics in
com mo n, such as the slow-growing colon ies, espec ially upon CA, and th e short
stipes, usuall y arising from tr ailing 01' prostrate cree ping hyphae , reminding
thos e species included within th e P. f ellutanum sub- series.

40. Penicillium kurssanovii ChaIabuda : fig . 40, co !o ur pIate 40 , a n d SEM

pl ate 7, (centre: right).

~ 00 0
o O

0 0

O ~O OO

000
0

lw.L!.u.J..U

10

)Jm

fR .

~ i-ll
I--J

100 ).1m

Fig. 40. Camera lucida

drawings of Penicillium
kurssa novii Chalabuda, CBS
625.67. A , Detailed drawings
of t he penicilli . B, Mature
conid ia. C, Habit sketches .

'oniclllium kurssanovii Chalabuda - Nov. Syst. Herb. Inst . Bot. Komarovii, Acad .
Sci. USSR . 6: 164. 1950. See also M.M. Kulik , ' A Compilation of Descriptions of
New Penici/lium Species'. Agric. Handbook No. 351. Agric. Res. Ser. U.S. Dep.
A gric . p. 48. 1968.

Colonies upon CA (colou r plat e 40, A a nd a) grow ing mod erately a nd

lI tlilining a di amet er of 35 mm in 14 days at roo m temperature (20-25 OC),
cousisting of a toug h , clos e-textured basal felt, with surface somewhat
u unon tose , plane ; thi n, eve n m argin surrounded by a fr in ge of submergcd
hyph ae ex te nd ing abou t 1- 2 mm beyond th e ae ria l growth; spo ring
moclcr at ely in pa le blue-green shades (Sguy, PI. XXX , 440) ; exudate and
od our lacking ; colony reverse in ligh t yellow shades (colour plat e 40, a); palc
umbcr-colo u re d pigm ent di ffu sin g into the su rrou nd ing agar; penicilli strictly
simple (fig. 40, A), with ve ry sho rt stipes, mostly 10- 30 p.m in length by
1.0 - 2.0 p.m in diame te r, with a pices enlarged in a ves icle up to 3.0 p.m in
rliam eter , a rising prim arily fro m interwoven trai lin g hyph ae , sm ooth wa llccl;
phialides usuall y few in number, 3-6 in th e ve rticil, 5.5 - 7.5 p.m , by

2. 0 -:~ . O

ttlll , tc rmln ut in g in a sho rt , na rrowcd neck: co n id ia a l Iirst c llip tica l,

thc n glo bosc , d clicut ely sp in u losc wh cn o hsc rve d undcr thc light microscope
(fig. 40, B), but cons picuo us ly rou gh cn ed wh cn ob serv ed under th c SEM
(plate 7, centre: right) , mostly 2. 2-3.0 ttm in diameter; conid ia l cha ins
forming loosely par allel , short eolu m ns, up to 40 ttm lon g (fig. 40, C).
Colonies upon C YA (eolour plat e 40, B a nd b) atta in ing a di amet er of
35 mm in 14 days at room temperature (20- 25 OC ), duplieating in texture
th ose grown on C A, but with th e colony surfaee deeply radiat ely wrinkled,
rais ed and depressed in central areas; abrupt m argin ; surfaee coloured in the
same light blue-gr een shade s (Sgu y , PI. XXX , 440); eolonies stro ng ly pulling
apart and cracking the su bs tratu m (eolour plat e 40, b); odo u r faint , indefinite ;
exuda te laeking; colony reverse (eolou r plat e 40, b) in einnamo n shade s; pale
am ber eoloured pigm en t diffu sin g int o th e surro unding agar; peni eilli
esse ntially as deseribed aboye .
C olon ies upon M EA (eolour plat e 40, C and e) att a ining a di amet er of
37 mm in 14 day s at room temper ature (20-25 OC), deepl y velve ty to
su b floccose , plane with centre slightly umbonat e ; thin m argin su rro u nde d by a
fringe of su b me rge d hyphae ex te ndi ng abo ut 3 mm beyond th e ae rial gro wth ;
sporing abu ndantly in pale blue- green shade s (Sg uy, PI. XXX , 440); odo ur
faint , aromatic; exuda te laekin g ; eolony reverse (eolour plat e 40, e) in pale
yellow shades; pale am ber-colou red pigm ent diffu sin g into th e su rround ing
ag ar; penieilli esse ntially as deseribed abo ye.
The species deseription is eente re d upon th e typ e spec ies CBS 625.6 7 = BKM F1244 = C M ! 129,965 = AT C C 18,387 = IJ FM 5045 , isolat ed from Ukrani an soil
in a Zea mais field in th e env iro ns of Ki ev , USSR , by T .V. C hala buda, in 1949.
The speeies as origin ally descr ibed , and as obse rved in ou r cultu res, however ,
sho w some di serep aneies, sue h as the eon id ia orname ntatio n. Whil e in
C hala buda's original di agn osis eo nid ia are deseribed as srnooth-walled, in our
own expe rienee th e conid ia are eonspieuo us ly rou ghened . Otherwise , th e
deser iption as presented abo ye is in satisfaetor y agr eem ent with th e origin al
di agn osis of the spee ies (1950).
No close relation ship between P. kurssanovii C hala buda and the spee ies
included wi thin th e P. decumbens is presumed here , bu t th ey are eons ide re d
together as a matter of eonven ience. They do , however, show eerta in
eharae te risties in eom mo n, sue h as the slow-grow ing eolon ies, th e tou gh , closetextured felt of fine hyphae , a nd th e sho rt stipes , m ostly arising as sho rt
braneh es from tr a ilin g hyphae, suggesting th ose Penicillium spee ies included
within the P. Jellutanum su b -series .
4 1. P enicillium cinereo-atrum C halab u da : fig . 41 , co lo u r pl a te 41 , and
SEM pl ate 7 , (bo tto m : left) .

:::::::--------

::.;::-oo"}

.,-,

~\" ',

.-. ,

~;

,.:,

'<i
-,.J

i.wWJ.uJ

10

jrrn

'1flJ1J
100

)Jm

Fig. 41. Camera lucida

drawings of Penicillium
cinereo-atrum Chalabuda,
CBS 222.66. A , Detailed
draw ings of the penicilli . B,
Mature conidia. C, Habit
sketches.

Penicillium cinereo-atrum Chalabuda - Nov. Syst. Herb . Inst. Bot. Komarovii,

Aead . Sci . USSR. 6: 167. 1950. See also M.M. Kulik, 'A Compilation of
Deseriptions of New Penicillium Species'. Agrie. Handbook No. 351. Agrie. Res.
Ser. U.S . Dep. Agrie. p. 26. 1968.

Co lon ies upon C A (colour plat e 41, A and a) attaini ng a diamet er of 45 mm

in 14 days at room temperature (20 -25 OC), cons isting of a tou gh , c1oseicx tu red ba sal felt , deepl y velve ty to lanat e , plane , slightly and grad ually
raise d towards th e ce ntre ; eve n m argin with a narro w fringe of su bme rge d
hyphae exte nd ing less th an 1 mm beyond th e ae rial gro wth; very poorIy
sporing, with colony surface cave red by a m ycelial ove rgrowth of closely
inte rwove n ae rial hyphae , influencin g the colony appeara nce, and lendin g it a
persist ently sno w wh ite colou r; exudate la cking ; odo u r faint , indefinite;
colony reverse (colou r pIate 41, a) in pale sulph ur yellow shades; yellowcoloured pigment diffusin g into th e surrou nd ing agar; peni cilli strictly sim ple
(lig . 41, A), with ver y short stipe s, m ostly 20-40 p.m by 1.5- 2.0 p.m , arising
primarily from trailing or prostrat e cree p ing h yphae , smooth -wa lled, with
apices slightly enla rge d up to 3. 0 p.m in di ameter ; phi alides com pa ra tively

small , Iew in number , 2 - ;> in th e verticil , 01' somutime c ro wdc d IIp lo 12 pcr
whorl , comruonly 5.0 -9 .0 ~ 1lI by 2.0 -:LO ~ nl , tcrtuinut ing in a short , rap er ed,
narrowed neck ; conid ia at lirst ellipso id a l, bccomi ng subglobosc, 2.5 -3 .5 ~m
in d iamet er , conspicuously echinulatc (lig . 41, B, and SEM plat o 7, bottom:
left) ; conid ial chains short , up to 30 ~m long, forming loosc1y d ivergen t
01' tangled colu m ns (fig . 41, C) .
Colonies upon CYA (colour pIat e 41 , B a nd b) a ttaining a di ameter of
30 mm in 14 days at room temperature (20 -25 OC ), cons isting of a tough,
close -textured basal felt, de eply and radiately wrinkled ; margin en tire ; poorly
sporing, with a colon y su rface remaining persistently white rosy ; ex uda te
lacking ; odour faint, ind efinite ; colon y reverse (colour pl at e 41 , b) in ligh t
cinnamon shade s; amber- coloured pigment diffusin g into th e surrounding
agar ; peni cilli esse ntially as des cribed aboye .
Colonies upon MEA (colour plate 41 , C and c) attaining a d iamet er of
40 mm in 14 days at room temperature (20 -25 OC ), conspicuously zonate,
ve1vet y to lanate , due to a loose network of fine ae rial white h yphae ; m argin
thin , with a wide fringe of submerged hyphae ex te nd ing about 3 mm beyond
th e aerial gro wth ; exud ate and odour lacking; sporing ab u nd a n tly in blu e-gray
shades (Sguy, PI. XXXIV , 50S); co n id ial structures somewhat obs cured in
the colo ny centre by the myce1ia l overgrowth of interwoven floccose hyphae ;
colony reverse (colour plate 41 , c) in crea m sh ades ; pale amber -coloured
pigment diffusing into the surrou nd ing a ga r; penicilli essen tially as described
aboye .
The sp ecies description is ce ntere d upon strain CBS 222.66 = BKM
F -S56 = IJFM 5024 , isolated from Ukranian soil in th e enviro n s of Ki ev , USSR ,
by T.V . Chal abuda, in 1949.
The sp ecies as originally described, and a s observed in ou r cu lt u res, agrees
satisfactorily in all respects, excep t that it has become somc wha t altered in
cultu ral aspect, and predominantly steril e upon CA and CYA, after 30 years of
laboratory cu ltivation .
Although a close re lationship between P. cmereo-atrum Chalabuda and the P.
decumbens series is not presumed here, th ey are con sidere d together, primarily
as a matter of convenience . They do , however , show certain characteristics in
com m on , su ch as th e com para tive1y slow-growing colon ies upon most
substrata , th e short , slende r stipes , arising from trailing hyphae, su ggesting
those species of Penicillium included within th e P. f ellutanum sub-series.

42 . Penicillium toxicarium M iyake ex Ramrez : fig. 42, coIour pIate 42,

and SEM plate 7, (bottom: rig ht) .
Penicillium toxicarium Miyake - Report on Rice Utilization l.ab, Japan. 1:1. 1940.
See also 1. Miyake . Jap. J. Bacteriol , 9: 1105. 1954; also S. Abe. J. Gen. Appl.

E) ... @
ttZ(\ ~
~ ~

~ ~
': ' ~.;."
/ ~
-!~; ~ Yit9
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Fig. 42. Camera lucida

drawings of Penicillium
toxicarium Miyake, ex
Ramrez, CBS 351.51. A ,
Detailed drawings of the
penicilli. B, Mature con idia .
e, Habit sketches .

Microbio!. Tokyo. 2: 57.1956; also M .M . Kulik , 'A Compilation of Descriptions of

New Penicil/ium Species'. Agric. Handbook No. 351. Agric . Res. Ser. U.S. Dep .
Agric. p. 75. 1968.

Penicillium toxicarium Miyake was originally isolat ed from contam inated

,Iapa nese rice, by 1. Miyake , in 1940 , who proposed th e name P. toxicarium a nd

dcscribed it without providing a Latin diagnosis. In order to va lidate the

species, a Latin diagnos is drawn from ou r obs ervations is given below . I t
dilf ers in m inor details from Miyake ' s original di agnosis . T he name of the
ori ginal au thor is maintained under Recommendation 46 C of th e Internati onal Code of Botanica l Nomenclature (Stafl eu (Ed.), 1978) .
Latin d iagnosis of Penicillium toxicarium Miyake ex Ramrez :
Coloniae in agaro Czapekii, post duas hebdomadas ad 20-25C , 45 mm in
diamet ro attingunt, velutinae , denso texto basali coacto mycelio, inaequal iter
prominentes, plicatae, fortiter radiatae plicatae et aliquantulum lanosae ad centrum,

cum margine asqualiter a hyphis subm ersis 2 mm lat ltudlne formatur; lenlter
sporulantes, primum albas, dainde vlrido pallido colorantur; odor abest; axsudatio
adest, succineo; reverso pallido fusco vel salmoneo; cum colore succineo fusco in
medio diffluente; neque ascomata neque sclerotia visa sunt; penicilli pleraque
monoverticillati, stipitibus erectis, brevis aut mediocriter longis , plurimum e hyphis
reptabundis exoriuntur, plerumque tenus 100-200 Lm longitudine per 2.0-2.5Lm in
diametro, cum apicibus tenus 3-4Lm inflatibus; phialides numerosae, usque 8-12
per verticillum, plerumque 7.5-10 Lm longitudine per 2.5-3.5 Lm in diametro;
conidia globosa, 3-5 Lm in diametro, pariete aspere; in catenis compactis tenus
25-50 Lm longis.
Coloniae in agaro Czapekii cum extracto levedinis, post duas hebdomadas ad
20-25C, 55-60 mm in diametro attingunt, velutinae, radiatae plicatae, albidae vel
griseo viridae; leniter sporulantes; odor et exsudatio absunt; reverso fu seo in
centrum et virido fusco in marginem; cum colore fusco atro in medio diffluente;
neque ascomata neque sclerotia visa sunt.
Coloniae in agaro hordeaceo celerior quam in agaro Czapekii crescunt, post duas
hebdomadas ad 20-25C, velutinae, planae, margine plerumque a hyphis
submersis formatur; copiosae sporulantes, superficiebus conidiorum griseo
viridibus; exsudatio et odor absunt; neque ascomata neque sclerotia visa sunt.
Reverso flavido.
Typus cultura cas 351.51 ex Driza sativa L., Japoniae, a 1. Miyake isolata est
anno 1940.
Miyake's original diagnosis follows:
'Colon y growth of this fungus is restricted , in a radial pattern; attarnmq a
diameter of 2.5-3.0 cm in 14 days. With central area raised and somewhat
irregularly folded, but with the rest of the colony surface remaining smooth and flat.
The raised part is cracked and lanate . The colony margin is 0.5-1.0 mm wide; the
colony shows yellow colours at the obverse as well as at the reverse. The
conidiophores are 100-200 Lm by 3.0-3.5Lm; penicilli are strictly monoverticillate;
phialides are crowded in c1usters at the apex of the conidiophores, 7.5-8.0 Lm by
3.0-3.5 Lm. Conidia are somewhat globose, 2 .0-2.5 Lm in diameter, and forming
chains 25-50 Lm long .'
(Translated from the Japanese by Miss Masae Yamamoto) .

Our not es folIow :

Colo n ies upon C A (colou r pl at e 42, A a nd a) gr owing m od era tely , attain ing
a di ameter of 45 mm in 14 days at room temperature (20 -25 OC ), co ns isting of
a tou gh , close- tex tu red ba sal felt, covere d by a m ycelial overgrowth of closely
in te rwoven ae r ial hyphae , lending a deepl y velvet y to lan ate a ppearance to th e
colony su rface; deepl y radiat ely furrow ed , so mewhat rai sed in th e colony
ce ntre; a ppear ing white in m arginal areas a nd pale blue-green (Sguy , Pi.
XXX , 438 ) coloured in conidial a reas, especialIy in ce nt ral a n d subcentral

arcas; thin , cvc n m argin surroundcd hy a l'ringe o l' suhmcrgcd hyph ac

111111 hc yon cl th e ucri al g ro wlh; oclo ur la ck in g ; ex ud a re in
.uuhc r-colou red dropl ct s ; co lo n y reve rso (colo u r piat e 42, a) in palc brown to
salrnon pink shades; brown-coloured pigrncnt dillu sin g in to th e su r ro u nding
a gar; penicilli str ictly sim p le (F ig . 42, A), with sho r t stipes, 50- 100 Jlm b y
'2 .0 - 2. 5 u tn ; smoo th wa lled, m ostl y a rising frorn tra ilin g hypha e , w ith a p ices
cnla rge d in a ves icle up to 4 Jlm in di ameter ; phialides , few in number, 3-6 in
I he ve rticil, 0 1' crowde d in clusters up to 12 in the whorl , m ostI y 7. 5-1 O tut: b y
'2 .5 -3 .5 iut , terminating in a sho rt , tapered , a n d n arrowed neck ; co n id ia
globose, 3.0 -5 .0 Jlm in d iamet er , wi th wall s de1icat ely ro ughe ne d, with
ro ug hness a r ranged in undul ating , parall e1 ban ds (S E M pl ate 7, bo tt om :
I'i ght ); co n id ia l chains in loosely p arall el to definitel y co m pact colu m ns u p to
'lOO Jlm long (fig. 42, C) .
Colo n ies upon C YA (colour pl a te 42 , B a nd b) attain ing a diameter of
~)5 - 6 0 mm in 14 da ys at room temperature (20 -25 OC ) , duplicat in g in tex ture
a nd cu ltu ral characteristi cs those grown upon C A, bu t show ing m ore rad ial
urrow s , a nd sporing less a b u n d a n tIy; ex u date a nd odour la cking; co lony
reverse (co lou r pla te 42, b ) sho wing d ark brown colou rs in ce n tral a nd
subc en tra l a reas a nd deep green- black sh ad es in th e m argin al a reas ; dark
brown-coloured pi gment d iffusing into th e surrounding agar ; penicilli
cssc ntially as described aboye .
Colon ies upon M EA (co lour pi a te 42, C and e) spreading broadl y , at ta in ing
a d ia meter of 70 mm in 14 days at room te m pe rature (20-25 OC ) , close
icxtured , vel ve ty to lanat e , co nvex, plane , wi th colo ny su rface covere d by a
111 ycelia l overgrowth of thin floccose , closely interwoven h yphae , lending a
lanate a ppearance to th e colony; heavily sporing through ou t in gr ay-green
shadcs (Sguy, PI. XXVII , 403); th in m argin su r ro u nded by a l'ri nge of
sub m erge d hyphae ex te nd ing large1y , up to 5 mm beyond th e ae r ial growt h;
ex u da re a n d odo ur lacking ; colony reverse (co lou r pl at e 42 , c) in yellow shades;
eolou red pi gment d iffusin g into th e surro u nd ing agar not ob served; penicilli as
desc r ibe d a boye, b u t with longer stipes a r ising primarily from th e su bstratu m ;
eon id ia l ch a in s for rnin g we ll-de fine d colu mns .
Sp ecies description is ce n te re d upon th e typ e stra in C BS 35 1.51, isolated
fro rn contam inated J apanese rice , by 1. Miyake , in 1940. lt has becom e
some what alte red d uring th e 40 years of co n ti n uo us laboratory cu ltivation.
The species as or igi nally descr ibe d, a nd as obse rved in ou r cu ltu res , di ffers
rna rkedly in several characte ristics ; name1y , th e ra te of growth , colony
colou ring , diffu sin g pigment , m easurements a nd sculpturing of co n id ia .
A close re la tions hip between P. toxicarium M iyak e a nd th e P . decum bens se ries
is not p res u med here , but th ey are co n sidered togethe r as a m atter of
conve n ie nc e . They do , how ever, show ce r ta in features in co rnmo n,
pa rticu la rly th e sho rt stipes , a r ising fro rn trailing fin e hypha e , a nd th e tough ,
cx tc n d ing a bo ut 1'2

close-tc xturc d b asal fclt, su ggc sting (hose specics includcd within th c P.
f ellutanum su b -se ries .

43. Penicillium gerundense Ramrez el Martnez: fig. 43 , colou r platc 43,

and SEM plat e 8 , (top: left) .

~~
OO O 1~
pm

00 0 0
O BOa
O o o

:
, J.". iJ/

00

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O
O

-----

"" , !7

i.' '
c

. 7;~'<:

L.......-....J

100 jJm

al

Fig. 43. Camera lucida

drawings of Penicillium
gerundense Ramrez et
Martnez , ATCC 42,234. A,
Detailed drawings of the
penicilli. B, Mature conidia.
C, Habit sketches.

Penicillium gerundense Ramrez et Martnez - Mycopathologia . 72: 181-191,1980 .

Colonies upon CA (colour pl ate 43 , A a nd a) gro wing restr ictedly, atta inin g a
di amet er of 20 mm in 14 days a t roo m temperature (20 - 25 OC ), co ns isting of a
tou gh , close -tex tu re d basal felt , stro ngly umbonat e ; white , a b ru p t, undulat ing
m argin , sulcated with few shallow wrinkles, velvety in a ppeara nce; colo ny
su rface remaining persistently white , covered b y a m ycelial overg row th of fin e
aerial hyphae forming a dense network in th e cen tral area; very slig h tly
spo ring; ex udate and odo u r lacking ; reverse (co lour plat e 43, a) in pale
sa lrnon -pin k shades; no coloured pi gm ent d iffu sin g in to th e su rro u nd ing agar ;
penicilli strictly sim ple, with th e sti pes of th e con id iopho re s sho rt and slender,
u su all y less than 100 11m in len gth b y 2.0 11m in di amet er, smooth wa lled,
arising com monly fro m the den se net work of trailing hyphae ; phiali d es bo ttle

4:~,

A), tc rmin utin g in a short-poi nte d ncck, in compact clusters 01'

10-14 in th c vcrticil, mostly 7.0 -10 .0 J-tlll h y 2.:l -:~ .!i J-tlll; conidia ovoid lo
d lipsoidal , usu ally 2.!i- :1.() J-tlll by 2.0 -2. :l J-trtl in diam eter , a p paren tly
xruooth-wallcd wh cn observcd under th e light mi croscope (lig. 43, B) , but
hnpcd (Iig .

ddi calcly rou gh en ed wh en ob se rve d under th e SEM (pl ate 8, top : left) ;
l'llllidial ch a in s formin g loosel y parall el co lu rn n s (li g. 43, C ) .
C olon ies upon CYA (co lou r plate 43, B and b) growing faster than those
Kro wn upon CA, attaining a di amet er 01' 30-35 mm in 14 da ys a t room
u-mp crature (20-25 OC), co ns isti n g 01' a similarly tou gh, close-tex tu re d basal
rc'll , with colo ny surface r emaining persistently white ; velve ty , stro ng ly and
II<-cpl y radiately furrowed , with ce n tre rai sed and irregul arly folded in a
rr. uc rifor rn pattern ; eve n ly, undulatin g m argin , surrounded by a narrow
rl'ingc 01' submerged hyphae ex te n d ing 1 mm beyond th e ae r ial growth; po orl y
spo ri ng ; ex u d a te and odour la cking ; colo ny reverse (colour pl ate 43, b) in de ep
snlmo n-pin k to cinn a m o n shades; no coloured pi gm ent diffusing into the
surro u nd in g aga r; penicilli essen tially as d escribed aboye .
Colon ies upon MEA (co lou r pl at e 43, C and e) growin g restrictedly ,
a lla in ing a diamet er 01' 30 mm in 14 days a t ro om temperature (20-2 5 OC ),
ru ns isting 01' a d ense network 01' trailing a n d close ly interwov en ae r ial hypha e
rove ring th e e n tire colon y surface, lending it a sublanate a ppearance; thin ,
uu dul ating margin su r rou n ded by a n a rrow fringe 01' su b merged hyphae
r-xtend ing 1 mm b eyond th e ae r ia l gro wth; sp oring abundantly throu ghout in
pale blue- gra y sh ades (S guy , PI. XXX , 445-450); ex u date a n d odo ur
l.icking ; n o co lou re d pi gment diffusing into th e surrou nd ing a gar; colony
rcverse (col our pl at e 43, c) in cin namon shades. P enicilli essent ia lly as
dcscrib ed above .
Penicillium gerundense a p pears m ost close ly rel ated with sp ecies in cluded
within th e P. decumbens ser ies on account 01' th e sho rt stipes, com mo nly a rising
Irorn trailing hyphae , th e velu tinous a p pearance a nd slow growth 01' its colo n ies
lHI most su bs trata .
The species di a gn osis is ce n tere d on th e typ e strain IJFM 5967 = ATCC 42,234,
isolat ed from th e ai r, Madrid , Sp ain, 1977 . It is a t present kn own o nly fro m
th e typ e strain .

44. Penicillium alicantinum R amrez et M artne z : fig . 44 , co lo u r pl ate 44 ,

and SEM pl at e 8 , (ta p: right) .
Penici//ium alicantinum Ramrez et Martnez - Mycopathologia. 72: 181-191, 1980.
C olon ies upon C A (colour pl ate 44, A a nd a ) gro wing m oderately , attain ing
di amet er 01' 35- 45 mm in 14 days a t room temperature (20- 25 OC ),
co n sistin g 01' a tou gh , close -textu red b as al felt , radiately furrow ed , with th e
l

l.lJ.J.l.1.JJ..

10 jsrn

0000

0 0 0
00

00 0 0

0 0 0

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L-..-.....J

100

jrrn

at

Fig. 44. Camera lucida

drawings of Penicillium
alicantinum Ramrez et
Martnez, ATCC 42,236. A,
Detailed drawings of the
penicilli. B, Mature conidia .
C, Habit sketches .

ce ntral area rai sed in a cra te riform pattern ; vel ve ty , with eve nly , white n arrow
m argin , 1 mm wide , surrounded by a narrow frin ge of su b me rged hyphae
ex te nd ing a bo u t 1 mm beyond th e aerial gro wth; colo ny surface at first white ,
lat er becoming yellow ish in non-sporin g areas and deep blue- gr een (S gu y, PI.
XXX, 44 6, 447) in marginal areas wh ere co nid ial structu re s a re abundantly
produ ced ; ex udate ab u ndan t to limited in a mo u n t, in yellow d ropl ets ; lacking
od ou r; colo ny rev erse (colour plate 44, a) in bri ght orange shades ; yellowora nge-co lo u re d pi gm ent diffusing into th e surro u nd ing agar; penicilli strictly
sim ple , with sho r t stipes, less th an 50 p.m in length by 1.5-2.0 p.m in di amet er,
smooth wa lled, with a p ices not enla rged in a ves icle , usu all y arising fro m
trailing hyphae; phialides bottle shape d, few in number , 2- 3 in th e verticil
(fig. 44, A), usually 6.0 -11 .0 p.m b y 2.0 - 3.0 tuu, terminating in a sho rt ,
tap er ed neck ; co nid ia globos e to subg lobose , smooth wa lled or nearl y so (S EM
plate 8 , top : right), m ostl y 2.5- 3.0 utt: in di amet er; con id ial chains forming
short, di vergent colu m ns .
Colon ies up on C YA (co lour plat e 44, B a nd b) a tta ining a di amet er of
45 mm in 14 d ays at ro om temper ature , du plicating in texture a nd cultu ral
charac teristics th ose grown up on C A , but sulcated with numerous and deep er

1:11

rud ial Iurrow s , co nspicuously zona tc nnd henvier spo riug , es peci ally in
ma rg inal a rcas in dcep blu e-g rccn shad<:s (S guy, PI. XXX , 447 ); cxud a tc
ubu ndan tly produccd in ycllow -orangc dropl cts; odou r lacking; colo ny reve rso
(colou r plato 44, b) in bri ght yellow -orangc shadc s; bri ght yellow- coloured
piglllent difu sing into th e surrounding ag ar; pcnicilli esse n tia lly as dcscribcd
a boye .
C olonies upon MEA (colour plate 44 , C and c) growin g m ore rapidly than
IIpon th e a boye substrata, attainin g a di amet er 01' 50 mm in 14 days at 1'00111
icmperature (20- 25 OC ), co ns isting 01' a tou gh , c1 ose-t extured basal Ielt ,
vclvet y to sublanate, slightly raised in ce n tra l a reas; thin , eve n margin ,
sur rou nd ed b y a frin ge of submerged hyphae exte nd ing about 3- 4 mm beyond
1he ae ria l growth; colon y surface remaining persistently white, due to th c
mycelial overg ro wth 01' fin e , close ly interwo ven ae ria l hyphae a nd th e pOOl'
sporula tio n up on th is substratum ; colony reverse (colour piat e 44, c) in pale
ycllow-o ra nge shades ; no colo u re d pi gment diffu sin g into th e surro u nd ing
"gar ; penicilli esse ntially a s d escribed aboy e.
Penicillium alicantinum a ppears most closely relat ed to species included within
lite P. decumbens series, on accou nt 01' th e slow-gro wing colonies, th e velv cty
appearance , a nd th e short stipes arising a s short branches from tr ailing
hyph ae .
The species diagn osi s is ce n te re d upon th e typ e strain IJ FM 7026 = AT C C
42,236, isolat ed from th e air, M adrid , Sp ain , 1978. It is at present kn own only
Irorn th e typ e strain.

15 . Penicillium malacaense R amrez el Martnez: fig . 45, colou r plate 45 ,

and SEM plate 8 , (centre : left ) .

Penicillium ma/acaense Ramrez et Martnez - Mycopathologia. 72: 181-191, 1980.
C olon ies upon C A (colour plate 45 , A and a) growing restrictedl y and
slowly , att ainin g a diamet er 01' 20 mm in 14 days a t ro om ternperature
(20- 25 OC ), co nsisting 01' a tou gh, com pact , close-t extured basal felt , velvety
in a p pearance , deeply furrowed in a radial pattern, raised in cent ral areas ;
wh ite, en tire, undulating m argin ; poorly sporing , not influen cing th e colony
a ppe a rance ; colony surface remaining persistently white ; exudate abu ndan tly
produced as topaz yellow, crys tal-clear dropl et s; od our lacking ; colo ny revcrsc
(colour pl ate 45 , a) in cin nam on shades; ambcr-coloured pi gmen t diffusing
slowl y into th e su rrou nd ing a ga r; penicilli strictly sim ple , with very short ,
slend er sti pe s, usuall y less th an 50 JJ-m b y 1.5 - 3.0 JJ-m , mostl y arising as short
branch es from tr ailing hyphae ; phialides (fig . 45 , A) bottle-sh aped , shortpointed , usually 6-10 per verticil , m ostly 7.0-10 .0 JJ-m by 3.0-3 .5 JJ-m;
co nid ia globose, 2.5- 3.0 JJ-m in di amet er, with wall s co ns picuo usly spin ulose

;:::::::-----

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8 (;;,

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'F:'

~r-~JJ$ f.R. .
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.i

Fig. 45. Camera lucida

draw ings of Penicillium
malacaense Ramrez et
Martnez , A TCC 42,241. A,
Detailed drawings of the
penicilli. B, Mature canidia.
C, Habit sketches .

(fig. 45, E , and SE M plate 8, ce ntre : left) ; conid ial chains forming sho rt, more
or less parallel colu m ns (fig. 45, C) .
Colon ies upon C YA (colour plate 45, B and b) a tt ainin g a di a m eter of
25 mm in 14 days a t roo m temperature (20-2 5 OC ), duplicating in tex ture and
cult ural cha racte ristics th ose grown on C A, but with better spo ru la tion; deepl y
radiat ely furrowed in m arginal and su bmarg inal zone s, with centre depressed ;
conid ial areas showing pale pink shades; ex udate abu nda n tly p roduced , in
crysta l c1ear, colourless drop s; odo ur pronounced , ' mould y ' ; colony reverse
(colour plate 45 , b) in reddish brown shades ; no coloured pigment d iffusin g
into the su rro u nd ing agar; penicilli esse ntially as described aboye.
C olonies upon M EA (colour plate 45, C and c) growing ve ry restrictedl y,
a ttai ning a di amet er of 15- 20 mm in 14 days at room temperature
(20- 25 OC ), velvety, plane, looser textured th an aboye; white, eve nly m argin,
su rro u nde d by a fringe of su bmerged hyphae extend ing abo u t 1 mm beyond
th e ae rial gr owth ; heavily spo ring throu gh ou t in deep gray-gre en, olive gree n,
or artem isia gre en shades (Sgu y , PI. XXIX , 421, 426, 428); exudate and
odour lacking; colony reverse (colou r plat e 45, c) in yellowish shades with a
gr een ca st sh owing through ; no coloure d pigment diffusing int o th e

surro und ing ngnr: pcni cilli csscntinlly aN dcscribed nhovc.

Penicillium 7IIa/aCfUm .l'C a ppe ars mo st closcly re lutcd with specics includcd
w ithin th e P. decumbens se ries, on a ccount 01' th e s lo w -g ro w ing co lo n ics o n m ost
su hstr. u a , th c vclve ty a ppeara nce 01' IIIC colo n y su racc a IHI th c sho rt, slcndc r
s tipcs a r ising frorn the tr ailin g hyph ac .
Thc species dia gnosis is ccn tcred u po n I he type stra in IJ FM 70 9 :~ - A 'I'C C
,12,24 1, isolated from th e air , M adrid , Spain , in 19790 It is a l pr csent kn own
onl y from the type stra in.
M embers 01' th e P. decumbens ser ies represent norm al co m po ne nts 01' th c
mycofl ora 01' soils . T hey a re cos mopo lita n in d istribution , a nd uppca r lo
pred ominat e in tr opi cal and sub tro pica l a reas. Since th eir grow th is gcnc rally
so rest rict ed in artificia l cult u res, th ey are ea sily ove rgrow n a nd obscurcd by
more ra pidly grow ing spe cies, hen ce th eir prescn ce and ab u ndance in natu re
Illay be underestimat ed .

Penicillium restricium se r ies

T he Penicillium restrictum se ries wa s ad m itted artificially in co nccp t a nd
design ed to cover certain for ms which co uld no t otherw ise be sati sfuctoril y
placed (Rape r and T ho m 1949). T wo species were inclu de d in thc se ries,
namely : P. restrictum G ilman el Abbott , and P.J uscum (So pp) R aper et Thom , In
I he present study seve n add itio nal species have been included in th e series, as a
matter 01' conven ience , namely : P. arabicum Baghdadi, P. atro-oirens Smith, P.
dim orp hospo rum Swart , P. mont anense Christensen et Hackus, P. raperi Smith , P,
striatisporum Stolk, and Po vasconiae R amrez et M art nez . Al th ough a close
rela tions h ip betw ee n the m is not pr esu med here , th ey do , howev er , sho w
ccrta in chara cte ristics in common, part icul arly the pred ominant floccosc
characte r 01' th e colonies, a nd th e usu all y coarsely roughene d conid ia .
K ey to the species

Page

A . Colonies predominantly floccose with funiculose character Iimited or

lackinq,
Porestrictum series
1. Conidia strongly echinulate or tuberculate, comparatively large , up to
5 p.m or more in diameter.
a. Colonies growing very poorly upon CA, mostly subrnerqed.
Po montanense Christensen et Backus .
b. Colonies growing restrictedly upon CA, attaining a diameter of
20-25 mm, but not submerged .
1'. Two forms of conidia produced: at first elliptical and smooth,
later tuberculate and qlobose.
P. dimorphosporum Swart
o

133

136

148

1:11
2'. Onlv achinulata, globosa conidia producsd .
aa. Colony reverse eolourless upon MEA.

P. fuscum (Sopp) Raper etThom .. . . 141

bb. Colony reverse in deep orange shades upon MEA .
P. vasconiae Ramrez et Martnez
2. Conidia not as above, comparatively small, up to 311m in diameter .
a. Conidia ellipsoid, with 6-9 longitudinal ridges, giving them a striate
appearanee.
P. striatisporum Stolk
b. Conidia not as above, rough-walled.
1'. Colonies growing restrietedly upon CA, attaining a diameter up
to 25 mm .
aa. Phialides eomparatively small , up to 7 11m or less in length.
P. restrictum Gilman et Abbott
bb. Phialides eomparatively large, up to 12 11m or more in length.
1". Penieilli strietly simple; eonidia globose, eonspieuously
roughened.
P. atro-virens Smith
2". Penicilli variable, mostly simple but oeeasionally forming
small verticils of 2-3 metulae, conidia delicately roughened.
P. raperi Smith
2'. Colonies less restricted upon CA , attaining up to 35 mm in diameter, conidia delicately roughened .

.. .. 152

.. .. 150

.. .. 134

.... 139

.... 145

P. arabicum Baghdadi ... . 143

46 . Penicilliu m restrtctum G ilman el Abb ott : fig . 46, co lo u r pl ate 46 , a n d
SEM pl ate 8, (ce n tre : ri ght).
Penicil!ium restrictum Gilman et Abbott - lowa State College J. ScL 1: 297. 1932.
See also Ch. Thom, 'The Penicillia', pp. 176-177. 1930; also Raper and Thom, 'A
Manual of the Penicillia', pp. 223-225. 1949.

Synonym
Citromyces griseus Sopp -

Monographie der Pilzgruppe Penicil!ium mit

besonderer Bereksiehtigung der in Norwegen gefundenen Arten . Videnskapselskapets Skrifter. 1. Mat.-Naturv. Klasse No. 11, pp. 119-120. 1912.

Co lon ies up on CA (colour plat e 46, A and a) gro wing rather restrictedl y ,
attaining a diameter of 25 mm in 14 days at room temperature (20-25 OC),
deeply velvety to floccose in appeara nce, 1 mm or mor e thi ck , con sisting of a
tou gh , close -tex tured basal felt , covered by a thin mycelial ove rg row th of white
slender hyphae, deeply radi ately fur rowed and rai sed in colony centre,

t(;'.~j
<::'i':.:

C>

<f ;

B t) C:>

t;.,:r. :/i
(7. {i.;,'

R .
'I(

e
./
\~i ~
~
100

jrrn

Fig. 40. Camera lucida

drawings of Penicillium
restrictum Gilman et Abbott,
eBS 367.48. A , Detailed
draw ings of the penicill i. B,
Matu re conid ia. e, Habit
sketches.

sIron gly pulling a part th e substratum and wit h colony surface cracked in the
ce ntre in a crateriform pattern; poorly sporing, re maining persistently whitc;
ex uda re and odour la cking; colony reverse (colo ur plate 46 , a) in ligh t yellow
shade s; penicilli str ictly simple (fig . 46 , A), with short stip es, mo stly 25 p.m 01"
lcss in len gth by 1.2 - 1.8 u ra , smooth walled , arising from aerial hyphae ,
occa sionally ir regul arly branched, showing some tim es one or two seco ndary
pcni cilli on the same st ip e; phialides u su ally d ivergen t, in sm all whorls of 6-8
in n umber, co mparatively small, about 5.0 -7 .0 tut: by 1.5- 2. 5 utt, bottle
shap ed , term inating in a short-pointed neck; conidia globose, about
'2 .0- 2.5 p.m in d iamet er with wa lls co nspicuously ro ughened (fig. 46, B, and
SEM pIate 8 , centre : right) ; co nidial chains tangled (fig . 46, C ) .
Colonies u po n CYA (colour pIate 46 , B and b) attaining a diamete r o'
:30 mm in 14 days at room temperature (20 -25 OC ) , dupli catin g in tex tu re
those grown upon CA, but deeply sulcated wit h numero u s rad ial furrows ,
with centre ra ised in a crat eriform pat tern ; u ndulatin g margin en tire; poorly
spo ring, with colony su rface remain ing pe rsisten tly white ; exudat e a bu nd an tly

pr oduced in topaz yellow d ropl ct s; a mhcr coloure d pigm ent dilu sin g inlo th c
su rro u nd ing agar ; co\ony reverso (coIour plat o 16, b) in c in narnon shades;
peni cilli cssc ntially as dcscribed ahoy e .
CoIon ies upon MEA (colour plat e 46 , C and e) gro wing less restrictcdIy than
up on th e aboye su bs trata, attaining a diarnet er of 40 mm in 14 days at room
temperature (20 - 25 OC), deeply floccose, plane , zonat e , with ce ntre urnbo nate ; colony surface cove re d by a my ce1ial ove rgro wth of close1y int erwoven slender aerial h yphae giving a powdery appe arance to th e coIony;
poorly sp or in g through out , with colony su rface remaining persistently white;
thi n m argin surra u nde d by a wide fri nge of su bme rge d hyphae extending
abou t 5 m m beyond th e ae rial gro wth; exudate and odo u r lacking; colony
reverse (coIour plat e 46, e) colourless to pale cream shades; pale arn be rcoloured pigm ent diffusin g into th e surrou nd ing aga r; penicilli esse ntially as
described aboye .
T he spe cies di agn osis is based upon cult ura l stud ies of strain CBS
367.48 = CMI 40 ,2 28= ATCC 1l,257=NRRL 1,748= QM 1,962=IJFM 5038 ,
isolated by P ra f. E .M . Gi Ibe rt , from H onduras soil, in 1937, a nd on G ilman
and Abbott's original description .
Stra ins show ing cultural a nd morphological charactcristics of thi s spec ies a re
frequently isolated fro m so il. (C olour plate 233 presents a representati ve strain
of P. restrictum, describcd by Kinoshita et al. as P. restrictum va r. kuensanii.) T hey
have a tenden cy to become sterile or nearly so afte r several tr ansfers on
arti ficial su bs trata .

47 . Penicillium monlanense C h r iste nsen el Backus: fig. 47 , colour plate 47,

and SEM plat e 8 , (b ottom : left).

Penicillium montanense Christensen et Backus - Mycologia. 54: 573-577. 1962.
C hris te nse n and Backu s' s original di agn osis fol1ows, since thi s ade q uate1y
presents th e species in question :
'On Czapek's agar growth very poor; colonies 2.2-2.4 cm in diameter in 14 days
at room temperature; mycelial mass thin, largely submerged, supporting uniform,
but sparse conidiophore production; conidia deep blackish green, becoming
fuscous; reverse uncoloured; exudate lacking; odour not distinctive .
Colonies on malt agar 4.6-5.4 cm in 14 days at room temperature; surface plane,
floccose to felt like , becoming velutinous with sporulation; margin irregular,
peripheral 3-4 mm white; sporulating surface dark olive, becoming brownish black
in age; reverse colourless or pale russet (especially in fresh isolates): no exudate;
odour not distinctive.
Conidiophores arising directly from the substratum and from low aerial hyphae up

2.5 - 4-.0 .UII in clia me tc r, smoo th wall cd , with upices enla rge d in a ves iclc up tu
6.0 Jl.m in di ameter: pcni cilli usu ally strictly sim ple , with a n occasiona l branch
(fig . 47 , A), com pa ra tivcly largc ; phi alidcs bo ttl e shapcd , mostl y in clus tc rs uf
6 - 14 e1eme nts in th e ve rticil, usually 10-1 2 Jl.m by 3.0 - 4. 0 Jl.m , terminati n g
in a sho rt, narrowed , tapered neck ; co nid ia at first subg lobose, becoming
globose, 4.0 -6 .0 Jl.m in d iameter, conspicuo us ly coa rse ly echi n ula te (fig. 4 7, B
and SEM plat e 8, bott om: left ), dark brown coloured; co nid ial chains in
loa se colu m ns 0 1' tan gled (fig . 47 , C).
Colon ies upon C YA (colour plat e 47, B a nd b) grow ing rapidly and
exte nd ing broadl y , attai n ing a diameter 01' 50 mm in 14 days at room
temperature (20- 25 OC), co ns isting a l' a close -text ured basal felt, cove re d by a
m ycelial ove rg rowth 01' loosely interwoven aerial hyphae , lendi ng a lanate
appe arance to th e colony surface; rad iat ely wrinkled, with cen tre irregul arly
fold ed ; thin m argin surro u nde d by a fringe 01' su bmerged hyphae exte nding
wide ly, abou t 4 mm beyond the ae rial growth; sparingly sporing , especially in
marginal a reas, re maining persistentl y wh ite; ex udate and odo ur lacking ;
colony rever se (colour plat e 47 , b) in pale N aple yellow shades (Sguy, PI.
XIV , 200); no coloured pigm ent d iffusi ng int o th e surro u nd ing agar; peni cilli
esse ntially a s described aboye , bu t with longer stipes aris ing directly from th e
subs tra tu m a nd also fro m the net work a l' floccose ae r ial hyphae ; con id ial
cha ins forming m ore 0 1' less com pact colu m n s.
Colonies u pon MEA (colour plate 47, C and c) spreadi ng bro adl y and
lu xu riantl y , attain ing a d ia met er 01' 65 mm in 14 days at room temperature ,
cons isting a l' a looser textured basal feIt, cavered by a mycelial ove rg ro wth 01'
floccose ae rial hyph ae , lending a lana te appeara nce to th e colony; thin and
wide margin, surro u nded by a frin ge a l' su bmerge d hyphae ex tend ing abo ut
5 m m beyond the aerial gro wth; heavily spori ng, espec ially in cen tra l and
su bce nt ral a reas in gray-gree n to brownish gree n sha des (Sguy, PI. XXV ,
373, 37 1) becom ing fusco us in age; odour lacking ; exuda te limited , in
colourless d roplets , enmes he d in the m ycelial felt ; no coloure d pi gm ent
d iffusin g in to the surro u nd ing agar; colony reverse (colour plate 4 7, c) in pale
yellow shades; peni cilli essen tia lly as desc ribe d aboye .
T he species d iag nos is is centered u pon cultu ra l stud ies 01' th e typ e strain CBS
3 10 .63 = C M I 99, 468 = ATCC 14,941 = IJ FM 503 4, isolated from a lod gepole pine
Dou glas-fir ( Pseudotsuga taxifolia) soil sa m ple collected in R avilli Cou nty ,
M ontan a , USA, by D I' M . C hristense n during th e su m me r 01' 1957.
Penicill ium montanense closely resembl es P. fuscu m (Sopp) R aper et T ho m.
H owever , P. montanense di ffer s from P. Juscu m in a number 01' mi cr oscopi c
characteristics, notably in its larger peni cilli , a nd its pOOl' grow th upon CA. O n
th e other h and , despit e its pred ominantly sim ple habit , P. montanense shows an
occasional character 01' the di varicat e peni cilli , which, in combination with the
dark bro wn colou r and th e stro ng ec hin ulation 01' th e conidia, a re rem in isce nt

11 1' P. meliuii T ho rn 01' the P. nigricnns se ries in th e Asytum cu -icn-idi varlcutu subsec tion ( R apc r and T ho m, 1!W J).

1B. l'enicillium airo -oirens Sm ith: lig. 4B, colour pla to 'lB, ami S EM plato
B, (bo llo m : right).

":'' ::.
~.~ \

?i::"'

' ..:.(. {\..,8i'::\

:~..;:~

L.wWwJ

10

).1m

tLttvif
l.-..-...J
).1m

100

Fig. 48. Camera luc ida

drawings of Penicillium strovirens Sm ith, CSS 326.59.
A , Detailed drawings of the
penicilli . S, Mature conidia.
C, Habit sketches .

Penicillium etro-virens Smith - Trans. Brit. Mycol. Soc o46: 331-337. 1963. See also
M.M. Kulik , ' A Compilation of Descript ions of New Penicillium Species' . Agr ic.
Handbook No . 351 . Agric . Res. Ser. U.S. Dep. Agric . p. 19. 1968.
Synonym

Penicillium decumbens Thom, varo atro -virens Abe - J . Gen. Appl. Microblol .,
Tokyo. 2: 70-71. 1956 (without a Latin diagnosis).

Colonies upon C A (colour pIate 48 , A and a) growing slowl y , attaining a

diameter of 30 mm in 14 days at room temperature (20 - 25 OC), cons isting of a
close -text u re d basal felt , velvety in appeara nce, deep ly rad iat ely wrinkled in

marginal ami submarginnl area s, but usually hcapcd in (he centre in a

cra te riform patt crn: undulating murgin cntirc , surroundcd by a IlUTOW fringc
01' submcrgcd hyphae cxte ncling lcss than 1 mm bcyond th e aerial growth;
strongly pulling apart and cracking the substratum underneath (colour plate 48,
a); heavily sporing in dull blue-green shades (Sguy, PI. XXXI, 461) ; exudate
and odour lacking; no coloured pigment diffusing into the surrounding agar;
colony reverse (colour plate 48 , a) colourless ; penicilli strictly simple but with
an occasional branch (fig . 48 , A), with th e stipes of th e conid iophores
com paratively long, up to 250 Jlm in length by 1.5- 2.5 Jlm in diamet er , with
wall s apparently smooth to deli cately roughen ed, with api ces slightly enla rged
in a vesicle up to 4.O Jlm in di ameter, arising either from th e substratum 01'
from trailing hyphae; phialides bottle shaped, mostly 9. O-13 .O Jlm by
2.5-4.0 iun , terminating in a short , narrowed neck ; con id ia ovoid ,
conspicuou sly roughened (fig. 48 , B, a nd SEM plate 8, bottom : right); con id ial
cha ins forming parallel to somewhat div ergent columns (fig . 48 , C).
Colonies upon CYA (colour plate 48, B a nd b) attaining a diamet er 01'
30 mm in 14 day s at room temperature (20 -25 OC ) , con sisting of a closetextured basal felt , velvety, strongly sulcate with numerous de ep radiate
furrows , and with th e centre heaped in a crate riform pattern ; poorl y sporing,
probably due to an excess 01' vitamins in th e substratum, with the colony
surface remaining persistently white ; undulating margin entire, with a narrow
fringe 01' submerged h yphae exte nd ing lcss than 1 mm beyond th e aerial
growth ; exudate and colour lacking; arnber-coloured pigment diffusing into the
surrounding a ga r; colony revers e (colour plat e 48 , b) in pale cre am shades;
penicilli essentially as described aboye.
Colonies upon MEA (colour plate 48 , C and c) attaining a diameter 01'
30 mm in 14 days at room temperature, con sisting 01' a c1ose-t extured ba sal
felt, plane , velv ety , with an evenly white margin, surrounded by a fringe 01'
submerged hyphae extend in g about 2 01' 3 mm beyond th e aerial growth ;
heavily sporing throughout in deep blue-green sh ad es (S guy, PI. XXX, 446) ;
exudate lacking ; odour faint, fragrant, aromatic; no colou re d pigment
diffusing into th e surrounding agar; colon y revers e (colour plate 48 , c)
colourless ; penicilli as described aboye , but with th e stipes mostly arising frorn
th e basal felt, and some from trailing hyphae ; conid ial ch ains forrning welldefined, com pact colu m ns.
The spe cies di agnosis is centered upon th e typ e strain C BS 326.59 = CMI
68,223 = ATCC 18,225 = IFO 6,228 = QM 7,307 = IJFM 5012 , isolated by S . Ab e
from Japanese soil in 1955, and des cribed by him as P. decumbens Thom, val'.
atro-virens Abe (without a Latin diagnosis) (1956) .
A close relationship between P. atro-virens Smith and th e P. restrictum series is
not presumed here. However, the y are conside red together , primarily as a
matter 01' conveni en ce. It shows certain affinities with th e P. decumbens seri es,

nnmcl y th c slow -growin g co lo n ics , IIH~ vclv cty a p pt:a ra nce , a m i lh e sripc

nrising

1'1'0111

lrailing hyph ae.

Penicilliumfuscum (Sopp) Rape!' el Thom: Iig. 49 , colour plate 49,

a nd SEM plate 9 (top: lcft) .

l!) .

~;9$_'~
QJ!
t t1J f':-
t::;p

(@

~ ~~ (~g' : '@ 'j

{J;

u.w.1.w.Ll
10 jirn

.K

Ji1
~

100 jrrn

Penicil/ium fuscum (Soppl Raper et Thom 226-227. 1949.

Fig . 49. Camera lucida

drawings of Penicillium
fuscum (Sopp) Raper et
Thom, CSS 147.45. A,
Detailed drawings of the
penicilli. S, Mature conidia .
e, Habit sketches .

'A Manual of the Penicillia', pp.

Synonym
Citromyces fuscus Sopp - In Monographie der Pilzgruppe Penicil/ium mit
besonderer Bercksichtigung der in Norwegen gefundenen Arten. Videnskapselskapets Skrifter. 1. Mat.-Naturv. Klasse No . 11 . pp. 120-122. 1912. See also
Ch. Thom, 'The Penicillia', p. 180. 1930.

C olonies upon CA (colour plate 49 , A and a) growing slowly , att am mg a

diameter 01' 30 mm in 14 days at room temperature (20-25 OC ), cushion like ,
1-2 mm deep , spongy 01' fleshy , with colony surface 01't en appe arin g wet in

locali zcd a rc a s , showin g' sh allow radi al lurrows in narrow sub margina l arca s,
co ve red by a nctwork 01' stc r ilc ac ria l hyph ac , sho wing' sorn e ro pinc ss in ce n tra l
a reas; thin rnargin su rroundcd by a [rin gc 01' suhmcrged h ypha c cx ie nd ing
a bou t 1-2 mm bey ond th e acrial gro w th ; po orly spo r ing, with co lo ny su rface
remaining per sist entIy w h ite in strain s long maint ained under artifici al
cu lt ivation, but m edium spo r ing throu gh out , hea vier in m arginal a nd su b m arginal a reas, a t fir st olive gray (S guy, PI. XXIX , 427), becoming d eep
gra yish to b rown (Sguy, PI. XXIX , 433) in a ge , in recen t isol at e s; exud a te
and od our la ckin g; pale amber pi gment slo wly diffusing into th e surro u nd in g
a gar ; colo ny reverse (c olo ur pl ate 49, a ) in p al e cream sh ade s; penicilli var ia ble
(fig . 49, A ) , ran gin g fro rn fra gmentary to str ictIy sim ple to irregularly
branch ed (s uggesting th e Asymmetrica-divaricat a ), with two 01' m ore m etulalike cells borne terminall y 0 1' a t low er se p ta, with th e stipes o f th e co n id iop hores
variable , a r isin g m ostI y fro m ae ri a l h ypha e a nd u su all y not exceedi n g
100 tm in length b y 1.5- 2.5 tm in di amet er, se p ta te, with wall s smooth 01'
deli cately rou gh en ed ; phi alides di vergent , in small clusters up to 6 e1ements in
th e ve rticil, frequentI y 2- 3, m ostIy 7-1 3 tm by 2.5 -3 .0 tui , te rm in a ting in a
co m parative ly lon g , narrowed , tapered neck ; co n id ia globose, a t first co lo u rle ss , becoming olive brown when m ature , co a rsely ech in u la te with prominent
dark tu bercle s (fig . 49, B, a n d SE M pl ate 9, top : left ) , m ostIy 3.5-5. 0 tutv , 0 1'
up to 7.0 tm in di amet er; con id ia l chains tangled up to 50- 100 tm in length
(fig. 49, C).
Colo nies upon C YA (colour pl at e 49, B and b ) gro w ing m ore r apidly than
upon CA , a tta in in g a di amet er al' 30 -35 mm in 14 days a t room temperature
(20- 25 OC), similar in texture and cu ltu ral ch aract eristi cs to th ose grown upon
C A, but with ropiness m ore ev ide n t th roughout , plane , 1 mm deep in th e
ce n tre; thin m argin surrounded by a fringe a l' su bmerged hypha e ex te nd in g
about 1 mm beyond th e ae r ia l growth; very poorly spor ing, with su rface
r emaining persist ently white ; odo ur and exu date lacking; pale a m be r -co lo u re d
pi gment diffusing into th e su rrou n d ing aga r ; co lo ny reverse (co lo u r pl at e 49,
b) in pale crea m shades; penicilli as de scribed a boye.
C olo n ies upon M EA (co lour pl at e 49, C and c) sp read in g broadly , a ttain ing
a diamet er al' 55 mm in 14 d a ys a t room te m pe rature (20- 25 OC ), looser in
texture , co m parative ly thin , vel ve ty in a ppearance, with co lony su r face
charac te r ize d b y tr ailin g a nd interwoven ae r ia l h yphae; th in, eve nly m argin ,
surrou nded by a wide frin ge a l' sub merged hyphae ex te nd ing m ore than
10 mm beyond th e aeria l growth ; heavily spor in g throughou t in d ark mouse
gray shades (Sguy, PI. XXXV , 513) 0 1' oli ve- green co lours (Sguy, PI. XXI,
301); ex udate a n d odour la cking ; n o colo ured pi gment diffusing in to th e
surro u nd in g agar; colony rev erse (colour pIate 49, e) in reddish brown to olive
gre en shades; penicilli essentiall y as described aboye but with the stipes al' the
co n id iopho re s somewhat longer th an upon th e a boye substrata a nd m ore

III 'qu elllly b ranchcd in th e mnnn cr 01' thc Asyuu uctri cn-edivnricnru.
spc c ics di a gnosis is cc n tc rcd II pon 1he sI rui liS e li S 147.45 :1:J1.4B = C M I 39, 747 = A T C C 10,447 - NI~I{ 7'21 .. 11"0 6, 09 7 ..

' 1'111:

Ind (: IIS

IJ FM
QM

:)0 15
1,92 5 ,

Hnlll l ecl rom T exas soil by Pr of. M .B . Morrow, frorn th e Univcrsity o f T exas,
11

111:10 .
'1'111: co r rcc t placern ent of thi s species is still op en to question , T he

IlI Hi~ II I1 I C II I lo

its present position , according to R aper and Thoms opinion

11:)11 11), is bas ed upon th e belief th at it ca n be keyed her e m ore readily than
t1IHC'wlll:l'e , altho ug h its closer relat ionship to th e P. nigricans seri es in thc
A"Y" lllletrica - d iva ricata su b-sec tion should not be overl ooked on account 01'
11 11' pn:sell cc of sorne di vari catel y branch ed stipes, espec ially upon M EA.

n. Prnicillium

ara bicum Baghdadi : fig . 50, calaur pl ate 50 , a n d SEM

plat e 9, (top: right ) .

(1))

0
00
O O
O O O OO
00 Bo 0 0
O O

0 0

lJ..wL.u.J.J

10 J.Im

{j
f"

.!

li(:.--/"
.-

~-

-t

100 }-1m

fR .

Fig. 50. Camera lucida

drawings of Penicillium
arabicum Saghdadi, CSS
414.69. A, Detailed drawings
of the penicilli. S, Mature
conidia . e, Habit sketches.

'enicil!ium arabicum Baghdadi - Nov. Syst. Niz. Rast. pp . 105-106. 1968.

..-,
Baghdadi's original diagnos is is as follows :
'Colonies upon Czapek's agar atta ining a diameter of 40 mm in 12 days , closs
textured , w rinkled, velvety to floccose , w ith c1avate, closely interwoven hyphae in
marginal areas, reminding c1eistothecia initials; nonsporulating areas in dirty white
shades; sporulating areas and reverse in gray-green colou rs; exudate limited in
amount, colourless or lacking.
The conidiophores arise from aerial hyphae, smooth walled or nearly so, 23-35
.tm by 3.3 .tm , sometimes up to 150 .tm in length; with apices enlarged in a vesicle
up to 3.5 .tm in diamete r; ph ialides bottle shaped, in c1usters numbering 6-8
elements in the verticil, 6-8 .tm by 3 .tm. Conidia globose, delicately roughened , 3
.tm in diameter.
Colonies upon malt extract agar attaining a diameter of 40 mm , close textured ,
plane, velvety, in lead gray shades, with centre white; colony reverse in darker
shades tha n upon Czapek's agar; heavily sporing throughout the colony surface .
Colonies upon corn meal agar attaining a diameter of 40 mm, c1ose-textured ,
velvety, wrinkled, in blue-g ray shades; exudate abundantly produced , in pale yellow
shades . The species is assigned to t he Monoverticillata stricta section. No closelv
related species are known . Several strains have been isolated from Syrian soil in the
environs of lrna and Darai, Damascus region , and in the environs of Es-Suveida , EsSuveida region (Svr ia), in August 1964, by the author.
The type culture T 16 has been deposited w ith the Cult ure Collection of the M.B.
Lomonossov University, Moscow, USSR , and with the Culture Collection of the
University of Syria , Damascus .'
(Translated from the Russian by Prof. L. Vilas).

Our notes follow:

Colonies upon CA (colour plate 50, A and a) attaining a diameter of 45 mm
in 14 days at room temperature (20 - 25 OC ), cons isting of a tough , leathery,
close -textured basal felt, velvety in marginal areas, surrou nded by a narrow
fringe of submerged hyphae extend ing about 1 mm beyond th e ae rial gro wth ;
sulcated with few, shaIlow, radial furrows , ot herwise almost plane in ce n tral
areas; poorIy sporing, with the colony surface remaining persistently white or
wh ite rosy ; exud are colou rle ss, in droplets en m eshe d in the mycelial felt ; pale
amber-coloured pigment diffusing into th e surrounding agar; odour lacking ;
when the K 2HP0 4 of the Czapek's medium is substituted by KH 2P0 4 , a
strongly fluore scen t yellow pigment is th en produc ed , diffusing into th e
surrounding agar; colon y reverse (colour plate 50, a) in pale cream shades;
penicilli strictly simple (fig . 50, A) , with th e stipes of th e conid iophore s short ,
ranging from 25 to 75 /lm in length b y 2.0-2 .8/lm in diameter, sligh tly
enlarged in th e apex in a vesicle u p to 4.0 /lm in diameter, wit h wa lls sm ooth or
nearly so ; phialides bottle shaped , sometimes in flated up to 7.0 /lm in width, in
clusters of 6-8 elements in the verticil, mostly 7.0- 13.0 /lm by 2.5-4.0 /lm ;
conidia ellip tical , subglobose to globose, delicately ro u gh ened (fig . 50, B, and

" I': M plat o 9, top : righ t), 2.5 - 1 .0 Lm by 2.0 - :I.:J Lm; conid ial cha ins shor t,
1/lII Io(lt:d (lig. 50, C ).
:lllon ics upon CY A (co lo u r pla to 50, B an d b) attain ing a diamc ter 0 1'
I ~ 11 \111 in 14 days a t room temperature (20 -25 OC ), consisting 0 1' a leathery
1II'1'11I:e, su lcate d with nu m erous, radial fu rrow s, and dcprcsse d in the centre;
th ln ruargin su rro u nded b y a frin ge 01' su b mc rged h yphae cxte nd ing 2- 3 mm
buyond th e ae r ial gr ow th ; colo ny su rfaee eovered with a thin mycc1i al
uvcr growth 01' white ae rial, closely interwoven , hyphae giving a su b la n a te
\pl'eal'a nec to th e eolony; med iu m sporing in pale blue-gra y shades (Sg uy, PI.
X X II I, 48 4), with n on -sp orulating a reas remainin g p ersistently white ;
xud.uc in eo lou rless d roplets; odou r laeki n g; ye llow -eoloured pi gmcnt
d fl'nsillg into th e surro u nding aga r ; eolo ny reverse (eolour plat e 50, b ) in
i-Ilow -cream shades; penieilli essentially as deseribed a boye, w ith th e stipcs 0 1'
11\1: con id iop hores mostly arising from th e net work 01' ae rial hyphae.
( :Dlon ies upon MEA (eo lour pla te 50, C a n d e) a ttaini ng a d iametcr of
I ~) 111m in 14 d ays at ro om te m pe rature (20- 25 OC), eonsisting 01' a typi cal1y
dC'('l' ly velvety, loose-t extured b asal felt, zo nate, plane , with th in margin ,
IIITDI!nded by a fringe 01' su b merge d hyphae exte nd ing abou t 2 - 3 mm bey ond
1111' ucrial growth; heavily sporing through out in blue-green shades (Sgu y , PI.
X X XI , 46 1); exudate limited in a mou nt, en meshed in the m yeeli al felt ; odo u r
Inrk iIIg; pale amber p igment d iffu sing into th e surro u n d ing agar; co lony
n -vcrsc (co lou r pla te 50, e) in yello wis h eo lou rs; penieil1i essen tial1y as
I h'sni bed a boye, bu t w ith the sti pes either a r ising from the su b st ratu m a nd
1'1'Il1I1 the ne twork 01' ae r ial hyphae ; co nid ial eh a ins in loosely paral1el colu m na
DI' lan gled .
' I' he speeies deseription is ee ntered u pon cu lt ural stu dies 01' th e typ e strain
1:Il S 11 4.69 = V K M F-I 0n = C M I 140 ,335 = ATC C 22 ,347 = IJ FM 50 14, isolatcd
I'I'DIl\ Syrian d esert soil by V . C h . Ba ghdadi , in 1964.
' I'he correet plaeement 01' the speeies is still ope n to question. It is as signed
1)(:1'(: tentat ively only as a m a tt er 01' eonven ienee .

.11 . Penicillium rapen' Sm ith : fig. 51, colour plate 51, a nd SEM plat e 9 ,
(centre: Ieft ).
Penicil/ium raperi Smith - Trans . Brit . Myeol. Soe. 40: 486-487. 1957. See also
M .M . Kulik, 'A Compilation of Deseriptions of New Penicillium Species' . Agrie.
Handbook No. 351. Agrie. Res. Ser. U.S. Dep. Agrie . p. 70. 1968.
Smith's original diagnosis is as follows:
'On Czapek agar grows restrietedly, 22-25 mm in 9-10 davs, and thereafter
spreading only slightly, matted floeeose about 1 mm thick, at first pale yellow
beeoming mauve in parches. tardily turning pale bluish green all over or almost all

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Fig. 51. Camera lucida

drawings of Penicillium
raperi Smith, CBS 281.58.
A, Detailed drawings of the
penicilli. B, Mature canid ia.
e, Habit sketches.

over, with drops small, colourless, and reverse dull brown with intense yellow edge,
but with no diffusion of the pigment into the agar, sporing tardily first in the mauve
areas, more freely when the greenish colour appears; on malt agar grows rather
more quickly, 30 mm in 9 days and 42-45 mm in 15 days, pale yellow and whitish
with only traces of pink or mauve and turning greenish only very slowly and
incompletely, with reverse deep dull yellowish brown; on potato-dextrose agar
grows rather more quickly than on Czapek, much as on malt agar, yellow at first
then with mauve centre but with a middle zone of pale bluish green, with reverse at
first brown w ith yellow margin, then with centre purple and margin zoned with
brown and deep greenish yellow; penicilli monoverticillate or. more often, irregular,
with two to three unequal metulae, or with various superimposed verticils
consisting of both metulae and phialides, but with the number of elements at each
stage very small; conidiophores arising from aerial hyphae, very short, mostly less
than 25 Jlm long and 1.3 to 1.5 Jlm in diameter; phialides few in the verticil, usually
divergent, 7-9 by 1.8~2.2 Jlm, narrowed at the base and with fairly long tip; conidia

Irrogularly ovate, sllghtly rough, 3-3.6 by 2.4-2.8 Lm, many appearing to be

lobose or subglobose, but these probably ovate conidia sean end-on; conidial
hains short, divergent or tangled; numerous sterile branches produced from aerial
hyphae, sometimes short and spur-like, sometimes longer, more or less clubheped and frequently nodulose at the tip, resembling abortive conidiophores.'

Sm iths original dcscription 01' the species is given sin ce this ad equatcl y
prcscnts th e spec ies in question whcn first isolat cd . The typc strain OIlS
" 11 1.58 = CMI 71,625 = IF O 8 ,179 = QM 7527 = IJFM 5007 , maintained in a rtificial
cultu re sin ce its isolation from th e soil of a cabbage field near Bedford, in
Septernber 1952, has become predominantIy poorly spor ing , rernaining pcrMistently white upon C A and CYA, not showing any longer th e typical cultura l
rh aracteri sti cs origina lly described by th e author, although still sho wing
cha racteristic det ails of morphology.
O ur not es follow :
Colonies u pon C A (colour plat e 51, A a nd a) a ttaining a di ameter of 35 mm
in 14 days at room temperature (20-2 5 OC ), ca ns isting a l' a close -textured
hnsal felt , velve ty to lanat e; poorly spo ring , remaining per sistentIy white ;
l'olony surface deepl y radiat ely furrowed and depresscd in th e cen tre; evcnly
IlIargin su rro u nde d by a frin ge of submerged h yphac exte nd ing about 1 mm
hcyond th e aerial gro wth ; od our and exudate lacking; pale am ber colou red
I,i).{rnent d iffusin g into th e su rro u nd ing agar ; colony reverse (colou r plat e 51, a)
in sulphur yellow shades; penicilli eithe r strictIy sim ple 01' irregul arly branched
i 11 a divaricat e manner (fig. 5 1, A) , with sti pes ar ising m ostl y as short branch es
I'I'OI!l ae rial hyphae , ranging fro m 25 to 100 .tm in length by 2.3 - 3 .0 uit: in
rlinme te r, with walls smooth, enla rged at th e apex in a vesicle up to 3. 5 .tm in
di.unet er; phialides in div ergent c1uster s of 6- 10 e1ements in th e verticil ,
iu ostl y 8 - 11 .tm by 3.0-3 .5 uui , terminating in conspicuo us, narrowed ne ck;
conidia elliptica l to su bglobose , 2. 5- 3. 3 .tm by 2.5-3. 0 uiu, delicately
roug he ne d (fig. 51, B, a nd SE M piate 9, ce nt re: left) ; con id ial chains loose to
tan gled (fig. 51, C ).
Colonies u pon C YA (colou r plate 51, B and b) attaining a di ameter a l'
;)0 mm in 14 days at roo m temperature (20- 25 OC), sim ila r in texture to th ose
gl'Own upon CA , bu t deep er radiat ely furrowed, with thin , yellow margin
sur rou nded by a fringe of su bme rged hyphae exte nd ing abo ut 1 mm be yond
lile ae rial grow th; stro ngly pulling and crac king th e agar underneath (colour
pla te 51, b); poorl y sporing wi th surface rem aining per sistentI y white with
I race s of pink ; exudate abu ndantIy produced in pink shades ; od our lacking;
hrown colou re d pigm ent d iffu sing into th e surroundin g agar; colony revers e
(colou r piate 51, b) in dark brown-purple colours; peni cilli esse ntially as
dcscribed abo ye , with stipes arising from th e net work 01' ae rial hyphae .

C olonics up on M EA (colou r plate 51, C and e) uua in ing a d ia mc rcr 01'

50 mm, co ns isting of a close-textu rcd hasal Ielt , pr cdominan tly la nat c , with
th in margin , surrou nde d by a frin ge 01' yellow subme rge d hyphae cxtending
ab out 4 mm beyond the ae rial growth, with centre urnbon at e; hcavily sporing
th roughout in pale gray shade s (Sgu y , PI. XXX , 445 ); exu dat c a nd odour
lacking ; yellow-gree n coloured pigm ent diffu sin g into the su rru u nd ing agar;
colony reverse (colour plat e 51, e) in greenish yellow shades ; peni cilli
essentially as described aboye, but with con idial chains mo stly in loosely
parallel colu m ns .
Penicillium raperi Sm ith is mo re closely relat ed to th e P. restrictum series on
acco u nt 01' its floccose colonies, with stipes a rising primarily fro m ae rial
hyphae. H owever, th e presen ce 01' divaricat e peni cilli , suggest ing affinities
with th e P. janthinellum series in th e Asymmetrica-divaricat a subsectio n sho uld
not be ove rlooked .

52, Penicillium dimorphosporum Swart: fig. 52, colour plate 52 , and SEM

plate 9 , (centre: right).

y ::-O 00
00 0

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Fig. 52. Camera lucida

drawings of Penicillium
dimorphosporum Swart,
CBS 456.70. A , Detailed
drawings of the penicilli. B,
Mature conidia (srnoothwalled). e, Spinulose
conidia. D, Habit sketches.

IIlci/lium dimorphosporum Swart - Trans Brlt. Mycol. Soco 55: 310-313. 1970.
S WHI'I 's

ori gin al dia gn osis

01' thc

spccies is givcn bclow sin cc thi s ade q ua tcly

pl'l':sc:n ls thc spccics in qucs tio n when first isolat ed . The typ e stra in CIIS 456 .70

tloes not show an y longer the typical cultu ra l cha rac teristics o rig ina lly
tlUMl'I'ihecl by th e a utho r , remaining persisten tly wh ite up on a11 media test cd ,
dlhlJlI /{h still sho wing chara ctcristic dctails 01' morphology.
' Colonies upon Czapek's agar slow-growing, reaching a diameter of 2 cm in 14
va. Colonies white at first , turning pale grayish green in the centre with the
lovolopment of the first conidia . Eventually the culture turns a deep reddish brown
xcopt for the white margino Mature colonies are slightly zonate, irregularly
buckled , with agar sometimes being pulled away from the bottom of the culture
lah. Exudate is abundantly produced , pale pink at first but soon turning deep red;
vorse in similar colours with pigment diffusing into the surrounding medium.
onidiophores short, srnooth, unbranched, arising as side branches from trailing
hyph ae at, or slightly aboye, the surface of the mycelial felt, rarely more than 30 11m
lonu by 2 Jlm wide. Phialides variable, in older cultures short, wide and thus
rllvorging . First-formed conidia are oval , srnooth, greenish in culture and measure
- 2.4 Jlm by 1.8-2 Jlm, but conidia produced later, from the moment the culture
turns brown, are globose, roughened by prominent tubercles and bars of pigment,
brownish, and average 3.1 Jlm in diameter with some conidia up to or even over 4
1,m . Some conidia are intermediate in size, shape, and surface, between these two
mnin types.
Colonies on malt extract agar grow slightly faster, produce more greenish conidia
md are slower to change colour . Pigmentation is not as intense and colonies are
browner with very little red at first . Exudate is limited and tinted amber or pale pink .
ccasionally a few patches of yellow encrusted hyphae are produced. The reverse
In yellow brown. Whereas on Czapek's agar the second type of conidium soon
dominares. on rnalt extract agar a considerable number of conidia of the first type
rre found . Intermediate conidia may be numerous as well.'

O u r notes follow :
Colonies upon CA (colour piat e 52 , A a nd a) a tta in ing a di ameter 01' 32 mm
in 14 days a t roo m temperature (20 -25 OC ), co ns isting 01' a close-textu re d
basal felt , irregul arl y folde d a nd radi at ely wrinkled a nd rai sed in th e centre ;
irregularly dissected margin surrounded by a frin ge 01' sub me rge d hyph ac
cx tend ing 3-4 m m beyond th e aerial growth ; heavily spo ring thro ug ho u t,
with colony surface rem aining persistently white ; ex udate lacking; odo u r fain t ,
'mouldy; no coloured pi gm ent diffus ing into th e surrou nding agar; colony
reverse (colour plat e 52, a) in crea m sha des; peni cilli strictly sim ple (fig. 52 ,
A), with sho rt stipes, abou t 25 11m in len gth by 1.8- 2.5 11m in diamet er ,

smoorh-wa llcd, arising Irom aeriul hyph ae: phi nlidcs bottl e-xhap ed , in clus tcrs
o l' 6- 10 elc mc n ts in th c vc r tic il, mostl y 11 lO I~III by :U l - 4,.:{ Jtlll, tcrmin a tcd
in a short-poin te d n cck ; co n id ia 01' two typ cs: The fir st type (Iig. 52, B) are
ellip tical , sm ooth walled , m ostl y 3.0 - 4 .5 Jtrn in lon g axis . The seco nd type are

globose, co nspicuo usly ech in ulate a nd brown colou rcd (lig. 52, e, a nd SEM
piate 9, cen tre : ri gh t), m ostl y 4.5 -5 .0 Jtm in di ametcr ; co nid ial chai ns sho rt ,
loosely parallel (lig . 52, D ).
Colo nies up on CYA (co lour plat e 52, B a nd b) attaini ng a d iameter of
37 mm in 14 d ays at room te rnpe ra ture (20 -25 OC), co nsis ti ng of a tough ,
close- te xtu re d basal felt, partiall y cove re d b y a thi n rnyceli al overgrowth,
giving a powdery appeara nce to th e colo ny ; deepl y rad ia tely wrin kled a nd
sulcate d conce ntr ically in th e cen tre which is irregula rIy folde d and lea th ery ;
poorI y spo ring, with su rface remaining persistently white; ex udate limited in
amo u nt; odo u r lacking ; pal e a mbe r-co lou red pi gm ent diffusing into the
surro u nd ing agar; colon y reverse (co lour plat e 52, b) in yellowi sh cream
shades; pe n icilli essc ntially as desc ribe d a boye.
Colonies upon MEA (colour plat e 52, C a nd c) attain ing a diame te r of
45 mm in 14 days at room tempera ture (20-25 OC ), consis ting of a looser
textured ba sal felt , with a velvety and powd ery a p peara nce, plane in
sub marg inal area and irregul arI y folde d and ra ised in th e ce nt re; thin,
irregul arI y di ssected rnargin , su rrou nde d b y a fringe 01' submerged hyphae
exte nd ing 10 01' more m m beyond th e aeria l growth; heavily spo rin g
throu gh ou t in pale gray shades (Sguy, PI. XXXV, 515); ex udate lacking;
odo ur faint, arom atic; no coloured pi gm ent diffu sin g into th e surrou nd ing
agar; colony reverse (colou r plate 52, c) uncoloured , bu t wi th ce n tre sho wing
th ro ugh green- blac k shades; penicilli esse ntially as described above b u t with
stipes aris ing directl y from sub me rge d h yphae in a radial pat tern ;
The species description is cen te re d u pon th e type strain CBS 456.70 = CMI
149,680 = ATCC 22783 = IJ FM 502 1, isolated fro m a m angrove swa m p soil near
W esternport Bay near M elbourne , V ictoria, Au stralia , by H .I. Swa rt, in

1969.
The co rrect placement 01' th e speeies a ppears to be m ost sa tisfac torily
ass igned to th e P. restnctum se ries on accou nt 01' th e slow growth on most m edi a,
th e sho rt stipes arising from ae rial hyphae , a nd th e presence 01' strongly
echin ula te globose conid ia.

53 . Penicillium striatisporum Stolk : fig. 53, colour plat e 53, and SEM
plate 9 , (bottom : left ).
Penicillium striatisporum Stolk - Antonie van Leeuwenhoek. 35: 268-270. 1969.

Colonies upon CA (co lour plate 53, A and a) grow ing restri ctedl y , attaining

rt

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ER.

Fig. 53. Camera lucida

drawings of Penicillium
striatisporum Stolk, CBS
705.68. A , Detailed drawings
of the penicilli. B, Mature
conidia. C, Habit sketches.

rli.uuc ter of 28 mm in 14 days at room temperature (20- 25 OC), consisting 0 1'

ICI\l gh , close-textured basal felt , abo u t 3 mm deep in th e colo ny ce n tre,
IZCl llate, not wrinkled , covere d b y a closely woven network of hyphae giving a
111)('('ose a p pe ara nce to th e colo ny su rface, white rosy coloured ; gro wing
IIl al'gin a b ru pt, surrou nde d by a nar row fringe of sub me rged hyph ae
"Xll:lld ing less th an 1 mm beyond th e ae ria l growth; very poorl y sporing, with
('C1 lol1 Y su rface remaining persistently white to white rosy ; ex udate a nd od ou r
lnckin g; no coloured pi gment diffusing in to th e surro u nd ing agar; co lony
n -verse (co lou r plat e 53, a) in very pale cream ro sy shades; penicilli strictly
simple , co m para tively small (fig. 53, A) , with very sho rt and slende r stipes
lll'ising fro m aerial hyphae as sho rt b ranch es, smo ot h walled , usu all y
;1.0- 15.0 tm in length by 1.0-1. 5 tui: in d ia m et er , very slightly enla rged at
Ihe a pex up to 2.0 {tm in d iamet er, terminating in a sim ple penicillus, bu t
uccasioriall y remaining ste rile 0 1' bear ing single phialides; ph ialides di vergent ,
in clus te rs of 2- 7 elements in th e verticil , m ostl y 5.0- 6. 5 tut: by 2.0 - 2.5 {tm ,
tcrrninating in a co ns picuo us ly narrow ed , sho rt neck ; co ni d ia ellipso id to
ovoid , show ing 6-9 lon gitudinal ridges givin g th em a st riate appeara nce, 01'
srcllate wh en seen in cross-sec tion , mostly 2.5 -4.0 tm by 2.0-2.5 {tm , with
ro nspicuo us co n nec tives between th em (fig . 53, B, and SEM pla te 9 , bottom :
I

;1

lcft): eonid iul chains looscly div crgcnt, up to 60 Jtm lon g (fig. 5:i, C ).
C olonies upon C YA (eolour piule 5:i , B and b) growing u littl e less
restrictedly than aboye, attaining a diamct er 01" :i O mm in 14 days at room
temperature (20-25 OC ), similar in texturc lo thos e grown up on CA, but
showing de ep radial furrows , margin raised and centre depressed, strongly
pulling and cracking the agar underneath (colour pia te 53, b) ; poorly sporing,
with colony surface remaining persistently white; exuda te abundan tly
produced in pale yellow shades; odour lacking; no coloured pigment diffusing
into the surrounding agar; colony reverse (colour plate 53, b) in cinn amon
shades; penicilli essentially as described aboye.
Colonies upon MEA (colour plate 53, C and e) attaining a diameter of
30 mm in 14 days at ro om temperature , consisting of a close-t extured basal
felt, deepl y velvety to floccose in appearance; white , eve n m argin surrounded
by a fringe of submerged hyphae exte ndin g abou t 2 mm beyond th e aerial
growth; colony su rface convex, abou t 1 mm deep in th e cen tre, covered by a
my celial overgrowth of white , slender hyphae , giving a lanat e appearance to
th e colony; heavily sporing through out , formin g conce n tric, differently
coloured zones , ranging from lila c (Sguy, PI. 1, 3) to pastel blue (Sguy, PI.
XXXII , 474) , pa ssin g through orange-yello w sh ad es, dep ending on th e degree
of maturity of con idia; exudate and odou r lacking; pal e yellow-colou red
pigment diffu sin g into th e su rrou nding agar; colony reverse (colour plat e 53, e)
in pal e yellow shade s; penicilli esse n ti ally as described ab oye.
The sp ecies description is cen tered upon th e typ e strai n C BS 705. 68 = CMI
151,749 = ATC C 22,052 = IJFM 5028, isolat ed from leaf litt er of A cacia karroo by
Prof. M .C . Pap endorf, D ep artment of Bota ny, Potchfstroom , Sout h Africa , in
1969.
Penicillium striatisporum Stolk appears mo st satisfac tor ily assigned to th e P.
restrictum series on accoun t of th e restricted growth and th e sho rt stipes arising
as branch es from ae rial hyphae . H owever, it shows a certa in relation ship with
th e P. adametz ii seri es sin ce sorn e strains produce strong ly funiculose colon ies
on M EA.
T he spec ies is probabl y a soil mi crofungus . Prof. M. C. Pap en dorf isolat ed
three addition al strai ns duplicating th e typ e strain in th e structure of th eir
penicilli a nd conidia, from soil taken at a depth of 15 cm. The colonies of th ese
strains on CA are less thi ck, they spo rulate better , in grayish shades, and up on
MEA th ey are thin and stro ngly funi cul ose .

54. Penicillium vasconiae R amrez el Martnez: fig. 54 , colou r plate 54 ,

and SEM plate 9, (bott om : right).
Penicillium vasconiae Ramrez et Martnez - Mycopathologia. 72: 181-191 , 1980.

15:J

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Fig. 54. Camera lucida

drawings of Penicil/ium
vasconiae Ramrez et
Martnez, ATCC 42,224 . A,
Detailed drawings of the
penicilli. B, Mature conidia.
e, Habit sketches.

C olonies upon C A (colour plate 54, A and a) growing restri ctedly , attaining
a d ia meter of 25-30 mm in 14 days at room tempe ratu re (20-25 OC),
uhho ugh sorne strains attain a diameter of 45 mm, consisting of a tough, closel c x t ur ed basal felt , velvety in appearance, radiately wrinkled, with white, eve n
u iurgi n, surrou nded by a fringe of su b merged hyphae exte nding less th an
1 111m beyond the aerial growth; raised in central areas; sporing abundantly in
pode blue -green shades (Sguy , PI. XXX , 49) in su bmarg in al zones , and in
hrown ish gray shades (Sg uy , PI. XXIX, 429); odour a nd exudate lacki n g; no
coloured pigment diffusing into the su rrou nd in g agar; colony reverse (colou r
plate 54, a) in pale cream ti nt s; penici lli strictly simple , wit h short stipcs,
usually borne as sho rt branch es from aerial hyph ae or from th e substratu m,
smooth wa lled , usu ally less th an 50 .tm in length by 1.5-2 .5 .tm in di am etcr;
phialides bot tle shaped, few in the vertici l, usu ally 2-3 eleme nts , m ea su rin g
D.O- 13.0 .tm by 2.5-3.5 uu, term inating in a conspicuously long, narrowed ,
lap er ing neck (fig . 54, A); con id ia globose , stro ng ly echi nula te (fig . 54 , B, and
SEM pla te 9, bottom : right), commonly 4.0 -4.5 .tm in d iameter, sho wing
hrown colou rs in mass.
C olon ies u pon CYA (colour pIate 54 , B a nd b) spre ad ing broadly, attaining

a diamctcr 01' 50 mm in 14 day s a l room rempcrnru re (20- 25 OC ), co ns isting a l'

a tou gh , close-tcxturc d , vc1vct y felt , rudiat cly furrowcd , with a whitc,
fimbriate margin , su rrou ndc d by a l'ringc 01' sub rncrgc d hyphac cxtend ing
largely up to 4 mm beyond thc aerial gro wth; abundantly sporing in marg-inal
are as, less abu nda ntly in th e ce ntre, in oli ve gree n shadc s (S gu y, PI. XXIX,
426); nonsporulat ed a reas sho w white to ochraceou s sha des; exuda te and
odour la cking; no colou re d pigment diffu sin g into th e surro u nd ing agar;
colony revers e (colour plat e 54, b) in cream shades ; penicilli essentially as
described aboye .
C olonies upon M EA (colou r plate 54 , C and e) gro wing rapidly , attaini ng a
diamet er 01' 50 mm in 14 days at room temperature (20-2 5 OC), cons isting of a
den se , close-textured basal felt , velvety in appearance, plane , with an
irregul arl y di ssected m argin surrounded by a wid e fringe 01' re dd ish browncolou re d, su bme rge d hyphae extend in g m or e than 10 mm beyond th e ae rial
gro wt h; heavily spo ring throu ghout in gray shades (Sguy , PI. XXV , 523);
exudate and odo ur lacking ; no coloured pigmen t diffusing in to th e
surrou nd in g agar; colony reverse (colou r pIat e 54, c) in sienna brown colou rs
(Sgu y, PI. XIV , 201); peni cilli esse ntially as described aboye, but with th e
stipes ar isin g eithe r fro m th e su bs tratu m and from aerial hyphae ; conidial
chains in loose colu m ns (fig. 54, C) .
The spec ies dia gn osis is ce ntere d upon th e typ e strain IJ FM 3008 = ATCC
42,224, and seve ra l add itional strains, all isolated fro m a n acid , was he d,
brown soil, in th e Spanish Provin ce 01' N ava rra, by DI' I. M ori yn , in April
1977 .
Penicillium vasconiae appe a rs mo st closely relat ed to spec ies included within
th e P. restrictum series on account 01' its restricted gro wt h upon C A, th e brown
colou re d, conspicuo us ly ech inulate conid ia , and th e sho rt stipes arising
primarily from ae rial hyphae.

Species of dubious position

Penicil/ium albo-simplex Nehira - J. Ferm. Technol. Jpn. 11: 854. 1933. See also
M.M. Kulik, 'A Compilation of Descriptions of New Penicil/ium Species'. Agric.
Handbook No. 351. Agric. Res. Ser. U.S. Dep. Agric. p. 15. 1968.

Penicillium albo-simplex Ne h ira is a species described in 1933 th a t is beli eved to

be closely relat ed to th e P. restrictum series. It is presented here as a species 01'
dubiou s position , since th e au tho r 01' th e present study was not able to exa m ine
th e type strain 01' th e spec ies, because all atte m pts to obtai n it from
Internation al Fungus C ollections failed. Ther efor e , it is not included in th e key
to spec ies .
Ne hiras original description follo ws :

tri !>
'Colonies upon Czapek's agar growlng slow ly, oppeorlng floccose; the penicilli
rru monoverticillate. The colony remains perslstently uncoloured in age at 20-25C.
After 30 days, the colony becomes slightly coloured, in part green, sometimes in
pnrt red-violet . The colony reverse is occasionally partially purple or deep purple.
'ho exudate is uncoloured or yellow, becoming purple in old cultures. The
onidiophores are generally monoverticillate, sometimes branched, 2 tm in
lIometer; phialides measuring 12 tm in length, the conidia are globose, 1.5-2.5 tm
In diameter, smooth walled.
Colonies upon malt extract agar with a wide, white margin, sometimes 5 mm in
w ldth; becoming partially purple-red coloured. It does now grow at 33C.'
(Translated from the Japanese by Masae Yamamoto). (The above description
W llS published without providing a Latin diagnosis).

l 'enicillium adametzii series

()I JTSTANDING F EAT URES

Colonies con sisting of a tough , close-textured basal felt , ap pearing

NInncwhat lanate and usually showing sorn e development 01' funicles or rop es 01'
hyph ae . C olony revers e frequently sho wing orange-red to brown or amber
('olours.
The stipes 01' the conid iophores are usually short, ari sin g m ustly from
I ra iling hyphae or ropes of hyphae , as perpendicul ar branches, com monly k ss
rhan 100 tm in length .
Pcni cilli strictIy sim ple, seldo m branched , with conid ial chains tangled or in
wcll-de fine d colum ns.
Conid ia glob ose or su bglobose, usually 2.0 - 3.0 tm in di ameter , with walls
~ ran u lar to conspicuo usly ro ughe ne d .
1n th e presen t study, eight additional species have been adde d to th e P.
adametzii seri es , namely: P. adametzioides Abe ex Smith, P. albo-cinerascens
C ha la buda, P. griseolum Smith, P. kazachstanicum N ovobranova , P. lilacinorclunulatum Abe ex Smith , P. resedanum M cL ennan , Ducker et Thrower , P.
.sacculum D ale , a nd P. spin ulo-ramigenum Sa saki ex Ramrez .
A close relationship between the m is not presumed here; they are cons ide red
logether primarily as a m atter 01' conve nicncc . With th e exception of Penicillium
sacculum D ale , th ey do , however, h ave ce rta in gen eral characteristir s in
cornmon , particul arl y the pred ominant funi culose characte r ofthe colon ies ancl
I he short stipes 01' th e con id iopho res borne as perpendicul ar branch es on rop es
01' hyph ae.
In our opinion, P. sacculum D ale ca nno t be clas sified sa tisfacto rily in a ny 01'
Rap er and Thom's (1949) series of Penicillium. Furthermor e , G . Smith (1961)
created th e m on ot ypi c ge nus Eladia, ba sed on P. sacculum D ale , on th e basis
t hat the spo rogenous cclls wer e phialides , a character sha re d wi th th e genus

Penicillium. O n th e othe r hand , P. sacculum uppeu rs closely related lo thc species

inclu d ed within th e P. adametzii seri es o n uccou nt 01' th e Iuni culosc ha b it 01' ils
colon ics, Therefore , a nd fo r th e time being , it is tcntatively placed here in a
new sub-series whi ch has be en erected primarily to accomo datc th is peculi a r
spe cies.

K ey to spe cies

Paga
1. Colonies with the funiculose habit predominant or well developed.
P. adametzii series . .. . 155
a. Colonies usually developing dull reddish orange to brown shades in
the colony reverse.
1'. Phialides comparatively small, usually less than 10 .tm in length .
aa. Conidia comparatively small, up to 3 .0 .tm in diameter.
1". Conidia smooth walled or nearly so, globose, up to 2.8 .tm in
diameter.
P. kazachstanicum Novobranova . . . . 173
2". Conidia delicately granular, globose to subglobose, ranging
from 2.5 to 3.3 .tm in diameter.
P. adametziiZaleski ... . 157
3". Conidia conspicuously roughened, globose , up to 2.5 .tm in
diameter.
P. albo-cinerascens Chalabuda ... . 175
bb. Conidia comparatively larger, up to 4.0 .tm or more in diamete r.
1". Phialides up to 10 .tm in length, con idia globose , ranging
from 2.5 to 4.0 .tm in diameter , conspicuously roughened.
P. lilacino-echinulatum Abe ex Smith . . . . 170
2". Phialides very small, ranging from 4.0 to 8.0 .tm in length,
with a narrowed, pointed neck; conidia globose, up to 5.0 .tm
in diameter, roughened, with conspicuously long connectives
between them.
P. griseolum Smith .. .. 104
2'. Phialides comparatively large, more than 10 .tm in length.
aa. St ipes of the conidiophores and con idia conspicuously
roughened, especially upon MEA.
1". Phialides atta ining a length of 14 .tm, conidia globose or subglobose , consp icuously spinulose .
P. spinulo-ramigenum Sasaki ex Ramrez .. .. 161
2". Phialides attaining a length of 12 .tm, conidia globose to subglobose , about 3.0-3.5 .tm in diameter, definitely rough .
P. terlikowskiiZaleski . .. . 159
bb. Stipes and conidia smooth-walled or nearly so.

1". Phialides terminating In a long, toporod, narrowed neck ;

conidia olliptical, very small, up to 2.6 Itm in long axis.
P. resedenum McLennan, Ducker etThrower . ...
2". Phialides terminating in a short, narrowed neck; conidia
elliptical, comparatively larga, up to 5.0 Itm in long axis.
P. adametzioides Abe ex Smith ....
h. Colonies not developing dull reddish orange brown shades in the
colony reverse, growing very poorly and restrictedly upon CA, penicilli
with small c1usters of phialides, not unfrequently single; conidia
conspicuously echinulate, up to 7.0 Itm in diameter.
P. sacculum sub-series
P. sacculum Dale
c . Colonies quickly developing deep vinaceous to purple red colours in
the colony reverse; similarly coloured pigment diffusing into the
surrounding agar.
P. vinaceum sub-series . .. .
1' . Phialides comparatively small, up to 8 Itm in length, terminating in
a tapered neck; conidia elliptical, up to 3.0 Itm in long axis.
P. vinaceum Gilman et Abbott . . . .
2'. Phialides up to 15 Itm in length, terminating in a tapered neck;
conidia up to 3.51tm in long axis, deep purple red pigment diffusing
into the surrounding agar .
P. phoeniceum van Beyma .. ..

168

166

177
178

180

180

182

Wi th in the series, individual strains vary from essen tially lanate , showin g a
limi ted development of ropes of hyphae, to d efinite1 y funicul ose .

:l:">.

Penicillium adametzii Zale ski: fig. 5 5, colour plate 55 , and SEM platc

10, (top: left) .

Penici/lium adametzi Zaleski - Bull. Acad. Polonaise ScL Math. Nat. Sr. B, pp .
507-509.1927. See also Ch. Thom, 'The Penicillia', pp. 194-195. 1930; also Rapar
and Thom, 'A Manual of the Penicillia', pp. 228-231. 1949.
Sy no ny m

Penici/lium niklewski Zaleski - Bull. Acad . Polonaise Sci . Math . Nat. Sr. B, pp.
504-505. 1927.
Colonies upon CA (co lou r pIa te 55, A a n d a) grow ing rather restrictedly,
a lta in in g a diameter of 28 mm in 14 days a t ro om temperature (20-25 OC ),
consistin g of a tough close-textu red basal felt , abou t 1 mm d eep in central
arca, zonate, a ppea r in g lanate , with colony su rface cove re d by a mycelial

B '

lwJ.lJ.wJ

10 )Jm

, >~~/. ~
t j" \)
L---...J

I..--...J

100 )Jm

100 )Jm

Fig. 55. Camera lucida

drawings of Penicillium
adametzii Zaleski, CBS
209.28. A, Detailed drawings
of the penieilli. B, Mature
eonidia. e, Habit sketehes.
D, Ropes of hyphae with
eonidial struetures arising
from them.

overg rowth of tr ailing sim ple hyphae and prominen t funi cles or rop es of
hyphae , eithe r prostrat e , tr ailing, or asce nding ; conid ial areas in pale blu egreen shades (Sg uy, PI. XXVII , 400); ma rgin su rro u nded by a fringe of
submerged hyphae ex ten d ing abou t 1 mm beyond th e aerial grow th ; exudate
lacking to abundantly produced , light am ber coloured; odo ur lacking or
indefinite ; a mbe r-co lou red pigm en t diffu sin g into the surrou nd ing agar;
colony reverse (colour pla te 55, a) in orange shades ; peni cilli str ictly sim ple
(fig . 55, A), rar ely showing on e to two branch es with seco ndary penicilli at
lower levels , with stipes mostly borne as perp endicul ar , short branch es fro m
tr ailing hyphae or rop es of hyphae (fig. 55, D ), com mo nly 20-3 0 /lm but
som etimes up to 50 /lm in len ght by 1.5- 2.0 /lm in diameter , smooth walled;
phialides in small clus ters of 3-8 eleme nts in th e ve rt icil, mostly 5- 7 /lm by
2.0 -2.5 /lm, terminating in a short , narrowed neck ; co nid ia globose to
subg lobose, 2.2-3. 0 /lm in diameter, with walls delica tely rou gh en ed (fig. 55,
B, and SEM plat e 10, top : left); conidial chains tangled , up to 50- 100 /lm in
len gth (fig . 55, C ), or in sho rt well-de fine d colu m ns (fig. 55, D ).

C olo n ies upon C YA (colour pl at e 55,13) nuuining a diamc ter oCiO mm in 14d ays a t room ternperature (20 -25 OC) , duplica ting in te xtu rc a no cu lt ural
cha racter istics th ose gro w n upon C A, hut up lo 2 m m d eep ; conidial arca s in
pale gray-gree n shade s (S guy , PI. XXll , :i:iO); poorIy sporing; exu date ano
odo u r lacking ; amber colou re d pigrnent diffusing in to the su rrou n d in g agar;
colon y rev erse in orange shades (colour plate 55, a); penicilli essentially as
dcscr ib ed a boye .
C olon ies upon MEA (colour pi at e 55, C and c) spread ing less restrictedly,
a tta in ing a diamet er a l' 45 mm in 14 days a t room temperature (20-25 OC),
consisting 01' a thinner felt , deeply velvet y to lana te in appearance , alm ost
plan e , sligh tly rai sed in th e co lony cen tre, cons picuous ly zonate; growing
margin su rrou n de d b y a w ide fringe 01' su b me r ged hyphae exte n di n g a bou t
:3 m m beyond th e ae r ial gro w th; h eavily spor ing in concen tr ic zo nes in very
pale gray-green shades (Sguy , PI. XXII , 330); exu date a nd od our lacking;
pale a m be r pi gment diffusing slowl y into the surrounding ag ar ; col on y reversc
(co lou r pl at e 55 , e) in pal e cream sh ades; penicill i esse n tially as d escribcd
aboye.
The spec ies d escription is centered upon the typ e strain CBS 209. 28 = CMI
:19 ,751 = ATCC 10,407 = N RR L 737= QM 1916=IJFM 500 6 , isola ted frorn soil
under a con ifer forest in Poland , by K. Zal eski.
Strains sh owing cu ltu ral a n d m orphologi cal characte r istics approximatin g
(hose described aboye are cornm only isolated from soil and from m at erial
subj ecte d to soil or dust-borne con ta m ination . Individual stra ins show com rnonly marked differences , with respeet to the degree ofropiness and the rate 01'
growth, without showing su fficie n t bases for separa t ion .

56 . P enicillium lerlikowskii Z al eski : fi g. 56 , colour plate 56 , and S EM

pl ate 10, (top : right) .
Penicillium terlikowskii Zaleski - Bull. Acad. Polonaise Sci. Math. Nat. Sr. B,
pp. 501-502. 1927. See also Ch. Thom, 'The Penicillia', pp . 203-204. 1930; also
Raper and Thorn, 'A Manual of the Penicillia', pp. 231-234. 1949.

Synonymies
Penicillium jentho-citrinum Biourge - Monographie, La Cellule. 33: 311-313. 1923.
Penicillium paczoskii Zaleski - Bull. Acad. Polonaise Sci. Math . Nat . Sr. B, pp.
505-507. 1927.

Colon ies upon CA (eolo ur pl at e 56, A a nd a) a ttain ing a di amet er 01' 45 mm

in 14 days a t ro om temperature (20 - 25 OC), eon sistin g 01' a tough , closetextured b asal felt , ove rl aid by a lanat e to somewhat fu nieu lose aer ial
myc eliu m eom po sed 01' trailing a n d interwoven hyphae a n d ropes 01' hyphae ,

":.',;-;'
,.:~:::

:1

." :0 ".",

:.:;::~"

10 ).1m

tS~r
L..--....J

100 jarn

fR.,

Fig. 56. Camera lucda

drawings of Penicillium
terlikowskii Zaleski, CBS
228.28. A, Detailed drawings
of the penicilli. B, Mature
canidia. C, Habit sketches.

somewhat zon ate, plane , with central area usually umbonate; undulating
margin sur rou nde d by a frin ge of subme rge d hyphae extend ing ab ou t 2 mm
beyond th e ae rial growth; medium to he avily sporing throu gh out or in sorne
strains in m arginal a reas only, in pale gr ay-green shades (Sguy , Pi. XXX,
450) , sometimes partially obsc ured by a white m ycelial overgrowth ; odo ur
faint , indefinite ; exudate abundantly produced , as amber coloure d droplet s, or
lackin g; bright yellow coloured pigment diffusin g into th e su rrou nd ing agar ;
colony revers e (colour plat e 56, a) in reddish orange shades; penicilli strictly
sim ple, seldom bran ch ed (fig. 56 , A) , with th e stipes of th e con idiopho res
arising primarily as short branches from aerial hyphae or ropes of hyphae,
usually less th an 100/lm in len gth by 2.0-4.0/lm in diamet er, with wall s
ranging from appare n tly smo oth to conspicuo u sly granula r; phialides in
clusters numbering 5- 10 elements in th e verticil, mostly 9-12/lm by
3. 5 - 4.5 tui , terminating in a con spicuously short , narrowed neck ; con idia
globose to subglobo se, mo stly 2.8 - 4.0 /lm in di ameter , with walls definitely
rou ghen ed (fig. 56, B, a nd SEM plat e 10, top : right) ; conidial chains

pro d ucing loos e lo well -d efincd co lu m na

IIp lo 50 .un 0 1' mo re in lcn gth.

Colonics upon C YA (colour plato B ami b) auaining a diamct er 01' 55 mm in
Ji! days a l room tcmperature (20 - 25 OC), co nsistin g 01' a similarly texturcd
bnxul fclt , deeply velv cty lo loccosc , but strongly and dceply radiately
[u rrowed , with cen tral area irregularly olded, overlaid by a Ianatc lo
Iuni culose mycelial overgrowth cornposed oftrailing and interlacing hyphae 01'
rn pcs 01' hyphae, partially obscuring the otherwise heavily sporing arcas in
hlu e-green shades (Sguy, PI. XXX, 442) ; with cven margin surrounded by a
I'riIl gc 01' submerged hyphae cxtending about 1 mm beyond th e aerial growth;
ex ud a re and odour lacking; deep orange-coloured pigment diffusing into th e
su rr ou n d in g agar; colony reverse (colour plate 56, b) in reddish orangc shade s ;
pcni cilli essentially as described aboye .
C olon ies upon MEA (colour piate 56, C and e) attaining a diamcter 01'
.1;1 mm in 14 days at room temperature (20- 25 OC), consisting 01' a loo sercxtured basal feit , velvety in appearance , plane with centre umbonate,
r onspicu ou sly zanate, with thin margin surrounded by a wide frin ge of
suh merged hyphae arising about 4 mm 01' more beyond the acrial growth;
cxud a te and odour lacking; deep amber-coloured pigment diffusing in lo the
su rr ou nd in g agar; colony reverse (colour plate 56, e) in brownish or an ge
xhades; penicilli essentially as described aboye but with phialides more
nurnerous in the verticil, with the stipes 01' the conidiophores mostly al'ising
primarily from the substratum, and the conidial chains in definitc1y com pac t
ro lu m ns, up to 300 tm in length (fig . 56, C) .
The species description is centered upon the type strain CBS 228 .28 ... C M I
!J2 ,232 = NRRL 752, isolated from conifer forest soil in Poland by K . Zaleski.
Penicillium terlikowskii Zaleski is regarded as occupying a rather terminal
position within the P. adametzii series, since it represents something 01' an
xtrcme in the variation normally encountered in the series. The colonies show
lite basic characteristics 01' P. adametzii but usually less definitely funiculose.
I.ike other members 01' the series , the species may be regarded as a typical soil

lorrn .

':> 7. Penicillium spinulo-ramigenum Sasaki ex Ramrez: fig. 57, colour plate

57, and SEM pIate 10, (centre : left) .
Penicillium spinulo-ramigenum Sasaki - J. Fae. Agrie. Hokkaido University. Japan.

49: 153. 1950.

Penicillium spinulo-ramigenum Sasaki was isolated from Japanese butter by Y .
Sasa ki in 1949, who proposed the name 01' P. spinulo-ramigenum and described
the species without providing a Latin diagnosis . 1'0 validate the species a Latin
diagnosis is given below ; the name 01' th e original author is maintained under

e
-:

'.

e
~

10,um

t11

.R..

100

jrrn

Fig. 57. Camera lucida

drawings of Penicillium
spinulo-ramigenum Sasaki
ex Ramrez, eBS 344.59. A,
Detailed drawings of the
penicilli . B, Mature conidia.
e, Habit sketches.

R ecommendati on 46 C of the Internati on al C od e of Bot ani cal N om en clature

(Sta fleu (E d.), 1972).
Latin di agn osis of Penicillium spinulo-ramigenum
Sasaki ex R amrez:
Coloniae in agaro Czapekii satis celeriter crescunt, post duas hebdomadas ad
20-25C, 53 mm in diametro attingunt, velutino-floccosae et funiculosae in centrali
area, denso texto basali coacto mycelio, fere planae, cum angusta margine ex submersis hyphis 1 mm long is circundata est; fortiter sporulantes, griseo viridae
colorantur; cum superficiebus fructificationibus conidialibus laxis tectae sunt e
hyphis flocossis albis et funiculosis; odor et exsudatio absunt; succineo colore in
medio diffluente; reverso aurantio; neque ascomata neque sclerotia visa sunt;
penicilli monoverticillati stricte vel ramigena, conidiophora levia vel aspera, 25-90
.tm longa per 2.5-3.5 .tm in dimetro. ex hyphis aeris reptabundis vel ex funiculis
exoriuntur, cum apicibus inflatibus tenus 6 .tm in diametro; phialidae lagoeniformae,
acuminatae, plerumque 6-12 per verticillum, 8-13 .tm per 3.0-4.0 .tm; conidia
globosa vel subglobosa, 3.0-4.3 .tm per 2.5-4 .0 .tm in diametro, leviter asperula.

1c;:

Coloniae in agaro Czapekii cum extracto levedinis, post duas hebdomadas ad

O-25 C, 50 mm in diametro attingunt, similiter in texturam sicut coloniae in agaro
pekii, sed fortiter radiatae sulcatae; parve sporulantes, albae colorantur; odor et
xsudatio absunt; reverso cinnamomeo; colore succineo in medio diffluente; neque
rscomata neque sclerotia visa sunt.
Coloniae in agaro hordeaceo, post duas hebdomadas ad 20-25C, 56 mm in diamot ro attingunt, velutinae, planae, denso texto basali coacto mycelio, cum margine
submersis hyphis circa 4 mm longis circundata est; fortiter sporulantes, griseoviridae colorantur; odor et exsudatio absunt; colore aurantio flavo in medio
cliffluente; reverso aurantio; neque ascomata neque sclerotia visa sunt.
Sasaki's original description follows:
'Colonies on Czapek's solution agar spreading broadly, 2.1-2.2 mm deep in the
centre, margin mostly submerged, velvety in marginal area, funiculose in the
contre, azonate, slate-olive (Ridgway, PI. XLVIII, later overlaid by white aerial
hyphae; reverse cream buff or honey yellow in the centre (Ridgway, PI. XXX), olive
gray in margin (Ridgway, PI. L1). Conidiophores arising from submerged hyphae, or
os branches from aerial hyphae, or ropes of hyphae, simple or ramified, with the
secondary branches, each terminating in a monoverticillate penicillus, with walls
conspicuously spinulose, 90-350 .tm in length by 2.5-4.5 .tm in diameter, enJarged
ot apex up to 5-6 .tm in diameter; phialides 10-12 .tm by 3-4 .tm, numbering 8 to 9
olements in the verticil; conidia globose or subglobose, 2.2 to 3.5 .tm in diameter,
delicately roughened; conidial chains parallel; ascomata or sclerotia not found.'

Our not es follow :

Colonies upon C A (colour pIat e 57, A a nd a) spreading broadl y , attaining a
d iameter of 53 mm in 14 days at ro om temperature (20- 25 OC), cons isting of a
close-textured bas al felt, de eply velvety to lanate , with colony surface ove rla id
by an ove rgro wth of aerial interwoven hyphae and ropes of h yphae , espec ially
in th e centre, partially obs curing th e othe rwise abundantly spo ring a re as
coloured in gray-green sh ades (Sgu y , PI. XXVIII , 417) ; eve nly margin
surrou nded by a fringe of su b me rge d hyphae exte nd in g ab out 1 mm beyond
the ae rial gro wth ; odour and exudate lacking; light amber colou re d pigment
diffu sin g into the su rro u nd ing agar ; colony reverse (colour plat e 57, a) in
ora nge shades; penicilli strictly m onoverticillate or sim ple , seld om branched,
but with occasional branch es terminat in g in a sim ple peni cillu s reminding th e
R aper and Thom's (1949) R amigen a series in th e Monoverticillata section ; th e
stipes of th e conid iophore s are smoo th walled or consp icuo usly roughened,
usual1y short, 25-90 /lm in length by 2.5-3.5 /lm in diameter, with apices
enlarged in a vesicle up to 6.0 /lm in diameter, mostly arising as short branches
from ae rial hyphae or from ropes of hyphae; phialides bottle- shaped , in
crowded clu sters of 6-12 elements in th e verticil , u su ally 8.0-1 3.0 /lm by

:3.0 -4.0 Ju n , terminating in a ShOI'I, tapcred ncck; conidia globoso lo

su bglobose, :~.0 - 4.3 u n by 2.5 -4.0 J.tlll in diumetcr, dcl icaic ly roughencd
(SEM plate 10 , centre : left , a nd ligo 57, 13); conidial cha ins in looscly pa rallcl
columns (fig. 57, C) .
Co lonies upon CYA (colour piate 57, B a nd b) attai n ing a diameter of
50 mm in 14 days at room temperature (20-25 OC), duplica ting in texturc
those grown u pon CA, but with the colony surface deeply radiately sulcate,
poorly sporing, and remaining persistently white; thin, undulating margin
with a narrow fringe of su bmerged hyphae extending about 1 mm beyond the
aerial growth; exudate and odour Iacking ; amber-coloured pigment diffusi ng
into the surrounding agar; colony reverse (coIour piate 57, b) in cinnamon
shades; penicilli essentially as described aboye.
CoIonies upon MEA (coIour pIate 57, C and c) attaining a diameter of
56 mm in 14 days at roo m temperature (20-25 OC ) , velvety in ap pearance,
pIane, with centre umbonate; heaviIy sporing throughout in deep gray-green
shades (Sguy, PI. XXX, 443); odour and exudate Iackin g; yellow-orangecoIoured pigment diffusing into the surrounding agar; colony reverse (colour
plate 57, e) in orange shades; pe nicilli essentially as described aboye, but with
stipes always conspicuously roughened (fig. 57, A), and conidial chains in welldefined columns (fig . 57, C).
Species diagnosis is centered upon the type strain CBS 344.59 = ATCC
18,486=CMI 68,6 17=IFO 5,359=QM 8, 152=IJFM 3758, isoIat ed from
Japanese butter by Y. Sasaki in 1949. It is known on ly from the type strain .

58. Penicillium griseolum Smith : fig. 58, colour plate 58, and SEM plate
10, (centre: rig ht) .
Penicillium griseolum Smith - Trans. Brit . Mycol. Soco 40: 485. 1957. See
also M.M. Kulik , 'A Compilation of Descriptions of New Penicil/ium Species'.
Agric. Handbook No. 351. Ag ric . Res. Ser. U.S. Dep. Agric. pp. 42-43. 1968.
Smith's original diagnosis follows, since this adequately presents the species in
question:
' Colonies on Czapek's agar spreading moderately quickly but extremely th in,
transparent, showing no visible myce lium but only scattered penicilli , giving a faint
greenish hue to the colony; on malt agar or potato-dextrose agar growing
moderately well, at first pale gray w ith broad white edge, turning darker gray,
matted funiculose about 1 mm thick; reverse brown with a greenish tone (somewhat Iike mustard but rather darker and duller) ; penicilli strictly monoverticillate;
conidiophores arising from aerial hyphae, very short, smooth, 12-35 by 1.2-2.0
.tm ; phialides few in the verticil, abruptly pointed, 6-7 .tm by 1.6-2.0 .tm; ripe

Jf

A~

y=vL

V0

kLl

~~

-==~
'.

.~
L...---...I

100

jrm

--- ~

lu.uJ.w..J
10 prn

R.

Fig. 58. Camera lucida

drawings of Penicillium
griseolum Smith, CBS
277.58. A, Detailed drawings
of the penicilli. B, Mature
conidia .
Habit sketches .

e,

conidia globose or nearly so, faintly verrucose, 3.1-4.0 .tm in diameter but mostly in
the range 3.2-3.5 .tm, in parallel chains or forming somewhat loose columns; young
conidia oidium-like, small , cylindrical, swelling only slowly and remaining in chains
attached to the phialides, even on mounting fluid.'

Our notes fol1ow:

Co lonies upon C A (co lour piate 58 , A) ex traord inarily thin, m ostly
subme rge d, att ain ing a di ameter of 45 mm in 14 days at room temperature
(20 -2 5 OC ) , cons isting of a loose-textured basal felt , transparent , with
scattered con idial structures; penicil1i (fig. 58, A) strictly sim ple , with sho rt
stip es ari sin g frorn ae rial hyphae and frorn rop es of hyphae , smo oth wall ed ,
usually less than 25 p.m in len gth by 1.0-1. 8 iuu in diameter; phialides
com para tively sm all , bo ttl e shap ed , com monly 5.0- 6 .0 utt: by 1.8- 2.8 pst ,
term inating in a lon g , narrawed, tap ered neck ; phialides few in number, 2- 5
ele me nts in the verticil; conidia elliptical whe n you ng, becoming globose at
maturity , 4.0- 5.0 p.m in di ameter, cons picu ou sly echin ula te (SEM plat e 10,
centre : ri ght); conidial ch ains in loosely parallel columns , with conspicuou s
con nectives b etw een th e con idia (fig. 58 , B a nd C) .
Colonies upon CYA (colour piate 58, B a nd b) attain ing a diameter of
35 mm in 14 days at room temperature (20 - 25 OC ), consi sting of a close-

textured basal elt , overl aid by a my cclial ovcrgro wth 01' closcly int crw oven
ae ria l and Iunicul ose hyphac , obs curing almos t tot ally the othe rwise
abu nda n tly produced con id ial structures, rc maini ng persistcntl y whitc,
showing th e gra y-coloure d sporing ar eas especially in m arginal zon cs ;
undulating, thin margin surrou nde d by a frin ge 01' su bme rged hyphac
extend ing abou t 2 m m beyond th e ae rial growth; exuda te and odo u r lacking;
pale bro wn coloured pigment diffu sin g into th e surrounding agar; colony
re ver se (colour plate 58 , b) showing brown colou rs; peni cilli essen tially as
descr ibed aboye.
Colonies upon M EA (colour pIat e 58, C and e) spread ing broadl y , attai n ing
a di ameter 01' 55 mm in 14 days a t room temperature (20-25 OC), cons isting 01'
a den sely textured basal felt , deepl y velvety to lan at e , ove rla id by a m ycelial
network of ae rial hyphae and ro pes of hyphae influen cin g partially th e colony
a ppeara nce; thin m argin su rro unde d by a wid e frin ge of su bme rge d hyphae
exte nd ing abo ut 3- 4 mm beyond th e ae rial grow th; ce ntral a nd su bce n tral
areas heavily spo ring in gra y shades (Sguy, PI. XXI , 310); conid ial structures
are obscured in m a rgin al areas by th e white mycelial ove rgrowth of floccose
ae rial hyphae ; ex udate and odo ur lacking ; no coloured pigm ent diffusin g int o
the su rro u nd ing agar ; colony rever se (colou r pIat e 58 , c) in orange sha de s;
penicilli esse ntially as described ab oy e .
T he species description is ce ntere d up on th e typ e stra in C BS 277 .58 = CMI
71,626 = ATCC 18, 239 = IFO 8.175 = Q M 7,523 = IJFM 5 113, isolated in] uly 1955
by Miss]uliet C . Brown , of the R oyal H ollow ay College, from th e ac id dunes
of Studla nd, Dorset , U K . It is kn own only from th e typ e strain.

59 . Penicillium adametzioides Abe ex Smith : fig. 59 , co lo u r pl ate 59 , and

SEM plate 10 , (b o ttom: left) .
Penicillium adametzioides Abe - J. Gen. Appl. Microbiol. Tokyo. 2: 68-69. 1956.
See also Smith, Trans. Brit. Mycol. Soco 46: 335. 1963; also M.M. Kulik, 'A
Compilation of Descriptions of New Penicillium Species'. Agric . Handbook No.
351 . Agric. Res. Ser. U.S. Dep. Agric. pp. 12-13. 1968.

C olonies upon CA (colou r pIate 59, A and a) growing ra th er moderately ,

attaining a di ameter of 40 mm in 14 days a t room temperature (20-25 OC ),
cons isting of a tough , close -tex tured basal felt , ve lve ty to funiculose, zonate
deepl y radiately wrinkled; th in margin surrounded by a narrow fringe of
su bme rge d hyphae extend ing abo ut 1 mm beyond th e ae rial gro wth; poorly
spo ring in ve ry pale blue-gr een shades (Sguy , PI. XXVII , 400), with th e
colony su rface rem aining persistently white ; exudate and odou r lacking ; no
colou re d pigm ent diffusin g into th e su rrou nd ing agar; colony reverse (colour
pIate 59, a) colou rle ss 01' in ve ry pale cream shades ; penicilli str ictly sim ple (fig .

i-l

~i4 \~ 1=-~~'-"'

'tr x

l-.-...-J

100 prn

iR.

'o

L '

.c>

Fig. 59. Camera lucida

drawings of Penicillium
adametzioides Abe ex Smith,
ces 313.59. A , Detailed
drawings of the penicilli . B,
Mature conidia . e, Habit
sketches .

59, A) , with stipes sho rt, ari sin g mostly from aeri al hyphae or rop es ofhyphac ,
sometime s from th e subs tra tum, com mo nly 25 p,m or less in len gth , by
2.2-3. 0 p,m in diamet er, with apices enla rge d in a ves icle up to 4 .5 p,m in
diame ter, smoo th walled , septate, phialides in cro wde d clu sters of 8 - 12
cIeme nts in the verticil, bottle sha pe d with a cons picuo us , sho rt, pointed neck ,
mostl y 10-1 2 p,m in len gth by 2.8-4.0 p,m in wid th; conidia broadly elliptica l
to subglob ose, smoo th walled or nearly so (fig. 59, B, and SEM pIate 10,
botto m: left ), mostly 3.0-4.8 p,m by 2.5-4.0 p,m; con idia l chains ta ng led (fig.
59, C) .
Co lonies upon C YA (colour plate 59 , B a nd b) attai ning a diame ter of
50 mm in 14 days at room temper ature (20 -25 OC), duplicating in textu rc
those grown up on CA, but deepl y radiat ely suleate with nume ro us furrows ,
and centre folded in conce ntr ical wrinkles in a cra teri form patt ern ; he avily
spo ring throu gh ou t in blue shades (Sguy, PI. XXX II, 477) , especially in
marg inal areas, a nd partially obsc ure d in central and subce n tral areas by the
white ove rgrowth of aeri al hyphae ; m argin thin , surrou nde d by a narrow
fringe of sub me rge d hyphae ex te ndi ng abo u t 1 m m beyon d th e aerial growth;

colon ics somctimes strongly pulling away thc agnr undcrncath, and crac king
it, thu s permitting the pr escn ce 01' air in the reverso a nd thc production 01'
abundant conidial structures (colour plate 59 , b) ; exudare lacking; odour faint I
indefinite; bright yellow-colou red pigment diffu sing into th e surrounding agar;
colony reverse (colour plate 59 , b) in deep orange shadcs; pcnicilli essentially
as described aboye .
C olonies up on MEA (colour plate 59, C and c) att ain ing a diameter 01'
35 mm in 14 days at room temperature (20 - 25 OC ), consisti ng 01' a closetextured basal felt , ve1vety with sorne centra l funicles , plane , with a thin
m argin surrounded by a wid e fringe ext ending about 7 mm beyond th e aerial
growth ; heavily sporing throughout in blu e shades (Sguy , PI. XXXII , 447);
exudate la cking; odour faint , aromatic ; revers e in ora nge sh ad es (colou r platc
59, c); bright yellow- coloured pigm ent diffusing into th e surrounding agar;
peni cilli esse ntially as described abo ye.
The species de scription is cente red up on th e type strain CBS 3 13.59 = CMI
68 ,227=ATCC 18306=IFo 6 ,055=QM 7, 312=IJFM 5112 , isolat ed from
J apanese soil by S. Ab e in 1955.

60 . Penicillium resedanum McLennan, Duckcr el Thrower: fig. 60 , colou r

plate 60 , and SEM plate 10, (bottom : right) .

UJJJ..l.u.i.JJ

10 jrrn

O O o O
O O Oo o
o 80 O
0 0
O 0

00

O 00

~r~
~

k,

0(';\ 1 ~

~:j

L--J

100 jrrn

Fig. 60. Camera lucida

drawings of Penicillium
resedanum McLennan,
Ducker et Thrower, CBS
181.71. A, Detailed drawings
of the penicilli. B, Mature
conidia. e, Interlacing
vegetative hyphae. D, Habit
sketches.

Ponicillium resedenum McLennan, Duckor et Throwor - Aust. J. Bot . 2: 360-361.

1954. See also M .M. Kulik, 'A Compllotlon of Doscrlptions of New Ponicillium
Species'. Agric. Handbook No. 351. Agr lc. Ros. Ser. U.S . Dep. Agric. pp. 70-71.
1968.

McLennan , Ducker and Thrower's ori ginal diagno sis of th e specics follow s
since this adequately presents th e spe cies in question when first isolated:
' Colonies upon malt extract agar spreading broadly, 5-6 cm in 14 days at 24C,
with mycelium forming a dense felt. The upper surface is very funiculose, and the
overall colour is mignonette-green (Ridgway, XXXI), fading towards the edge of the
colony to lemon yellow (Ridgeway, IV), with a tendency to zonation of colour. The
margin of the colony is regular, thin, and white . The reverse of the colony is flat,
pale, cream-coloured (Ridgeway, XVI) with a buff centre.
Colonies on Czapek's agar grow more restrictedly than on rnalt, attaining a
diameter of 2.5 cm in 14 days at 24C. The funiculose character is more pronounced
than on malt; the colonies resemble little nests, the brim formed by a creamy white
lanose growing margin which surrounds the sunken mignonette-green centre. The
reverse is cream to avellaneous in the older areas.
Colonies on Raulin's agar grow more rapidly than on Czapek's agar, 4.5 cm in 14
days at 24C. The nest-like character is more pronounced with a small coloured
sunken centre (Kronberg's green, Ridgway, XXXI) surrounded by a very lanose
white area, 1.5 cm bread , with an abrupt margin oThe reverse is slightly buckled, the
centre antimony yellow, darkening to yellow ochre in age.
Morphology as on malt agar. The colony is characterized by a profuse
development of ropes 10-15 .m wide, consisting of coloured hyphae from which
numerous smooth conidiophores arise, 50-150 .m long. Each conidiophore is
sparsely septated uniformly 2 .m bread , the apex is slightly swollen and bears a few
(3-9) phialides; the spore chains are twisted into short columns. The phialides
which are strongly tapered at their apices are 9-12 by 3 .m. The conidial chains
break up readily, the conidia are smooth and tinted, broadly elliptical, 2-3 by 1.5-2
.m.
This isolate is a strictly monoverticillate form and produces neither ascomata nor
sclerotia, even when newly isolated. The phialides are long and tapering,
resembling those found in the Biverticillata-symmetrica section, also the general
colour scheme is suggestive of this group . However, we have never found other
than monoverticillate conidiophores. The colonies are characterized by the
development of prominent funicles or ropes of hyphae, and hence the form must be
grouped in the P. adametzii series. It differs, however, from the other forms in that
series in the broadly elliptical spores, of the type found in the members of the P.
lividum series.'

Our not es Iollow:

Colonics upon CA (colour plato 60 , A ami a) attuinin g a diameter 01' 20 mm
in 14 days at room temperature (20 - 25 OC), co nsisting 01' a tou gh, closetextured basal felt , overlaid by a mycclial ovcrgrowth of intcrwoven aerial
tr ailing hyphae and rop es 01' hyphae; white, thin, cvcnly margin, somewhat
lanate, surrounded by a fringe ol' submerged hyphae extend ing about 1- 2 mm
beyond th e aerial growth ; very poorly sporing, not influen cing th e colony
appearance; exudat e abundantly produced as large , colourless drops ; odour
lacking; coloured pigment diffusing into th e surrounding aga r not produced;
colony reverse (colour piat e 60, a) with orange shades showing through the
otherwise uncoloured reverse ; peni cilli strictly simple (fig . 60, A) , with the
stipcs 01' the conidiophores mostly arising from rop es 01' hyphae (fig . 60, D),
smooth walled , with api ces enlarged in a vesicle up to 5.0 p.m in diamet er,
usually 50- 150 p.m in len gth by 2.0 p.m in width ; phialides in crowde d verticils
01' 8-12 elements in th e whorl , com monly 9-13 ust: by 2 .8-3.0 usn,
terminating in a conspicuo usly tapered, narrowed neck; conid ia comparatively
sm all, elliptical, with walls smooth or nearly so (fig. 60 B, and SEM plate 10,
bottom: right) , mostly 2.5-3. 2 p.m by 1.2-1. 8 p.m ; conidial chains in short
loose colum n s (fig. 60, D) .
Col oni es upon CYA (colour piate 60 , B and b) attaining a di ameter 01'
35 mm in 14 days at room temperature (20-2 5 OC ) , duplicating in texture
tho se gro wn upon CA ; very poorly sporing; exudat e abundantly produced, as
large slimy drops, colou rle ss; odour la cking; no colou re d pigment diffusing
into th e su rrou nd ing agar; colony rev erse (colour piate 60, b) colourless, with a
pale or ange cas t showing through; peni cilli essentially as des cribed aboye .
C olonies upon M EA (colour plate 60 , C and c) attaining a diamet er 01'
45 mm in 14 days at room temperature (20-2 5 OC ), cons istin g 0 1' a leathery
basal felt , very thin, plane , mostly submerged , colou re d in yellowish brown
shades, with a thin , eve nly m argin surrounded by a fringe 0 1' submerged
hyphae exte ndi ng ab out 3 mm beyond th e ae rial growth; colony surface
overlaid by a nctwork 0 1' interwoven aeri al and fu niculose hyphae , especially in
th e colony ce ntre ; poorl y spo ring; exudate a nd odour lacking ; no colou red
pigment diffusing int o the surrounding ag ar; colony rever se (colour plat e 60, c)
in reddish brown shades; peni cilli essentially as des cribed aboye.
The spcc ies description is cen te re d upon th e typ e strain CB S 181. 71 = ATC C
22, 356 = C M ! 62, 8 77 = NR R L 578 = IJ FM 503 7, isolated from th e A l horizon 01' a
sandy podzol supporting a cha racte ristic heathland flora , at Fr ankston ,
Vi ctoria , Au stralia , in 1952. It is at present kn own only from th e typ e spec ies.

61. Penicillium lilacino-echinulatum Ab e ex Smith: fig. 61 , colour plate 61 ,

and SEM plate 11, (top: left) .

L::::=-----

t
.

,\\

i)

" ~

/,/ (\

c~:;8 f)

n (\/)

()()B
';;:'.... ," '"
.-,
;;.c"

.. w.w..u.w

10).lm

fR .
e

~ f rDJ
100).lm

~~

Fig . 61. Camera tucidn

drawings of Penicillium
lilacino-echinulatum Abo /IX
Smith , CBS 332.50. A,
Detailed drawings of th e
penicilli . B, Mature con idia.
C, Habit sketch es.

Penicillium lilacino-echinulatum Abe - J. Gen. Appl. Microbiol. Tokyo. 2: 54. 1956.

See also G. Smith. Trans. Brit. Mycol. Soco 46: 335. 1963; also M.M . Kulik , ' A
Compilation of Descriptions of New Penicillium Species' . Ag ric . Handbook No.
351 . Agric. Res. Ser. U.S. Dep. Agric. p. 49-50. 1968.

Penicillium lilacino -echinulatum Ab e ex Smith was prese nted as a new specics

by S. Abe in a communication read before th e Agricultural Chemical Society
ofJ apan (Tokyo) , in April1954, and th e description ofthe spec ies a ppe are d in
th e Journal of G en eral and Applied Microbiology in 1956 , without a Lati n
diagnos is. G. Smith validated the spe cies by providing a Latin diagnosis ( 1963)
of it. Abe' s original d iagnosis of P. lilacino-echinulatum follows since this
ade q uately presents th e species in question when first isolated :
' Colonies on Czapek's agar growing rat her restrictedly, about 30 mm in 10-12
days at 25C; the cent ral area is raised and sulcate , the surface appearing typically
fun iculose, in pale gray -green shades near Mineral Gray or Pale Olive Gray,

changing to Tea Green; exud ate abundant , colourloss; roverse in near Iight
vinaceous lilac in 6-7 days, changing to yellow shades, and th e surrounding agar In
pale yellow shades; penicilli strictly monoverticillate; conidiophores short, up to
20-40 .tm in length by 1.2-1.9 .tm in diameter, with walls smooth or nearly so, and
apices enlarged up to 3.0-6.2 .tm in diameter; phialides in crowded c1usters,
numbering 6-15 in the verticil, mostly 5.0-7.0 .tm or rarely 9.3 .tm in length by
1.5-2.5 .tm in diameter, acute type with smooth walls or nearly so; conidia globose
to subglobose, w ith typically echinulate or verruculose walls, mostly 2.1-2.8 .tm in
diameter; chains of conidia in parallel or loase columns, commonly up to 30-60 .tm
in length; and conidia showing conspicuously echinulate or verruculose walls in
electro n microscopv .'

Our not es follow :

Colonies u pon CA (co lou r plat e 6 1, A and a) a ttaining a di ameter of 25 mm
in 14 days at room temperature (20- 25 OC ), cons istin g of a tou gh , closetex tured bas al felt , abo u t 2 mm deep in th e centre , st rongly and deeply
radiately wrinkled , with ce ntre irregularly fold ed in a crateriform pattern;
undulating m argin ab ru pt, surrounded by a frin ge of submer ged hyphae
exte nd ing abo u t 2-3 mm beyond the aerial gro wth ; poorl y sporing in pal e
gray-gree n shades (S guy, PI. XXX, 445) not influen cing the colony
appe arance, whi ch remains per sistently white ; exudate as minute dropl ets in
pale yellow colours; light yellow coloured pigm ent diffu sin g into the
su rrou nd ing agar; colony reverse (colou r piate 61, a) in pale yellow shades ;
peni cilli strictly simple (fig . 61 , A) , with sho rt stipes arising m ostl y from
tr ailing hyphae and from rop es of hyphae, smooth walled , less th an 25 utt: in
len gth by 2.8 -3 .5 Ilm in d iamet er , with apices enla rge d in a ves icle up to 6 Ilm
in di amet er; phialides in cro wde d clu sters of 8 - 12 eleme nts in th e ve rticil,
bottle sh ap ed , terminat ed in a sho rt, narrowed neck , mo stly 7.0-10 .0 Ilm by
3.5 - 4 .5 Ilm ; con id ia glob ose to subglob ose , 2. 5- 3.0 Ilm in di amet er ,
cons picuo usly spinulose (fig . 6 1, B, and SEM piate 11, top : left) ; conid ial
cha ins forming well-d efined , short colu m ns up to 75 Ilm in length (fig . 6 1, C ) .
C olonies upon C YA (colour piate 6 1, B and b) att a inin g a diamet er of
35-38 m m in 14 days at room temperature (20- 25 OC ), co ns isting of a tough,
close- tex tu re d basal fe1t, ove rla id by a thin m yceli al network of white floccose
and funi cul ose aerial hyphae influ en cing th e colony ap peara nce, sulca ted by
n ume ro u s radiat e fur rows, and raised in th e colony cent re in a cra teriform
pattern ; co ns picuo us ly zonate , with a n eve nly th in margin su rrounded by a
narrow fringe of sub me rge d hyphae exte nd ing about 1 mm beyond the ae rial
growth; sporing poorl y in olive gree n shades (Sguy, PI. XV , 221), espec ially
in m arginal areas; exu date abundantly produced as pale yellow dropl et s ;
sim ila rly coloure d pigm ent diffusing into th e su rro und ing agar; colony revers e
(colou r plate 61 , b) in pale ora nge shades; penicilli essentially as describe d
~ h rl'\ TP

:jo(onics upon MEA (colour plato 6 1,

und e) att ainin g a di am ct er 0('
111111 in 14 days at room rcmperaturc (20- 25 OC), co lIsistillg' 0 1' a rough,
d llse-Icxtllrcd basal felt , plan e , with a th in lIIargin surrou ndc d by a wid c f'ringc
ul' suln nc rgcd hyphae cxtc nd ing a bou t 10 111111 hcyond th e nc r ial growth, with
l'cJlon y surfacc a ppcaring gra n u la r owin g lo numerous fu nicles disposed in
ru ncc u trical circles sho wing thus a somcwh at zonate surface ; heavily sporing
I hrou ghout in paIc olive green colou rs (S gu y , PI. XIX , 279); od our and
I'XIHl at c la cking; no colou red pi gm ent diffusing into th e su r ro u nd ing agar ;
I'lllo llY reverse (eolour plat c 61 , e) uneoloured 01' nearl y so; penieil1i esse n tial1y
liS dcscribed a bo ye, but with eo nid ial ehains tangled .
T he correet plaeement 01' th e speeies is still some what ope n to question ,
ilrhough it a ppears to be m ost sa tis fae torily ass ignable to th e P. adametzii series
llll nccou n t 01' th e simple penieil1i and th e funieul ose charac te r 01' its eolonies.
' I'he spee ies deseription is ee n te re d upon th e typ e stra in CBS 332.5 9 = CMI
1:11,B99 =ATcc 18, 309=I FO 6, 231 = QM 7,289=IJFM 5033, isolat ed from
.la pa nesc soil b y S. Ab e in 1953. It is a t present known only from th e typ e
suai n.

(j. Penicillium kaza chstanicum N ovobranova : fi g. 62 , colo u r plate 6, a nd

S EM plate 11, (top: ri ght) .

00
0 0 0 O
00 0 0 0
BOO

o 0
l.WJ1J.=J

10 jrrn

,~

Fig. 62. Camera lucida

drawings of Penicillium
kazachstanicum
Novobranova, ATCC 24,722.
A, Detailed drawings of the
penicilli. B, Mature eonidia.
C, Ropes of interlae ing
hyphae with eonidial
struetures arising from them.

Penicillium kezochstonlcum Novobranov o - Nov . Syst. Nlzhnlkh . Rost . 11: 225-226.

1974.
Novobranov a ' s origina l di agn osis Iollows, sino : litis ude q ua tely presenta th..
spec ies in question :
'Colonies upon Czapek's agar atta ining a diameter of 25 to 30 mm in 12-14 day
at room temperature, velvety, slightly raised , radiat ely wrinkled, more densely
text ured at margin than in the centre, slightly zanate in marginal areas, at f irst
white, becoming grayish-violet after 10-12 days. Slowly and sparsely sporing in
zones in ligh t gray-green shades . Exudate hyaline, colourless, Iimited in amount.
Reverse wrinkled , w ith centre brown and margin in hazel-nut shades, marg in in pale
sandy shade s. Odour indefinite . Penicilli very small , str ictly monoverticillate, with
the st ipes very short, smooth walled , arising as short branches from aerial hyphae ,
8.0-18.0 .tm by 2.0 -2.2 .tm . Phialides in c1usters of 5-6 elements in the verticil,
4.0-6 .0 .tm by 1.5-2.0 .tm . Con idia globose to subglobose, very small , 2.0-2.5 .tm
sometimes up to 3.0 .tm in diameter .'

Our not es follow:

Col onies upon C A (colou r plat e 62, A a nd a) a tta ining a di ameter of 35 m m
in 14 days a t room tem perat ure (20- 25 OC ), cons isting of a tou gh, closetex tured basal felt, with colony surface overlaid by a m ycelial network of ae rial
hyphae and ropes of hyp hae , giving to it a gra n ula r appeara nce, rad iat ely
wrinkled with sha llow fu rrows , a nd slightly ra ised in th e cen tre; pract ically
sterile, sho win g whitish cream colou rs ove r the entire su rface; ex udate and
odour lackin g; no coloure d pigm ent d iffu sin g in to th e su rrou nding agar ;
colony reverse (colour pIate 62, a) in cream shades; peni cilli strictly sim ple,
very sm all (fig. 62, A) , arising m ostly from rop es of hyphae , with short stipes,
usually ran ging from 10- 20 tut: in len gth by 1.8 -2 .0 f.Lm in di ameter , smooth
walled; phi alides in sm all ve rticils of2-6 eleme nts in the whorl , bottle shape d ,
mo stly 5 - 8 f.L m by 2.0 -3. 0 tui , terminating in a ve ry short, pointed neck ;
conid ia globose to su bglobose, usually 2 .4- 3.0 f.Lm in di ameter, with walls
smoo th or nearly so (fig. 62 , E , a nd SEM plat e 11, to p: right); con idial chains
short, d ivergent.
C olonies u pon CYA (colou r pIate 62 , B a nd b) attaining a di ameter of
25 mm in 14 days at room temperat ure , cons isting of a den se netw ork of rop es
of hyp hae with a leath ery appearance, irregularly folded a nd raised in the
colony centre; colony surface appearing ' we t ', in fleshy shades ; ex udate and
od our lackin g ; no coloured pi gm ent d iffu sin g into the su rro u ndi ng agar;
colony revers e (colour plate 62 , b) in salmo n pink shades to cream colou rs;
alm ost ste rile.
Colon ies upon MEA (colour plat e C and c) att a ining a diameter of 40 mm in
14 days at room temperature , cons isting of a close-t extured b asal felt , overlaid

by H network (JI' ropes and trailing hyphac, disponed in a zonu te pattcrn, wit h
1II1I I'!{ina l a rcas plano an d ce n tra l zo nes decp ly rud iatcly wrinkl cd ; thin,
1"1'l:glllar1y dissccted ma rgin su rroundcd b y a widc fringc 01' su b rne rgcd hyph ac
exic nd ing abou t :~ mm beyon d the aerial grow th; colony su rface appear ing
rucmbrunous ; sparse ly spo ring; exudate and odo u r lacki n g ; no coloured
pigrm:nt d iffu sing into th e su r ro u ndi ng agar ; colo ny rcv c rse (colour pia te 62 , e)
in cin na rno n sh ad cs ; penicilli esse n tially as d cscribed aboye , bu t more
rbu ndant ly produced , a r ising m ostly frorn trailing hyphae and from rop es 01'
hyph ae (lig. 62 , C ).
The species diagn osis is ce n te re d upon th e type st rain AT C C 24 ,722 = C IlS
1!). 74 = VKM F-1750 = C M I 174 ,720 = IJ FM 7112 , isolated from Pirus malus L.
xu rlacc in th e Alma-Ata region, Sov iet Soc ia list R epublic a l' K azach stan ,
USS R, by T. I. N ov ob ranova, July , 1967 . It is actually kn own on ly frorn the
I ype strain.
The co rrect placement al' this species a p pears to be m ost sa tisfac to r ily
lIssigna ble with in th e P. adametzii se ries on accou n t of its str ictly sim ple
pcnicilli , a nd the strong fu n icu lose characte r a l' th e colo n ies .

(;:1. Penicillium albo-cinerascens C halab uda: fig . 6 3 , coIo ur pI ate 63, a n d

SEM pI ate 11, (ce n tre: Ieft).
Penicillium albo-cinerascens Chalabuda (not Maublane) - Nov. Syst. Crypt. Herb .
Inst. Bol. Komarovii. Aead. Sci. USSR. 6: 166. 1950. See also M.M. Kulik, 'A
Compilation of Deserptons of New Penicillium Species'. Agre. Handbook No.
351. Agre. Res. Ser. U.S. Dep. Agric. p. 14. 1968.

Colonies u pon CA (co lour pi ate 63 , A a nd a) growing restrictedl y , attain ing

a di a m eter 01'25 mm in 14 days a t room te mper a ture (2 0 -25 OC ), co ns isti ng 01'
a tough , close-textu re d b asal felt , deep ly vclvety to lanat e in a pp ea ra nce , at
Iirst white, then becoming pale blue -green (Sguy, Pi. XXX, 445), radiat cly
wrin kled in marginal zo nes, a n d ra ised in the co lony centre; evenly m argin
su r rou nded by a n a r row fringe al' su bmerged hyphae exte nd ing less th a n 1 mm
hcyond the aeria l growth; with th e colo ny su rface ove rlaid b y a m ycclial thin
overgrowth al' closcly in te rwoven trailing hyphae and rapes al' hy phac,
rc maining pe rs iste n tly wh ite througho ut , influencin g the co lo ny a ppearance ;
poorl y spori ng, not influencing the colo ny aspect; odou r la cking ; ex u date
limi ted in amou n t , as small co lou rless drople ts enmeshed in the m yceli al felt;
110 colou red p igmen t di ffu sin g in to the su rrounding agar; co lony rever se
(colour pla te 63, a) in cream shades wit h a gre en cas t showing throu gh in the
centre; penicil1i str ictly simple 01' showing a n occasional bra nch (fig. 63, A),
with sho rt stipe s a r ising m ostl y from tra iling hyphae, ropes 01' hyphae (fig . 63,
E), 01' from strongly coiled ae rial hyph ae (fig . 63, D) rem in isce nt of th e traps 01'

o
0 0
O O O
0 0 0 80
C\
0
'O'G
Uw.l=J

10pm

/y

0
ty ~ Y
y

5i.:".fR
.

Pc

100 jim

.;''>--,

fR..

Fig. 63. Camera lucida

drawings of Pen;c;/!;um albo cinerascens Chalabuda, CBS
219.66. A, Detailed draw ings
of the penieilli. B, Mature
eonidia. C, Habit sketehes.
D, Coiled hyphae with
eonidial struetures arising
form them . E, Ropes of
inter lacing vegetative hyphae
with eonidial struetures
arising from them.

nemat ophagou s fu ng i, usu ally less th an 50 p,m in len gth by 1.8 - 2.5 p,m in
diamet er, wi th apices enlarged in a vesicle up to 3. 0 usi: in d ia me te r, smooth
walled ; phialides com paratively small, bottle shaped, in clus te rs of 3- 10
eleme nts in th e verticil, com monly 6-9 p,m by 2. 3- 3.0 p,m , ab ru ptly tap er ed
in a short , n arrow neck ; co nidia globose, u su ally 2.5 utt: in di amet er ,
delicatel y sp in ulose (Fig. 63, B, and SEM plate 11, cent re: left); con id ial
chains loosely parallel to tangled (fig. 63, C ) .
Colonies upon CYA (colour plat e 63, B a nd b ) duplicating in texture those
grown up on C A, but spreadi ng mor e rapi dly , attaining a diameter of 35 mm
in 14 days at room temperature (20 - 25 OC ), stro ng ly and deepl y ra diately
wri nkled, with conce nt rical ridge s in the colony centre, whi ch is also
irregul arl y fold ed in a crateriform patte rn, with surface cracked; better sporing
in pale blue-gre en shade s (Sg uy, PI. XXX , 44 5), espec ially in marginal a reas ;
eve n margin su rrou nde d by a narrow frin ge of submerged hyphae exte nd ing
less than 1 mm beyond th e ae rial growth; odo u r lacking; exudate as small

~Y)

~@@

@@
@
~

10pm

R.

Fig. 64. Camera lucide

drawings of Penicillium
sacculum Dale, CBS 229.61.
A , Detailed drawings of the
penicilli. B, Mature conidia .

hyph ae and rapes of hyphae, remaining persistently white ; od our a nd exudate

lncking ; no colou re d pi gment diffusin g into th e surro undi ng a ga r; colony
rcverse (colou r plat e 64 , a) uncoloured ; peni cilli cons isting of ve ry poori y
differen tiated structu res (fig. 64, A) with stipes short, ran ging fro m 10 lo
10 .tm in length by 2.5- 4 .0 tui: in d ia meter, septa te, m ore or less a rticula te,
smooth walled, arising m ostly fro m aerial hyphae or ra pes of hyphac; phialides
in small clusters or single, bottle shaped, 5- 7 .tm by 3.0- 4.0 uit, terrnin atin g
in a sho rt-pointed neck ; conid ia globose, 6- 7 .tm in di ameter , stron gly
cchinulate (fig. 64, B, and SEM plate 11, ce ntre : right) , brawnish colourc d;
conidial chains short, di vergent.
Colon ies upon C YA (colour plate 64 , B a nd b) growing rcstrictedly,
al taining a d iameter of 15 mm in 14 days at raom temperature , cons isting of a
lou gh , close- textu re d bas al feIt , m embranous , ra ised in ce ntral a rea, radiat ely
wr inkled, with surface funiculose , espec ially in th e ce nt re; eve nly margin
surro u nde d by a frin ge of su bmerge d hyphae ex te nd ing 1- 2 mm bcyond th e
acrial gro wth; almost ste rile , re maining persistently whi te cream colou red ;
odou r a nd exudate lacking ; no coloure d pigment d iffusin g into the surround ing agar; colony reverse (co lou r plate 64 b) in cream shades
C olonies upon M EA (co lou r plat e 64, C and e) spread ing broadly, atta ining

t1 ropl ct s e n mc shed in thc my ccli al lcl t; pu le nmbcr pigm c nt dilfu sin g into the
urro u nd ing agar ; colon y rev ers o (co lou r platc 6:1, b) in orange lo cin namo n
ha rlc; pcnicill i csse ntially as dcscribcd a b ovc ,
C olo n ies upon M EA (col our plato 6:i , C a n d e) grow ing m ore rapidly,
ru uini ng a c1iamet er 01' 50 mm in 14 day s al room tcmperaturc (20 -25 OC ),
etlllsisting of dense basal fel t ovc rla id by a close ly wov en network ofla n atc ami
lun icul osc hyphae lending a lanate appearance to the colony surfacc, planc,
wh h I ult s of white aerial hyphac in the ce n tre; thin evenly margin su r ro u ndc d
by a wid e fringe of subrncrged hyphae ext ending about 5 mm beyond th e acr ia l
ro wt h; heavily sporing throughout in blue-gray shades (Sguy , PI. XXX ,
111:1 , 188) ; exudate and odour la cking; no coloured pigment diffusing into the
au rro u ncl in g a ga r ; colony reverse (colour pIate 63 , c) in brown shades; pen icilli
cssc n tia lly a s des cribed aboye, but showing more divaricate penicilli.
'I'he species description is ce n te red upon the type strain CBS 219.66 = AT C C
11I ,:ln=vKM F-1037=IJFM 7051 , isolated frorn uncultivated soil in the
cuviro ns of Ki ev, USSR, b y T .V . Chalabuda in 1949 . It is at p rescnt kn ow n
o ldy rom the type strain .
' I'he co r rect placement of the species appears to be m ost sat isfactor ily
lssig-n able within th e P. adametzii series on account of its pred ominant simple
Iu-uicilli and the strong funiculose charac te r of the colonies , but the possibllitlca
I hal P. albo-cinerascens Chal abuda m ay represent a sp ecies tra ns itio nal het wcc n
11u: Monoverticillat a and th e Asymmetrica-divaricata Penicilli a, sUPPOI'lt:d
m.rinly by th e presence of divaricat e penicilli upon sorne su b st rat a ( M EA),
shou ld not be overlook ed .
l'cnicillium sacculum Sub-series , su b -ser ies nov .
Hyphae sterilia reptabundis, crassae, hyalinae, septatae; cum stipitibus brevis,
irregularit er ramosis, ramis ultimis phialidis ferentibus; phialidae dense compressae ,
brevia, plusminusve ovoideae; conidia singulares vel in catenis duorum, triorum;
asperula.
Species typicalis : Penicillium sacculum Dale.

The Penicillium sacculum sub-series is ad m itte dly artificial in concept a n d is

dcs igned, a s a m atter 01' convenience , to accommodate peculiar species which
ran not oth erw ise be satisfactorily plac ed in any of R aper a nd Thom' s (1949)
Penicillium series, namely Penicill ium sacculum Dale.
In 1914, Miss E . Dale described very briefly a fungus isolated from soil
which she tentatively regarded a s a new species 01' Penicill ium but did not n ame
it , The n ame P. sacculum was first published in Biourge' s Monographie (192 3) ,
who referred to it with a figure but without diagnosis . In 1926 , Miss Dale
published a short note giving n ames 1'01' three sp ecies 01' Penicillium which had

rcrn ain cd u nidcntified since 1914, one 01' the: num es bc ing P. .sacculum. T ho m
( 1930), alt er cxa mi n ing it , regarded Mi ss Da lc's spccies as Scopulariopsis repens
Bain ier. R aper and Thom (1949) assign ed P. .sacculum to Scopulariopsis on th e
b asis 01' T homs earl ier findings , a nd without cxa m in ing a nd stu dy ing any
fresh m at erial. This species wa s list ed by C hes te r a nd Thornton (1956) as
Scopulanopsis sacculum (Dale).
H u gh es ( 1953) h as sho wn th a t Scopulanopsis is n ot rel at ed a t all to Penicillium,
since th e two ge ne ra have entirely d iffere n t m ethod s 01' spo re formation . I n
Penicillium th e spores a re phiall ospores, while in Scopulariopsis th ey are
annellospores .
A ca re fu l exam inatio n 01' P. sacculum has fa iled to det ect a n nelatio n in th e
sporogenou s cells, wh ich on th e othe r hand show all th e characte ristics 01' true
phiali d es.
In 1961 G . Smith create d th e new genus Eladia for th is fun gu s, on the b asis
01' th e fact th at th e spo re- pro ducing cells wer e phialides.
In our op in io n thi s spec ies m a y be cons ide re d as a ve ry primitive form of
Penicillium, co ns isting 01' poorl y d iffere n tia ted sti pes, septate, m ore or less
a rticu la te, b earing monoverticillat e clu sters 01' phial id es, producing
consp icuously stron gly ech in ula te co n id ia.
The species shows a cer ta in relati on shi p with th e P. adametzii series on
account 01' the funicul ose characte r 01' th e co lon ies, espec ia lly when cu ltiva ted
on M EA .

64. Penicillium sacculum D al e: fig . 64 , co lo ur pi a te 64 , a n d SEM

pl at e 11 , (ce ntre: ri ght).
Penici//ium sacculum Dale - Ann . Mycol. Berln. 12: 33-62. 1914. See also Biourge's
Monographie, La Cellule 33: 323. 1923; Also E. Dale, Ann. Mycol. Berlin. 24: 137.
1926; also C. Thorn, 'The Penicillia ,' 1930; also Raper and Thorn, ' A Manual of
the Penicillia'. p. 840. 1949; also G. Smith, Trans. Brit. Mycol. Soco 44: 42-50.
1961.

Synonymies
Scopulariopsis sacculum (Dale) Chester et Thornton - Trans. Brit. Mycol. Soc o
39: 301-313. 1956.
Eladia saccula Smith - Trans . Brit. Mycol. Soco 44: 42-50. 1961.

Colon ies upon CA (co lour plat e 64, A a nd a) gro wing ve ry restrictedl y , mostly
sub me rge d, a ttain ing a di ameter 01' 15 mm in 14 da ys a t room tem pera ture
(20- 25 OC ), co nsist ing 01' a loose-textured basal felt , ve lve ty , whi te, somewhat
za nate, with th in margin largely su bmerged (4- 5 mm wide) ; sparin gly
spo ring, with co n id ial fru ctification s a r ising from loosely in terwov en ae r ial

--

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@) ~ @
~
@
~

10

jrrn

eR.

Fig. 64. Camera lucida

drawings of Penicillium
sacculum Dale, CSS 229.61.
A, Detailed drawings of the
penicilli. S, Mature conidia .

hyph ae a nd rop es of hyphae , remainin g persistently white ; odo ur a nd ex udare

lac:k in g; no coloured pigme nt di ffus ing in to th e surround ing agar; colony
n -verse (colour plate 64, a) uncoloured ; pe n icilli cons isting of vc ry poorly
rliff crentia ted structu res (fig . 64, A) wi th st ipes short , ranging from 10 lo
'lO /-l m in length by 2.5-4 .0 .tm in d iamet er , septate, m ore or less a rticulatc,
smooth wall ed , arising m ostly from aerial hyphae or ro pes of hyph ae ; phialidcs
in small clu sters or singl e, bottle shaped, 5-7 .tm by 3. 0-4.0 u sn , termi nating
i 11 a sho rt-pointed neck ; co n idia globose, 6- 7 .tm in d ia mete r, stro ngly
ech inulate (fig . 64, B, a nd SEM plate 11, ce nt re : right), brownish colourccl;
conidial chains short, di vergent.
Colonies upon CYA (colour pIate 64, B and b) growing re strictedly,
u rtaining a diameter of 15 m m in 14 days at room temperatu re, consisting of a
lou gh , close-textu re d ba sal feIt , me mb ranous, ra ised in cen tral area, radi ately
wr inkled, wit h surface funiculose, especially in th e ce ntre ; eve nly m argin
surrounde d by a frin ge of su bmerge d hyphae exte nd ing 1- 2 m m beyond th e
ncr ial growth; almost sterile, rem aining persistently white cream coloured;
odour a nd exu da te lacking ; no coloure d pigment di ffusin g in to th e surroundi ng agar ; colony reverse (co lour plate 64 b) in cre am shades
Colon ies upon M EA (colour plate 64, C and c) spread ing broadly , attaining

a di amctcr 0 1' 5:)-60 IlIIl1 in 14 day s al room remp crnturc (20-25 OC),
cons isting 01' a loosc-t cxturcd basal clt , vclv et y in a ppcu ru ncc, co nsp icuously
za na te, showing a thin ove rgro wth 01' sno w whi te, looscly int crwoven acrial
hyphae a nd rop es 01' hyphae from whi ch arise the abu ndan tly produced
con id ial structures, rem aining per sistently white; th e colony surfa ce plan c ,
with gro wing m argin surrounded by a wid e frin ge 01' su bmerged hypha
ex tend ing up to 5 mm beyond th e ae rial gro wth ; odour and ex udate lackin g;
no colou re d pigment di ffu sin g into th e surrounding agar; penicilli abu nda nt ly
produced throughout , esse ntially as described aboye; colony reverse (colour
plate 64 , c) in ora nge shades.
The species diagnosis is ce ntere d on strain c as 230 .6 1 (white variant),
isolated by W . Gams from Austrian soil. The othe r strains 01' P. sacculum Dale
kept in th e Internati onal Fungus Collect ions duplicate this strain in cult ural
and morphological characte ristics with th e exception 01' th e green colou r sho wn
by th ese forms in stead 01' th e persistently white colou r 01' strain css 230 .61 .
Smith's ori ginal di agnosis of Eladia saccula follows, sin ce thi s ade q uately
presented P. sacculum D ale:
' On malt agar growing moderately well, dull gray-green with white edge during
the growing period, gradually becoming brownish green, velvety, slightly wrinkled;
reverse grayish; colonies on potato agar growing somewhat more slowly, otherwise
similar; colonies on Czapek's agar thin and sparse, growing very slowly; on cornmeal agar growing slightly better than on Czapek's agar, with colonies definitely
arachnoid; on malt agar with 40% sucrose growing slowly but colon ies eventually
typical, gray-green, smooth velvety; superficial hyphae funiculose, hyaline, coarse,
septate, more 01' less artic late. 3.5 to 6 /lm in diameter.'

Penicillium vinaceum su b -se ries

Two spec ies are included here , Penicillium vinaceum G ilman el Abbott and P.
phoeniceum va n Beym a , characterize d by th e production 01' inte nse vin aceou s to
deep purple-red colours in th e colony reverse and di ffusin g into th e
surrou nd ing agar.
Both spec ies, whil e pos sibl y not closely relat ed gene tica lly, are however ,
cons ide re d together as a m atter 01' conve nience .

65. Penicillium vinaceum G ilman el Abbott: fig. 65 , co lour plate 65,

and SEM plate 11, (b ottom: left ).
Penicillium vinaceum Gilman et Abbott - lowa State College J. Sci . 1: 299.1927.
See also Ch. Thom, 'The Penicillia', pp. 195-196. 1930; also Raper and Thom,'A
Manual of the Penicillia', pp. 234-236. 1949.

~~~

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,

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100 JJm

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Fig. 65. Camera lucida

drawings of Penicillium
vinaceum Gilman et Abbott,
CSS 514.73. A , Detailed
drawings of the penicilli. S,
Mature conidia . C, Habit
sketches.

Colonies upon C A (colour plate 65, A and a ) gr owing rather restrictedl y ,

.u ta ining a diameter of abou t 25 mm in 14 days at room temperature
( ~W- 25 OC), consisting of a clos e-t extured basal fe!t , com para tively de ep , up to
~ mm in th e colon y centre, whi ch is depressed more or less in a cra teriform
pattern , with su rface deepl y and radiately furrowed ; with undulating m argin
xurro u nde d by a fringe of submerged hyphae ex tend ing abo u t 1 mm beyond
(he ae rial gro wth, and sho wing sorne funicul ose arra n geme nt of veget ative
hyph ae ; gen erally very slightly sporing throughout in pale gray- blue to pal e
gray -green sh ades (Sguy, PI. XXX, 450, and PI. XXVI , 400); with colony
su rface remaining persistently wh ite ; ex udate abu ndantly produced , as mi nu te
d ropl et s , in pale yellow to deep vinaceous shades, dep ending on th e stra in ;
odo u r faint or lacking; pale yellow coloured pigment diffu sin g slowl y into th e
surro u nd ing agar; colony reverse (colou r pIate 65, a ) in vinaceous sha des;
peni cilli (fig . 65, A) usu all y simple, seldo m branched, but occas iona lly
prod uc ing div erging rami suggestive of th e PenicilliumJanthinellum series; with
th e stipes of th e con idiophores sm ooth walled , com mo nly borne from ae rial
hyphae , short , seldom exceed in g 50 ust: in len gth by 1.5- 2.0 .tm in diam et er ,

with apices vcry slightly c n la rgcd ; phialldcs usunlly COJlsisling 01' a cluster 01'
2-5 ele rnc n ts in th e verti cil , 6.0 -7 .51t1ll by 1. :> -2.0 Itlll, tcrminat in g in a
conspicuous tape red neck , so rncwha t div crgent ; con id ia globoso to su bglob ose
when mature, with ends apiculate when young, 2 .0-2 .5 tun, occasionally up
to 3 tui: in diamet er , with walls appearing slightly irregu lar 01' in sorne strains
definitely roughened (Fig. 65, B, and SEM plate 11, bottom: left); co n id ial
chains very short a nd loosely divergent (fig . 65, C).
C olonies upon CYA (colour pi ate 65 , B and b) attaining a diamet er 01' about
20 - 25 mm in 14 da ys at room temperature (20- 25 OC ), con sisting 01' a clos e
textured and tough basal felt, duplicating in texture those gro wn upon CA, but
abruptly raised at the colon y centre in a crateriform pattern; exu da te very
abundantly produced , either as large topaz yellow drops 01' minute droplets;
colon y surface sulcated b y numerous radi al furrows, remaining persistently
white , and slightly sp oring, with undulating margin surrounded by a very
narrow fringe not exceed in g 1 mm beyond th e aerial growth ; arnber-co loured
pigment diffusing into th e surround in g aga r ; colony reverse (colour piat e 65,
b) in deep vin aceous shades; penicilli essentiall y as described aboye.
Colonies upon MEA (colour plate 65, C and e) attaining a diamet er 01' abou t
35 mm in 14 days at room temperature (20 -25 OC ), consisting 01' a looser
textured basal felt, with a wide transparent m argin , su rro u nd ed by a frin ge 01'
su b me r ged h yphae extend in g more than 10 mm be yond th e aerial growth ; no t
furrowed , plane , more 01' less floccose, with ce n tre slightly raised a nd fold ed ;
more 0 1' less zonate ; h eavily sporing in cen tral a nd subcentral zone s in
artemisia green shades (Sguy, PI. XXVII, 404) ; exudate and odour lacking;
colony rev er se (colour plat e 65 , c) in brown sh ades with a green cast show in g
through, rather than vinaceous ; penicilli as aboye .
The spe cies d escription is cen te re d upon strain C BS 514.7 3 = NRRL
2,063 = IJFM 7262 , isolated in 1945 by Prof. G .W . Martin frorn th e sidewall 01'
a tent in th e area 01' Hollandia, N ew Guinea, which represents an extre m e type
by producing conidia that a re consp icu ously ech in ula te. In colony ch a rac te ristics , this cu ltu re duplicates Gi lm an and Abbott 's type, originally isolated
frorn Utah soil, in 1927, CES 389 .48 = NRRL 739 = ATCC 10 ,514 = IJFM 5123 .

66 . Penicillium phoeniceum van Beyma: fig . 66, colour plate 66, and SEM

plate 11 (bottom : right).

Penicillium phoenice um van Beyma - Bakt. Parasitenk . Inf. Hyg. 1I Abt. 88: 136-137
1933. See also Raper and Thom, 'A Manual of t he Penicillia'. pp. 236-239 . 1949.
Colonies upon CA (colour plate 66, A) a tt a in in g a di ameter 01' 30 mm in 14
d ays at room temperature, consistin g 01' a tough, clos e-textured basal felt ,
sulcated by shallow furrows in a radial pattern , strongly raised in th e cen tre,

00

. 0 0 00
00 o O
O

000
00

00 O
B 00 0 O
O O 00 0
00 O

lLw.J..wJJ

\,

'~

10 prn
'<ji

1
\U

('-" ~~'~r;'.1 l;~~,

"~- ' C'-'I' 'S'"Io

~
~

R .

100 prn

dccpl y velve ty in appearance ; eve n m argin surrounded by l IHIIW' ,,1

su bme rge d hyphae exten d ing abo ut 1 mm beyond th e ae rial gro wth ; heuvll
spor ing zones in pale blue-gr een shades (Sguy, PI. XXX , H :I). 1t' 11I1
sporula ted a reas in white rosy colou rs; exudate and od ou r lack ing : el!'."!,
purple-red -coloured pigm ent diffusin g into th e su rro u nd ing aga r ; (: U l l l ll~'
reverse in th e sa me shades; peni cilli strictly sim ple (fig . 66 , A), with th c NII!'I'
uf th e con id iopho res usu ally erec t, arising primarily fro m th e su bstratu m UI' I hl1
basal felt , up to 200-250 /lm in len gth by 2.0-2. 5 /lm in diametcr, with wull
srnoo th 0 1' delicat ely rough en ed , with ap ices enla rge d in a vesicle up lo 4 .0 /UII
in di amet er ; phialides in clusters of 3- 12 elements in th e verticil , rou ghl y
parall el 0 1' slightl y divergent , mostly 11-1 5/lm by 2. 2- 3.0/lm , gradu all y
tap ering lo a narrow ed , long neck ; conid ia elliptica l to su bglobose, usuully
:~ . 0 - 4 . 0 tui: by 2.0- 3.0 /lm in di ameter, with walls sm ooth 01' finely granula r
(SEM pla te 11 , bottom; right, a nd fig. 66 , B); conid ial chains loose 01' tan glcd
(fig. 66, C ) .
Colon ies upon C YA (colour plat e 66, R a nd b) a ttaining a diameter of
30 mm in 14 days at room temperature (20 - 25 OC ), con sisting of a tou gh ,
close -tex tu re d basal felt , deeply velve ty to lanat e , deepl y ra diately fur row ed ,

with ce n tre rai scd and irrcgularly Ioklcd in a cra tcrifor m patt crn ; cven margin
surrounded by a fringe 01' su b rne rgc d hyph ae extc nd ing about 1 mm beyond
th e ae rial growth; heavily sporing es pec ially in m arginal arc as in pale bluegreen sh ades (S guy, PI. XXX , 443) , with th e remaining zones obs cured by
th e white ove rg ro wth 01' close ly interwoven ae rial hyphae ; odour faint,
indefinite ; ex udate lacking; a m be r-c olou re d pi grnent diffusing int o th e
su rro u nd ing agar; colony reverse (colour plate 66 , b) in pal e brown to
cin namo n shades , with th e colo ny stron gly pulling away a nd crac king th e agar ,
permitting ae ro bic cond itions underneath a nd allowing a b u nd an t sporu la tion
in th e colo ny reverse (colour plat e 66, b) ; penicilli esse n tially as described
aboye .
C olonies upon MEA (colour plat e 66 , C and e) attain ing a di ameter of
40 mm in 14 d ays at ro om temperature (20 - 25 OC ), cons isting of a less d ens e
basal felt , plane in marginal areas and fold ed and raised in th e cen tre, with
thin m argin surro u nde d b y a wid e fringe 01' submer ged hyphae exte nd ing
abou t 6 mm beyond th e aerial growth; vel vet y; hea vily spo ring in ar te m isia
gre en shades (Sguy , PI. XXX, 438); odo u r and exudate lacking; purple-redcolou re d pi gment diffu sing into the su rrou nd in g aga r; co lony reverse (colour
plat e 66, c) in deep purple-red sh ad es; penicilli esse n tially as described aboye ,
but with th e stipes 01' th e con id iop ho res m ostly arising from th e su bstra tu m and
from tr ailing ae rial hyph ae .
The spec ies description is ce n te red up on th e type strain CBS 249 .32 = CMI
40 ,85 8= ATCC 10,481=NRRL 2,070= QM 7,608=BKM F- 321=IJFM 5 122 ,
isolat ed from soot y m ould of a palm (Ph oenix sp. ) at Baarn , H olland , by F .H .
va n Beyma, in 1933.

Species of dubious position

Penicillium ellipticum Sasaki - J. Faet. Agrie. Hokkaido Univ. Jpn. 49: 151. 1950.
See also M.M. Kulik, 'A Compilation of Deseriptions of New Penicillium Species'.
Agrie. Handbook No. 351. Agrie. Res. Ser. U.S. Dep. Agrie. p. 37. 1968.

Penicillium ellipticum Sasaki is a species described in 1950 tha t is believed to be

closely related to th e spec ies 01' th e P. adametz ii ser ies . However , it is presented

her e as a species 01' dubiou s po sition since th e au tho r 01' th e presen t study was
not able to exam ine and study th e typ e stra in 01' th e sp ecies in question ,
because all attempts to ob ta in it from Internation al Fungus Collecti ons failed.
Therefore , it has not been included in th e key to sp ecies.
Sa saki ' s original description follows :
'Colonies upon Czapek's solution agar growing restrietedly, with eenter 1.5-3
mm deep, wrinkled, funieulose, margin white, about 1 mm wide, eonidial areas tea

Ill rJ

green (Ridgeway, PL. XLVII), azonate; exudate abundantly produeed as large,

eolourless drops; eolony reverse in olive -buff shades (Ridgway, PI. XL) to olive gray
or dark oliva gray (Ridgway, PI. LO; agar not eoloured. Conidiophores mostly arising
from trailing hyphae or ropes of hyphae, simple or oeeasionally ramified, with
phialides in cluster, smooth-walled, 2.0-2.5 p.m in diameter, and up to 200 p'm long
when arising from submerged hyphae, measuring up to 10 to 40 p.m in length when
borne as branehes from aerial hyphae; phialides 10-16 p'm by 2.5-4.0 p'm,
numbering 2 to 3 elements in the vertieil; eonidia elliptieal, delieately spinulose ,
3 .0-4.5 by 1.7-2.8 p.m in diameter; eonidial ehains parallel or divergent; sclerotia or
perithecia not produeed.
On Koji extraet gelatin growing abundantly, furrowed, mineral gray (Ridgway, PI.
XLVII) to deep grayish olive (Ridgway, PI. XLVI); eolony reverse dull citrine
(Ridgway, PI. XVI) to Kronberg's green (Ridgway, PI. XXX); gelatin Iiquefied,
colourless.
On potato growth abundant, furrowed, velvety, white with aerial hyphae.
Milk alkaline but not peptonized .'
(The above description was published without providing a Latin diagnosis).

The Ramigena ser ies

AII Penicillium species with con id iopho re s m ostly irregularly ramified , but
with each ramification bearing a terminal , wel1-defined simple penicillus were
included b y R aper a nd T ho m ( 1949) in this series.
OUTSTAN Dl NG FEATU RES

Colon ies gr owing usuall y ra ther restrict edl y , cons isting of a close-textured
basal felt , wi th su rface a p pearing velve ty or n early so ; fre quen tly p oorly
sporing during th e first 8-10 d ays ; sometimes spread ing broadly , tending to
be lanat e , rangin g from slig htly to heavily sporing .
The stipes of th e co n id ioph ores eithe r prostrate or ascend ing , arising
primar ily from th e su bs tratu m or from creep ing ae ria l hyphae , typi call y
ra m ified, w ith th e ramifications eithe r one celled (metula-like) or lon g a nd
sep ta te, co m monly unequ al in length , sim ple or rebranch ed , not producing
de fin ite a p ical ve rticils or m etulae .
Penicil1i typi cally simple , b orne te r mi nally on th e a pe x of the ramified
stipe s, but keeping always its monoverticillat e characte r , altho ugh rep eat ed
d iva ricate ram ifica tion at var ious levels p recludes a definite or ganizat ion .
C onidia co m para tively small, ranging from globose to stro ngly ellip tical ,
a nd frorn smooth to definitely ro u ghe ne d, dep ending upon th e spec ies .

K ey to species
Pag
B. Stipes of the conidiophores mostly ramified , occasionally twice, each
branch bearing a terminal and simple penicillus but not arranged as a
definite apical verticil of metulae .
RAMIGENA Series . . . . 186
1. Colonies growing restrictedly upon all substrata , mostly 15-25 mm in
diameter in 14 days.
a. Conidia defin itely elliptical, smooth-walled .
1'. Colony surface spor ing in gray-green shades; conidia strongly
elliptical to narrowly cylindrical , with rounded ends, not apicu late.
P. capsulatum Raper et Fennell .... 187
2'. Colony surface spor ing in blue-green shades; conidia elliptical
wi th ends somewhat pointed .
P. cyaneum (Bainier et Sartory) Biourge .... 189
b. Conidia globose, ovate , or slightly elliptical, w ith walls roughened .
1' . Conidia globose or nearly so; conidial chains divergent, not
form ing well-defined columns.
P. waksmaniiZa leski .. .. 191
2'. Conidia ovate to slightly elliptical; conidial chains parallel,
forming well -defined compact columns.
P. charlesii Smith .. . . 193
2. Colonies growing more rapidly upon most substrata , usually 45-50
mm in diameter in 14 days.
a. Conidia roughened , echinula te; colony reverse in vinaceous to
reddish to reddish brown shades.
P. velutinum van Beyma . . . . 195
b. Conidia smooth walled; colonies producing brigh t yellow pigment
in the reverse, diffusing into the surrounding agar. Velvety .
P. citrinum series . .. . 351
Isee The Asymmetrica-velutinasubsection)
c. Conidia smooth wa lled; colony reverse in orange shades ; deeply
floccose.
P. sartoryiThom . .. . 197

T he Monoverticilla ta - ramigena series includes a number of forms first

descr ibed by Bainier and Sartory (19 12,1 9 13) as species o fCitromyces and sorn e
add itional spec ies that have been described since th at tim e , in whi ch sim ple
peni cilli are charac te ristically produced , but with th e stipes of th e
con id iopho re s a rising as sho rt branch es at several septa fram cree ping or
asce n di ng fertile hyphae in a ramigenous pattern .

67. Penicillium capsulatum Raper et Fcnncll: fig. 67, colou r platc 67, and
SEM plat c 12, (top : lcft).

lw.J..l=1J

10

.:

jrrn

fR. ,
-,

~.~
~

100

jrrn

Fig. 67. Camera lucida

drawings of Penicillium
capsulatum Raper et Fennell,
CSS 301.48. A , Detailed
drawings of the penicill i. S,
Mature conid ia. e, Habit
sketches.

Penicillium capsulatum Raper et Fennell - Mycologia . 40: 528-530. 1948. See also
Raper and Thorn, 'A Manual of the Penicillia', pp . 242-244. 1949.

C olonies on CA (colour plate 67, A and a) growing restricted ly, attaini ng a

diameter of28 mm in 14 days at room temperature (20-25 OC ), consisting ofa
tough , close-textured basal felt , with surface plane, usu ally raised and folded in
(he ce ntre in a crate riform pattern , velvety or very slightly lanate in
appearanc e, overlaid by a white, thin mycel ial overgrowth of close ly
interwoven aerial hyphae , influ encin g th e colony appearance ; white , eve n
margin surrounded by a narrow fringe of submerged hyphae exte nd ing less
(han 1 mm beyond the aerial growth; slightly sporing throughout in pale bluegreen shades (Sguy , PI. XXX , 449), partially obs cured in zones by the thi n
rnycelial overgrowth of wh ite aerial hyphae ; exu da te and odour lacking; pale
ambe r colou re d pigment d iffusing into th e surrounding agar; colony reverse

 \IV

(colour pia te 67, a) un colou red 01' in palo yellow-grec n sha dcs ; peni cill i
typi call y sim ple , borne on stipes 01' va ryin g lcn gth (Ii g. 67, A a nd C ), and
occas ionally formin g clu ste rs, but cons iste n tly ret ainin g th cir mon overticilla tc
character; th e stipes a l' th e ca n idiaphores arise prim aril y frorn cree ping 01'
interl acin g aerial hyph ae (fig. 67, C), rangin g frorn very sho rt up to 100 p.m in
len gth by 2.0- 3.0 p.m in di am eter, smooth walled ; phialides in whorls,
rangin g from 4- 5 up to 8- 10 ele me nts in th e verti cil, u su al1y paral1el, mostly
8-11 p.m by 2- 3 p.m , terminatin g in a short , tap er ed neck; con id ia stron gly
el1ipt ical, with ro unde d ends, mostly 3.0 - 7.0 p.m by 2.0 - 2.8 p.m in diameter ,
smooth wa lled 0 1' nearl y so (SEM pIate 12, top : left) ; conid ial chains sho wing
cons picuo us ly lon g ca nnec tives, and adhe ring in lon g, well-defined colu m ns
(fig. 67, C ), a nd (fig. 67, B) .
Colonies up on C YA (colour pIate 67, B and b) atta ining a diameter 01'
35 mm in 14 days at room temperature (20-25 OC) , duplicating in texture
those grown up on C A, but with the colony surface deepl y wrinkl ed in a radial
pattern , a nd with the centre depressed in a crateriform way , a nd cons picuo usly
crac ked, zonate; medium sparing in pale blue-green sh ad es (Sguy , PI.
XXXI , 463); eve n m ar gin su rro u nde d by a wide frin ge 01' subme rge d hyphae
exte nd ing abou t 2 mm b eyond the aer ial growth; exudate a nd odo u r lackin g ;
pale ambe r -coloured pigm en t diffusin g into the surrou nd ing agar; colony
reverse (colour pIate 67, b) in yellow-cream shades; penicil1i esse ntially as
described aboye, but with stipes arising m ostly fro m trailing ae rial hyphae and
from ropes 01' hyphae.
Colonies upon M EA (colour plat e 67, C and c) attaining a diameter 01'
35 m m in 14 days at roo m te m pera tu re (20 - 25 OC ) , cons isting 0 1' a loaser
text ured felt , appearing deepl y velve ty, not furrowed , abo u t 2 mm deep in the
centre; heav ily spo ring throu ghout in deep blue-gr een shade s (Sguy , PI.
X XX , 446); gra wing margin wh ite , surrou nde d by a wide fringe extend ing
abo u t 2-3 mm beyond th e aeri al grow th; exudate and odo ur lacking ; ca loured
pigment not diffusin g into the surrou nd ing agar; colony reverse (colour pIa te
67, c) uncoloured ; peni cilli esse ntially as described aboye, but with th e stipes
u suall y longer , up to 200 p.m in len gth , ar ising primarily from th e substratum ,
fro m tr ailing hyphae 0 1' from rope s 0 1' hyphae .
T he spec ies description is cente red upon th e typ e stra in C BS 301.48 = C M I
40,576 = ATCC 10,402 = NR R L 2,056 = Q M 4,869 = IJFM 5120, isolated in th e
P an ama Canal Zon e fra m an optical instru ment by DI'. W. G . Hu tchinson in
1945.
Add itional strains representing this spec ies have been rep eat edl y recovered
from deteriora tin g military eq ui pme nt in tropical and sub-tropi cal countries,
thu s suggesting its probable wid e distribu tion in such areas.

11

fiU . Penicillium ey(weum (Baini cr el Sartory) Biourgc: fi g. 60, culuu

platc 68, a n d SEM plat c 12, (top: right).

R.,

~'

1" e 1V'
~\;~::&>~

j)l-.-.....J

100

jrrn

Fig. 68. Camera lucida

drawings of Penicillium
cyaneum (Bainier et Sartory)
Biourge, CBS 315.48. A,
Detailed drawings of the
penicilli. S, Mature conidia.
C, Habit sketches .

Penicillium cyaneum (Bainier et Sartory) Biourge - Monographie, Liste onomastique, La Cellule. 33: 102. 1933. Emend. Ch. Thom, 'The Penicillia', pp. 226-228.
1930. See also Raper and Thom, 'A Manual of the Penicillia', pp. 244-246. 1949.
Synonymy

Citromyces cyaneus Bainier et Sartory - Bull. Trim. Soco Mycol. France. 29:
157-161 .1913.

Citromyces musae Bainier et Sartory - Bull. Trim. Soco Mycol. France. 29:
154-157. 1913.

Colonies up on C A (colour plate 68 , A and a) gro wing restrictedl y , attaining

a diameter of 35 mm in 14 da ys at roo m temperature (20- 25 OC), con sisting of
a tou gh , clos e-tex tu red basal felt , not furrowed , with cent re com mo nly ra ised,

1- 2 mm dcep , dccply velvet y lo lan at e in appcnrn ncc from an ovc rg rowth 01'
white aerial sterile hyph ae inllucu cin g grca tly th e colony ap pcura ncc; azo na le;
m argin entircly surro u nde d by a very narrow fringc 01' su bme rged hyph ae
exte nd ing less th an 1 mm beyond th e ae rial gro wth; poorl y sporing in palc
gray-green shades (Sguy, PI. XXIX , 429) but with th e colony surface
remaining persistently white th rou gh out owing to th e m yceli al ove rg rowth;
exudate a nd odo ur lacking; coloured pigm ent not dil'fusin g into thc
su rro u nd ing agar; colony reverse (colou r plat e 68, a ) colou rless 01' becoming
yellowish in age ; penicilli sim ple (fig. 68, A) , with th e stipes a ris ing from the
su bs tra tu m 0 1' as branch es fro m ascending ae rial hyphae , smooth walled , up to
100- 200/lm in length by 2.0 -2 .8 /lm in d ia mete r , irregul arly bra nc he d,
occasionally bearing apica l gro u ps 0 1' penicilli ; phialides comparatively large ,
mo stly 13-16/lm by 3.0-4.0 iuu , terminating in a cons picuo us ly tapering
narrowed neck , u sually in clusters 01' 3- 10 ele me nts in th e ve rt icil; conid ia
elliptica l with mor e 0 1' less point ed ends , sm oo th walled 01' nearly so (fig. 68, B,
and SE M plat e 12, top: ri ght) , mostly 2.5 -5 .0 p.m by 2 .0-3 .0 p.m ; conid ial
cha ins loosely parallel 01' ta ng led (fig . 68 , C) .
Colonies upon C YA (colour plat e 68, B and b) attaining a d iam et er a l'
37 mm in 14 days at room te m pe ra tu re, cons isting 0 1' a tou gh , close- tex tu red
basal felt , deep velvet y in ap peara nce, stro ngly and deeply sulcated with
numerou s radial fur row s, rai sed in th e centre in a crate riform pattern , ove rlai d
by a thin m ycelial ove rg ro wth 0 1' closel y interwoven ae rial hyphae , infl uen cing
th e colony appearance; margin entire su rro u nde d by a very narro w fringe of
su bme rged hyphae , exte nd ing less th an 1 mm beyond the ae rial gro wth ;
spo ring abu nda ntly in deep gree n shades (Sguy, PI. XXVIII , 4 16) in
m argin al areas, with non-sporulating a reas remaining persistently colou re d in
cream shades owing to th e m ycelial ove rg ro wth; exudate a nd odo u r lacking;
am be r-co lou red pi gm ent diffusin g into th e su rro u nd ing agar; colony reverse in
yellowish sha des (colour plat e 68, b); penicilli essentially as described aboye.
Colon ies upon M EA (colour pIate 68, C and e) attain ing a d iamet er 01'
40 mm in 14 days at roo m temperature (20 -25 OC), looser tex tu re d than the
preceding , deepl y velvety to la nat e , somewhat deepe r in th e ce ntre, up to
3 mm thick , overla id by a m ycelial overgrowth 0 1' loosel y interwoven floccose
ae rial hyphae influencin g the colony ap peara nce; thin , eve n m argin
surrou nde d by a fringe 0 1' submerged hyphae exte nd ing abo ut 2 mm beyond
th e ae rial growth; heavily spo ring in th e colony cent re in blue-gr een shades
(Sguy, PI. XXX , 443) ; exu date lacking; odo u r faint , aro matic; pale am be rcoloured pi gm ent d iffusin g into th e su rround ing agar; colony rever se (colour
pIate 68, c) in yellow ish sh ades; peni cilli esse ntially as described abo ye , but
with th e stipes eve n longer th an on C A, sho wi ng fewer phialides in th e ve rticil.
T he species description is cente re d upon th e typ e strain CBS 3 15.48 = CM I
39, 744 =ATCC 10,432= NRRL 775=IFO 5,33 7 =QM 7,516=IJ FM 3759, isolat ed
from soil by Ba inier in 1913 .

69. Penicillium uiaksmon ii Zalcski: fig. 69, colou r plat c 69 ,

pla tc 12, (ce nt re : le n ) .

/l ll el

llH

!\,t i-t

J;~V

~~~. .

::=1?
(" 1'..'

"'T iOOtf "~~ ' :t~1(t;<~j~tt

/iJ?"., ..
F'

':fu1

: ..

.'t- . o

Fig. 69. Csmom lil e/el,

drawings of Penlclll/IIIII
waksmanii Zaleskl,
230.28. A , Detailed drowln
of the penicilli. B, Mat ur
conid ia. e, Habit sketchus.

Penicillium waksmani Zaleski - Bu!. Acad. Polonaise Sci. Math . Nat. Sr. B, pp .
468-469. 1927. See also Ch. Thom, 'The Penicillia', pp. 230-231; also Rapar and
Thom, ' A Manual of the Penicillia' , pp. 246-248. 1949.

Probable synonymies
Citromyces affinis Bainier et Sartory - Bul!. Trim . SOC oMyco!. France. 28: 39-43.
1913.
Citromyces brevis Bainier et Sartory - Bull , Trim . Soco Myco!. France. 28: 43-45.
1912.
Citromyces minutus Bainier et Sartory - Bull. Trim . Soc oMyco!. France. 29: 137144. 1913.
Citromyces ramosus Bainier et Sartory - Bul!. Trim. Soco Myco!. France. 29:
144-148. 1913.

C olon ies upon C A (colour plut e (;9, A and a) growing mod cratcly , auainin g
a diameter 01' 40 1II111 in 14 days at room tcmperaturc (20 -25 OC), co nsisting 01'
a close-textured basal ICIt, ovcrlaid by a clos cly wovcn nctwork 01' deli cat e
hyphae, giving a deep vclvety appearance to the colony suracc; thin margin
surrounded by a fringe of submerged hyphae extending about 2 mm beyond
the aerial growth; colony surface deeply furrowed in a radial pattern, raised in
the centre, appearing crateriform ; heavily sporing throughout in deep bluegreen shades (Sguy , PI. XXX, 447), obscured in sectors by the thin
overgrowth 01' white sterile aerial hyphae; exudate abundantly produced, as
large yellow drops; odour lacking; pale amber-coloured pigment diffusing into
the surrounding agar; colony reverse (colour piate 69, a) in bright yelloworange shades; penicilli monoverticillate , 01't en appearing singly, sometimes in
irregular clusters 01' 2-3 01' more elements, borne on separate branches and
retaining their individual monoverticillate character (fig. 69, A and C), with
the stipes arising from the basal felt as ascending branches, ranging from very
short to 100-200 tm in length by 1.5-3.0 tm in diameter, with apices
somewhat enlarged in a vesicle up to 3.0 tm in diameter; phialides
comparatively small, in compact clusters 01' 6-10 elements in the verticil,
mostly 7-11 tm by 2.5-4.0 uit , terminating in a short, narrowed neck;
conidia globose to subglobose, mostly 2.5-3 .0 tm in diameter, delicately
roughened (fig . 69 , B, and SEM plate 12, centre : left); conidial chains
divergent, becoming tanglcd, 50-100 tm in length (fig . 69, C) .
Colonies upon CYA (colour piate 69, B and b) attaining a diameter 01'
55-60 mm in 14 days at room temperature, duplicating in texture those grown
upon CA, but strongly and irregularly furrowed, raised in the centre in a
crateriform pattern, and cracking; heavily sporing in the same blue-green
shades (Sguy, PI. XXX, 447) as the preceding, especially in marginal areas,
but conidial areas obscured in central areas by the thin overgrowth 01' white,
closely woven aerial hyphae overlaying the colony surface; thin, even margin
surrounded by a wide fringe 01' submerged hyphae, extending about 3-4 mm
beyond the aerial growth; odour faint, aromatic; exudate abundantly produced
as large, deep amber drops ; amber-coloured pigment diffusing into the
surrounding agar; colony reverse (colour plate 69, b) in orange to cinnamon
shades; penicilli essentially as described aboye .
Colonies upon MEA (colour plate 69, C and c) attaining a diameter 01'
50 mm in 14 days at room temperature (20- 25 OC), consisting 01' a looser
textured basal felt, conspicuously zonate, overlaid in sectors by a thin mycelial
overgrowth 01' white floccose aerial hyphae; colony surface plane, slightly
furrowed in the centre; thin, even margin surrounded by a wide fringe 01' submerged hyphae extending 10 mm 01' more beyond the aerial growth ; heavily
sporing throughout in gray-green shades (Sguy, PI. XXIX, 429); exudate
and odour lacking; coloured pigment not diffusing into the surrounding agar;

I ~ ;l

oony rev crse (colour plat e 69 , e) ;n pu le yellow sha dcs : pcn ici11 cssc n t ially a
:)cscribed abovc but more co n unon ly show ing' ramifi cati on s in a terminal
c rticil and co n id ial cha ins lon gc r , up lo 100 ~1Il 01' m ore , tan glcd.
Spcc ics d escription is cen tc red upon the type stra in C BS 230 .2B = C M I
:19 ,746 = AT C C 10,516 = NRRL 777 = 11'0 7, 7'3 7 = QM 7,681 = IJFM :1757, isolat cd
rom forest so il in Pol and by K . Zal eski , in 1927 .
Penicillium w aksmanii a ppe a rs to be clos ely related to P. corylophilum Dierckx , both
in its basic cu ltu ral ap peara nce a nd in th e ge neral m orphological charactc rs ol'
its penicilli. H owe ver , in th e former species the penicilli a re mure regul arly
mon overticill a te , a nd th e co nid ia a re globose and co ns p icuous ly rou gh en ed ,
wh ile th e la tt er species shows smooth -walle d, elliptica l to su bglobose co nidia,
Accord ing to Raper a nd Thom ( 1949) th e tw o species sh ould be regarded as
close ly rel ated , a n d as p ossibly forming a bridge between th e M on overticillata
a nd th e Penicill ium citrinum series.

70 Penicillium charlesii Smith: fig. 70 , colou r plat e 70 , a nd SEM piate 1~,

(centre: ri ght).

10 urn

iR.

P /Jr
c-L

100 J.lm

Fig. 70. Camera lucida

drawings of Penicillium
charlesii Smith, CBS 304.48.
A, Detailed drawings of the
penicilli. B, Mature conidia .
e, Habit sketches.

Penicillium charlesii Smlth - Trans. Brlt. Mycol. Soco 18: 90-91. 1933. See also
Raper and Thom, 'A Manual of the Penicillia', pp. 248-250. 1949.

Colonies upon C A (colour plat c 70 , A and a) att ainin g a diam eter 01' 38 mm
in 14 days at room temperature (20- 25 OC), consisting 01' a tough , closetextured basal felt , deepl y velve ty in a ppeara nce to sublanat c , strongly rai sed
in th e ce ntre up to 3 mm deep, with a depression in a crate riforrn pattern ; eve n
m a rgin su rro unde d by a fringe 01' submerged hyphae exte nd ing about 2 mm
beyond th e ae rial growth; colony su rface radiat ely furrowed, rem aining
persistently sno w white throu ghout ; m edium sporing throughout , heavier in
submarginal are as, in a rte m isia green shade s (S guy, PI. XXIX , 425) , not
influencing th e gener al colony appearance; exudate and odo ur lacking; no
coloure d pigmen t diffu sin g into th e su rro u nd ing agar; colony reverse (colour
pIate 70, a) strong ly pulling away and cracking th e ag a r, allowing th e air to
pen etrat e underneath and a good sporu la tion, othe rwise colourle ss; peni cilli
sim ple (fig. 70, A), with stipes arisin g from cre eping or closely interwoven
ae rial hyphae , sim ple or variou sly ramified , occasionally showing fairly
com pact clus te rs 01' m etula-like branch es , eac h ramification bearing a
monoverticillate peni ciJlus ; th e stipe s 01' th e conid iopho res are variable in
length , ranging from very sho rt up to 100 Ilm or more by 2.0-4 .0 ust , with
apices enlarged in a ves icle up to 4.5 Ilm in di amet er, smooth wall ed (r eported
as slightly rough en ed by G . Smith) ; phialides in com pac t clus te rs, m or e or less
parallel , numbering 6- 12 eleme nts in the ve rticil, m ostl y 8-10 Ilm by
2.9-4 .0 psi , terminating abruptly in a short-pointed neck ; conidia ovate to
su bglobo se, deli cately roughened (Fig. 70 , B, and SEM pIate 12, ce ntre:
right) , mostly 3.0- 4. 0 psi: by 2.5-3 .0 tui ; conid ial chain s producin g welldefin ed colu m ns (fig. 70, C) .
C olonies upon CYA (colour plat e 70, B and b) attain ing a di amet er 01'
28 mm in 14 days at room temperature (20-25 OC ), consisting al' a tough ,
close-t extured bas al felt , velv et y in appearance , overlaid by a thin mycelial
ove rg ro wth 01' white , ste rile, closely interwoven ae rial hyphae, influen cing
gre atly th e colony appe arance; colony su rface strongly wrinkled , sulcated by
numerou s, deep , radial furrows , with centre raised in a crateriform pattern ;
undulating margin surrounded by a narrow fr inge a l' submerged hyphae
exte nd ing less th an 1 mm beyond th e aerial gro wth; colonies strongly pulling
away and crac king th e a ga r , remaining persistently white owing to th e thin
m ycelial ove rgro wth ; slightly spo ring throughout , not influen cing the colony
appearance ; odour lacking ; exudate not obs erved ; amber- coloured pigm ent
diffusin g into th e su rrou nd ing agar; colony reverse (colou r plate 70, b) in
cinn namo n shades ; peni cilli essentially as described aboye.
C olonies upon M EA (colour plate 70, C and c) attain ing a di ameter 01'
45 mm in 14 days at room temperature (20-2 5 OC), cons isting 01' a close-

195

textured basal fclt , co ns picuous ly zonu tc, slig hlly raised and sulca ted by
sha llow radial Iurrow s, a p pearing vclvery, with thin , even rnargin surrounded
by l wide frin ge 01' su b rne rge d hyphae cx ic ud ing 5 mm or m ore beyond the
ncr ial gro wth; colony su rfacc rcmaining pe rsi ste n tly snow white in colou r;
spo ring spa ringly, not influen cing the co lo ny as pec t; od ou r faint , indefini te;
cxudate lackin g; pale a m be r-c oloured pi gm ent diffusing into th e su rro u nd ing
agar; colo ny rever se (colour plat e 70, c) in pal e cre a m sh ad cs; pcnicilli
essc n tially as d escr ibed aboye but with stipe s arising mostly frorn su b me rgc d or
ascc nd ing ae rial hyphae , co m monly 200 ,tm or m ore in len gth , loosely
ramified , with penicilli u su all y well separated a nd p roducin g co nid ial chains in
lon g narrow well-defined colu m ns 100- 200 ,tm in len gth .
T he species d escription is ce n te re d upon th e typ e strain CBS 304 .48 = C M I
10,232 = AT C C 8, 730 = NR RL 1,887 = Q M 6,83 8 = IJFM 3756, isolat ed from Ital ian
mai ze (Zea mays L. ), by G. Sm ith in 1932.
St ra ins showing th e general cult u ral characte ristics a nd details al'
morphology 01' Penicillium charlesii Smith a re occasionally isolat ed from soil an d
[rorn organic su bs tra ta ex pose d to soil co ntam inatio n, a s fabrics, tents am i
othe r military equipmc n t undergoing deterioration in trop ical a nd su btro pica l
zones .

71. Penicillium velutinum v an Beym a : fig. 71, co lo ur pl ate 71 , a n d SE M

pl ate 12 , (botto m : le ft ) .
Penicillium velutinum van Beyma - Bakt. Parasitenk. Inf. Hyg. 11 Abt. 91: 353. 1935.
See also Raper and Thorn , 'A Manual of the Penicillia', pp . 250-251. 1949.

Colonies upon CA (co lour pl at e 71, A and a) growing fairly rapidl y,

a ttain ing a di amet er al' 35 m m in 14 days at ro om tempe rature (20-25 OC),
consisti ng 01' a tou gh , close-textured basal felt ove rlaid by a m yceli al
ove rgrowth 01' closely woven , fin e aeria l hyphae p ro duc ing an almost lan at e
a ppeara nce; colon y surface radiately a nd irregul arly wrinkled ; th in, even
margin surro u nde d by a fringe 01' su bmerged h yphae ex te nd ing abo u t 1- 2 mm
beyond th e ae rial grow th; heavily spo ring throu gh out , es pecially in m a rgin al
zones, in gree n shades (Sguy, PI. X X IX, 427); conid ia l areas parti all y
obscured b y th e m ycelial ove rg ro wth, es pec ia lly in centra l a nd su bcen tra l
areas, remaining per sistently white or d irty white in th e mentioned are as;
exud ate lacki n g; odo u r p ron ounced , suggesting mi stl et oe (Visc um album L. );
no co loured pigme nt d iffu sin g into th e su r ro u nd ing agar ; co lony reverse (co lou r
pla te 71, a) in cinnamon shades ; penicilli sometimes st rictly simple (F ig. 71,
A), u sually ramigenous and irre gu la rly ramified , ra nging from a te r m inal
clus ter al' m etula-like b ranch es to one, two , or more asymmetrica lly arranged
elemen ts, unequal in lengt h , appearing indep enden t and retaining th ei r mono-

fR.,

\':\1
l.--.....J

100 jirn

Fig. 71. Camera lucida

drawings of Penicillium
velutinum van Beyma, CBS
250.32. A, Detailed drawings
of the penicilli. B, Mature
conidia . e, Conidial
structure. D, Habit sketches.

verticillate cha racter, occasionally a rising directly fro m th e substratum, mostly

50- 100 .tm in len gth by 1.5- 2.0 .tm in di ameter , sm ooth wall ed ; phialides
comparati vel y sm all , usu all y in clu sters of 3-8 or 10 ele me nts in th e verticil,
m ostly 5.0- 10 .tm by 2. 5- 3.0 .tm, terminating abruptly in a sho rt-poin ted,
narrowed neck ; conidia globo se or nearly so , abo ut 2.8- 3.5 .tm in diameter ,
with wall s con spicuou sly ro ughe ne d (fig . 71, B, a nd SEM plat e 12, bottom:
left) ; con id ial cha ins up to 100 .tm in length , usually tangled, some times
loosely parall el (fig . 71, D ).
Colonies upon CYA (colour pIate 71, B a nd b) attaining a di ameter of
66 mm in 14 days at roo m temper ature (20- 25 OC ), duplicating in cultural
ch aracteristics a nd det ails of morphology th e previously described spec ies , but
m or e irregu la rly fold ed th roughout, with th e othe rw ise abu ndantly sporing
areas tot all y obsc u red by the overl ayin g m ycelial growth com po sed of closely
int erwoven floccose , white , sterile aeri al h yphae, lending a lanat e appe aranc e
to th e colony su rface; exudate lacking ; odo u r strong, su ggesting a m m on ia; no
coloured pigment d iffusin g into th e surrou nd ing agar; colony reverse (colour

1 ~I I

plnt c 71, h) in cin na mo u shadc s ; pcui cill i cHHI:nlial1 y as dc scribed aboy e.

C olo n ics lIpOIl M EA (colour plutc 71, C an d e) sp n :ad illg as upon C Y A ,
attainin g a diamct cr 01' 66 111111 in 11 day s al J"(HlIlI tcrupcr.uure (20 -2 5 OC ),
co nsisting 01' a close-tcxtu red basal lclt , uppcarin g d ccply velv ety lo lanat c ,
owin g to th c ove rlay ing n etwork 01' fine lloccosc acrial hyphae , inllu cn cin g
I-; n :a tly th e co lo ny appcarance , ami obs curin g parti ally th e othe rw ise heavily
sporing arcas ; con id ial areas in deep olive grecn shades (S gu y , PI. XXVIII,
'116); odour faint , indefinite ; ex u da te lacking; no coloured pigment c1iffusin g
into th e su r rou nd ing agar; colo ny reverse in orange shades (colour plate 71 , e) ;
pcnicilli essentiall y as described ab oye .
Th e species description is ce n te red upon th e typ e strain C BS 250.32 = C M I
40 ,571 = ATCC 10 ,510 = NRRL2 ,069 = QM7,686 = VKMF-379 = IJFM3762, isolated
Irorn sputum by F .H . van Beyma in 1932 . It is ac tu ally kn own only frorn Ihe
I ype species.
This spe cies was placed by Raper and Thom (1 949) in th e M onoverticillata rarnigena series on th e b asis 01' its penicilli , not presuming a close relation ship
lo othe r members 01' th e seri es . In contra st to th e other spec ies assigned hcrc ,
which are rather slow-gro w ing, P. velutinum is charac te rize d b y its very rapidly
a nd broadly spreading colonies.

72 . P enicillium sartoryi Thom: fig . 72, colo u r plate 72 , a n d SEM plat e 12 ,

(b o tto m : ri ght) .

Penicillium sartoryi Thom - in 'The Penieillia', p. 233. 1930.

Synonymy
Citromyces subtilis Bainier et Sartory - Bull. Trim. Soe. Myeol. Franee. 28: 46.
1912.

C olon ies upon CA (colour plate 72, A a n d a) growing rather restrictedly ,

a tta in in g a di amet er 01' 35 mm in 14 days a t room temperature (20-2 5 OC ),
consistin g 01' a c1ose-t extured basal felt , deepl y velv et y to su bla n a te, with
margin abrupt, surrounded by a very narrow fringe 01' submerged hyphae
exte nd in g less th an 1 mm beyond the aerial growth; col on y surface remaining
persistently sno w-wh ite throughout , irregularly sulcated with shallow furrow s,
a nd raised in th e ce ntre ; slightly sporing , es pecially in m arginal zones in pal e
blue-green sh ades (Sguy , PI. XXX , 438 ) ; ex u da te a nd od our la cking; no
colou re d pi gment diffusing into th e surrounding agar ; colony rev erse (colour
piate 72, a) uncoloured, with a reddish cast showing through ; penicilli
com m on ly str ictly simple or ramigenous with stipes arising either directly from
tr ailing aerial hyphae 300-400 11m long by 2 .0 11m in di amet er , or as branches
from aerial hyphae 100 -200 11m long, variously ramified with th e secondary

o 0 0

O O O
OO

O O
O

00 () Q

OB

00
O
O

O
~

10

jrrn

)11'1"1
L--......J

100

jirn

fR.

Fig. 72. Camera lucida

drawings of Penicillium
sartoryi Thorn , CBS 172.44.
A, Detailed drawings of the
penicilli. B, Mature conidia .
C, Habit sketches.

ramificati ons some times resembling apica l ve rticils of m etul ae , eac h te rrnin ating in a m onoverticillate penicillu s (fig . 72, A) ; phialides in clu sters of 3- 8
ele me nts in th e vertici l, mostly 9 - 13 p.m by 2. 4-3 .8 psi , te rm in ating in a
sho rt-po inted narrowed neck ; conid ia ellip tica l to su bg lobose, m ostly
2.5-3.5 p.m by 2.0-2 .5 p.m in di ameter, with walls smooth or nearly so (fig.
72, B, a nd SEM pIate 12, bottom: ri ght) ; co nid ial chains forming par all el or
loose colum ns (fig . 72, C ) .
Colonies upon CYA (colour pIate 72, B and b) a ttain ing a d iameter of
40 mm in 14 days at room temper a ture (20 - 25 OC), cons isting of a closetextured basal felt, deepl y velvet y , ra di ately wrinkled, with ce ntre depressed in
a crat eriform pattern ; eve n m argin surro u nde d by a ve ry narrow fringe of sub merged hyphae exte nd ing less th an 1 mm beyond th e ae rial gro wth; heavil y
sporing in subcentral are as in deep blue-gr een shades (S guy, PI. XXX , 44 7);
non-sporulating areas remainin g sno w-wh ite in colou r; exudate lacking; odo ur
faint , indefinite ; amber-col oured pigment di ffusin g in to th e surro und ing agar ;
colony reverse (colour pIate 72, b) in yellowish shades; penicilli esse nt ially as
descr ibed aboye .

I ~) \

;olon ies up on M EA (co lour plntc 7'2,

a m i e) alt ainin g a di nmct er 01'
in 14 days a l 1'00 111 tc mperu ture (~() - ~~ OC), co nsislin.{ 0 1' a looscr
icxtu rcd basal k it, co nspicu ously zo nutc , with (hin ma rgin su rrou ndc d by a
wid c Irin gc 01' su bmergcd hyph ac ex te nd ing IIp to 4 m m beyond th e acrial
g-l'Owth, a ppcar ing vclvet y, planc , with lllfl s 01' w hitc, floccosc ae r ia l hyph ae
ovc rlaying th e colony surface in secto rs; heavily spor ing throu gh ou t in dcc p
blue-green shades (Sguy , PI. XXX, 443); ex udate lackin g ; odo u r faint ,
aromatic; pal e am be r co lou re d pi gm ent di ffusin g in to th e su r ro u nd ing agar;
colo ny reverse (co lou r pla te 72 , c) colou rless to pale yellow; penicilli esse nt ially
as descri bed a boye, with th e stipes arising primarily fro m th e substratum 01'
Irorn trailing hyphae , co n id ial chai ns formin g we ll-defi ne d co mpact to looscly
parall el colu m ns.
The species descri pt ion is ce n tere d upon th e st ra in CB S 172.4 4 = IJF M 51 21,
isolat ed as a cu ltu re co ntam inan t .
T he m embers 01' th e M onov erticilla ta -ramigen a se ries regul arly occur in
soil, and may be isola ted also from a va r iety 01' organ ic su bstrata u ndcrgo ing
slow de com pos ition in the field . They h ave been encounte red freq uc n tly
a mo ng the m oulds iso la te d from fab r ics, optical instru ments, a nd mili ta ry
cq u ipm en t d et eriora ting in tropical a nd su btropical a reas, espcc ia lly P.
capsulatum th at wa s re peatedly encou ntered fro m the se sourccs a nd rcg ions,
hen ce bein g beli eved to be w idely di stri buted in th e soils 01' war me r a nd hum id
cou n tries . P. waksmanii Zales ki a nd P. charlesii Smith, we re simila rly isolal cd
fro m det eri orating mate rial.
4~ 1II111

Species of dubious position

Penicillium vitis Novobranova - Nov. Syst. Niz. Rast. 11: 225-234. 1974.
Penicilliu m vitis N ovob ranov a is a species d escribed in 1974 th at is b eli ev ed to
be closely relat ed to th e R amigena series . But since th e author 01' th e present
stu dy was n ot able to exami ne th e typ e strain 01' the species, b ecause all
a tte m pts to ob ta in it fro m Interna tion al Fungu s Collec tions fail ed , it is
p res en te d here as a sp ecies 01' dubious position a nd is not in cluded in th e key to
spe Cles.

Novobranova's original diagnosis follows.

Colonies on CA attaining a diameter of 10-20 mm after 24 days, irregularly raised
and folded, sporulating very scarcely and slowly, at first white, with centre sandy
coloured, becoming gray-green as sporulation proceeds, with central area showing
deep red shades, later becoming gray-green, with zones in ash-gray colours, and
margin gray-green. Exudate scarce or lacking; reverse folded, with agar cracking
underneath the colony, at first cream coloured , then becoming Iight grayish

violaceou s, showing little spots deep red in colour, w ith centr e in dark brown
shades. Penicilli simple, wi th tangled chains of conidia. Conidiophore of the
ramigena type , smooth walled , 10.0-30.0 (60.0) .tm by 2.0-2.5 .tm, with apices
inflated; phialides in verticils of 5-8 elements, slightly divergent, 6.0-9 .0 (11 .0) .tm
by 1.8-2.2 .tm. Conidia subglobose, smooth walled , 2.0-2.5 .tm. It belongs to the
Monoverticillata-ramigena. No related species are known.
Type: isolated from stored grapes, in the Alma -Ata region , USSR, 16-X-1967 by
T.1. Novobranova.
(Translated from the Russian by Prof . L. Vilas).

"'/1

hapt cr 4

The Asymmetrica

RAP ER ANO THOM (1949) IN CLUOEO I N THI S SE CTI ON AL L T H E PE NI CILLI A WIII GII
pr oduced two- or three-stage branched penicilli , in whi ch th e ramifi cation
sys te rn was typically irregular, on e side d , or asymmetrical.
T he ge ne ra l typ e 01' penicillus d escribed is 01' an arb itra ry characte r,
co rnmon to seve ra l more or less well-defined groups , co nside re d by th e authors
mentioned as su b-sectio ns . Each 01' th ese su b-sec tions em b raced two or more
se ries whi ch were beli ev ed to represent natural grou ps 01' spec ies. Fiv e su bscctio ns we re recognized and are m aintained here as fol1ows:
DIVARICATA . Charact eri zed by th eir stro ng ly divaricat e or bi fur care
penicilli , co m mo nly presenting th e a ppe ara nce 01' irregular and a pical clus te rs
01' penicilli ret aining alw ays th eir monoverticill at e characte r.
VELUTINA. Grouping spe cies producin g penicilli one - or two -stnge
branched , es pecially characte rized by th e producti on 01' ab u ndan t co nid ial
struc tu res with th e stipes 01' th e con id iop ho res a rising primarily in a dense
stand from th e substratum or the ba sal felt , giv ing a velvety a ppeara nce lo th c
colony surface.
LANATA . G rouping species producing penicilli similar to th e preceding
gro u p, but charac te rize d by th e producti on 01' a loose-textured network 01'
ae rial hyphae , from w hich a rise largely the stipes 01' th e conid iophores,
pr esentin g a lan at e or floccose overa ll as pec t.
FUNICULOSA. Grouping species producing penicilli , as previ ousl y , but
with th e m yceli al ove rg rowth sh owing definite aggre ga tes 01' m yceli a or rop es 01'
hyphae or fu nicles, with th e stipes 01' th e con id iophores com m only arising a s
shor t branch es from su ch furrieles .
FASCICULATA. Grouping species producing penicilli sim ila r in pattern to
th e preceding , but arising ch aracteristicall y from syn ne mato us co nid iop ho re s.

Asymrnetrica-divaricat a su b -sec tio n

The penicilli are stro ngly divaricat e in most species, co nsisting 01' an apical
gro u p 01' irregul arl y a rranged branches and/or metulae terminating in wh orls
01' phialides , presenting a n ov erall as pect 01' a group 01' sim ple penicil1i. The
terminal clus te rs usu all y cons ist 01' a prolongation 01' th e m ain axis, or a n

ind cfinitc numbcr 01' b runch es arising' irre gui arly at lowc r lcvclsto co mprise a
looscly a rrangcd two-st uge branc hcd a nd asym me tric pcni cillu s. In a nu mbcr
01' spee ies th e br an ch es arisc at onc Icvcl in a mctula-likc m anner , comprising
fairly eom paet but definitely d iva riea te pen icilli .
M embers 01' th e gro up a re eom manly isolat ed frorn soil, vege ta ble rem ains,
a nd or ganie mat eri als in th e lat er stages 01' decompositi on. They a re fre q ue ntly
eneo unte re d on fab ries a nd othe r military equ ipme nt undergoin g det eri or ati on
in th e field.
Altho ug h the Asymmetriea- diva rieata are re garde d as rep resen tin g, in
gene ral , a na tural grou p 01' Peni eillia , th ey sho uld be eons idered, however , as
oeeupy ing a somewhat intermedi at e posit ion in th e genus as a who le , sinee
different m embers assigne d here show affinities to ot he r well-defined gro u ps .
In eonst rueting th e key , we had to eree t th ree add ition al series to
aeeomo date seve ral 01' th e numerous spee ies 01' D iva rieata deseribe d sinee 1949
th at could not satisfaeto rily be c1assified in any 01' R ap er and T ho m' s ( 1949)
kn own series, nam ely:
T he P. godlewskii series, inc1uding th ose form s gro wing rather restrietedly,
with penieilli praduein g st ro ng ly diver gent eo nidial eha ins, a nd eomparatively
small eonid ia .
T he P. atro -sanguineum series, ereete d to inc1ude those for m s developing,
up an sorne substra ta, deep pu rpl e 0 1' blood-red eolou rs in th e colony reve rse,
an d diffu sin g int o the surrou nding agar .
The P. brasilianum series , inc1uding form s spreading broadl y on all subs tra ta,
appearing deepl y floeeose .

General key to th e Asymmetrica-divaricata sub-secti on

Penicilli typically once- or twice-branched below the level of the phialides,
asymmetrical, irregular; strongly divaricate, with individual elements
strongly divergent, often appearing simple but so arranged as to produce
the appearance of a monoverticillate or simple penicillus.
Page

1. Colonies producing sclerotia or masses of thick-walled cells.

A . Colony surface velvety or nearly so upon most substrata; stipes
arising directly from the substratum or from aerial hyphae.
P. raistrickiiseries .. .. 208
1. Conidiophore walls coarsely roughened; sclerotia well developed,
very hard.
a. Scerotia very hard, stony, white to pale pink in colour; vegetative
hyphae always colourless .
P. raistrickii Smith . ... 210
b . Sclerotia fairly firm, yellow to Iight brown in colour; vegetative
hyphae not always uncoloured.

1'. Vegetative hyphae somet imos encrusted with yellow

granules; conidia subglobose, up to 4.5 ILm in long axis.
P. pulvillorum Turfitt . . . . 212
2'. Vegetative hyphae always enerusted with yellow granules;
eonidia elliptical, up to 3.5 ILm in long axis.
P. pedemontanum Luppi-Mosea et Fontana .. . . 221
2. Conidiophore walls delieately roughened, true sclerotia laeking;
small rounded masses of thiek-walled eells present upon all substrata and partieularly upon malt extraet agar .
P. soppiiZaleski . . . . 214
3. Conidiophore walls smooth or nearly so.
a. White or pink sclerotia reported.

P. rolfsiiThom . . . . 217
b. Blaek sclerotia reported .

P. mirabile Beljakova et Milko .... 224

Small masses of thick-walled eells (as in P. soppi produced in
some strains.
P. miczynskii Zaleskl .. , .
(see P. janthinellum series)
B. Colony surfaee eommonly showing fascieulation, with conldiophores
aggregated into more or less well-defined synnemata or tufts.
The Asymmetrica-fasciculata (in part)
1. Sclerotia abundantly produced in recent isolates, often charaeteri zing the colony at temperature above 20C.
P. gladioli McCulloch et Thom .... 461
2. Sclerotia reported but seldom produced abundantly .
P. italicum Wehmer, var italicum Samson et al. . . . . 474
11. Colonies not producing sclerotia or masses of thick-walled cells.
A. Colonies not showing green , gray-green, or blue-green colours with
the ripening of eonidia .
1. Colony surface deeply velvety or lanate, becoming lilac, vinaceous,
avellaneous, or brown to ochraceous in colour.
P. lilacinum series . .. . 228
a. Colonies lanate, showing lilacinum to vinaceous shades, w ith
reverse similarly coloured.
P. lilacinum Thom ... . 229
(Paecilomyces lilacinus Samson)
b. Colonies not lanate, appearing velvety, comparatively th in, not
developing liIacinum or vinaeeous shades.
1'. Colonies spreading broadly upon most substrata, developing
pinkish-buff to avellaneous shades.
P. humulivan Beyma ... . 231
C.

2'. Colonias growing very restrlctedly upon Czapek's agar and

Czapek's yeast extract agar.
aa. Colonies developing avellaneous shades upon malt extract
agar.
P. argil/aceum Stolk, Evans et Nilsson
bb. Colonies developing brown to ochraceous shades upon
malt extract agar.
P. cy/indrosporum Smith
B. Colonies showing some shades of green, gray, gray-green, or bluegreen with the ripening of the conidia.
1. Penicilli with the divaricate character well marked; growing rather
restrictedly, at least on some substrata; with metulae scattered on
the conidiophore, or usually partly aggregated into true verticils.
a. Colonies developing typically pale blue-green or gray-green
shades with the ripening of the conidia; colony reverse often
brightly coloured.
1'. Conidial chains strongly divergent and/or becoming tangled
in age, not tending to form compact columns.
aa. Phialides abruptly tapering to a long, narrowed neck;
colonies commonly not funiculose.
P. janthinel/um series
1". Conidia conspicuously roughened, with roughness
arranged in spiral or transversal bands.
aaa. Conidia globose to subglobose .
P. da/eae Zaleski
bbb. Conidia conspicuously apiculate .
P. asturianum Ramrez et Martnez
2". Conidia usually roughened but with roughness not
arranged in spiral or transverse bands .
aaa. Vegetative hyphae and colony reverse strongly
coloured (orange-red, reddish purple, etc.) in recent
isolates .
P. janthinel/um Biourge
bbb. Vegetative hyphae uncoloured or slightly buff to
peach; colonies sporing sparingly or tardily; colony
reverse colourless or in yellow or orange shades.
1"', Conidiophores delicately roughened; penicilli
usually consisting of a terminal verticil of divergent
metulae; conidia ovoid to subglobose, delicately
roughened; colony reverse yellow to colourless.
P. simp/icissimum (Oudemans) Thom

. . .. 233

.. . . 236

. ... 238

.. . . 243
. . .. 254

. .. . 240

. .. . 246

2''' , Conid iophoros consplcuously w arty; panicllli irregular; conidia olllptlco l, smoot h walled or delicately
roughened; colony reversa developing orange
colours .
P. ochro-chloron Biou rge ... . 248
3'''. Conid iophores smooth walled or nearly so; penicilli
commonly irregular; conidia mostly subglobose,
conspicuously echinulate; colony reverse in cream to
light shades.
P. piscarium Westling . ... 250
4' '' . Conid iophores smooth walled or nearly so; penicilli
irregular; conid ia elliptical to subglobose, smooth
walled or nearly so.
aaaa. Conidia comparatively large, up to 6.5 {t m in
diameter; phialides large, up to 19 {t m in length.
P. aragonense Ramrez et Ma rtnez . .. . 256
bbbb. Conid ia compara tively small , up to 2.5 {t m in
long axis; phialides small , up to 10 {t m in length.
P. miczynskiiZaleski . . .. 262
bb. Phialides not abruptly taper ing to a long, narrowed neck;
penicilli cons isting of c1uste rs of strongly divergent phialides,
producing divergent columns of conidia; conidia compara tively small, usually not surpassing 3.0 {tm in diameter.
P. godlewskii series ... . 258
1". Conidiophores smooth walled or nearly so.
aaa. Conid ia smooth walled or nearly so.
1' ", Colonies growing very restrictedly upon Czapek's
agar, attaining up to 20 mm in diameter.
aaaa. Colon ies appearing definitely funiculose.
P. godlewskiiZaleski . . . . 259
bbbb. Colon ies appearing velvety
P. eben-bitarianum Baghdad i . . . . 266
2''' . Colon ies atta ining a diameter of 30 mm on CA .
aaaa. Conidia comparatively small, up to 3 {t m in diameter.
1"". Exudate and reverse in deep orange shades .
P. gorlenkoanum Baghdadi . . . . 269
2' ''' . Exudate colourless and reverse yellowish to
colourless.
P. botryosum Batista et Maia . .. . 261
bbbb . Conidia larger, up to 4 {t m in diameter.
P. citreo -virens Abe ex Ramrez ... . 279

3"'. Colonlos attalnlng a dlamotor 01 40 or more upon

CA.
aaaa. Conidia comparativelv small, up to 2.5 ,tm in
diameter.
P. sizovae Baghdadi
bbbb. Conidia comparatively large, up to 4 ,tm in diameter.
P. harmonense Baghdadi
bbb. Conidia conspicuously spinulose.
P. damascenum Baghdadi
2". Conidiophores conspicuously roughened.
aaa. Colony reverse developing typically strawberry red
shades upon MEA.
P. novae-caledoniae Smith
bbb. Colony reverse developing yellow shades upon MEA.
P. baradicum Baghdadi
2'. Conidial chains tending to produce compact columns, at least
in young cultures; conidia globose to subglobose.
P. canescens series
aa. Colony reverse developing deep red or brown shades;
not tending
towards
penicilli strongly divaricate,
ramigemous.
1". Colonies up to 0.5-1 .0 mm deep, with surface loose,
more or less floccose; conidial areas in dull blue-green
shades; conidia about 2.0-2.5 ,tm in diameter; colony
reverse orange, becoming brown in age.
P. canescens Sopp
2". Colonies deeply floccose, up to 2.0-3.0 mm deep;
conidial areas in brighter greenish glaucous shades; conidia
about 3.2-3.6 ,tm in diameter; colony reverse in deep red
shades near maroon.
P. nalgiovense Laxa
3". Colonies appearing velvety.
aaa. Conidiophores smooth walled; conidia conspicuousIy roughened .
P. turolense Ramrez et Martnez
bbb. Conidiophores
delicately
roughened;
conidia
strongly echinulate.
P. murcianum Ramrez et Martnez
bb. Colony reverse orange, in dull peach shades or colourless,
not developing dark colours; penicilli often appearing somewhat ramigenous.
P. jensenii Zaleski

.... 273

. . .. 271
. ... 268

.. .. 275
. .. . 264

. . .. 283

. . . . 285

... . 287

.. .. 293

.. .. 291

.... 289

b, Colonies developing typlcolly dull groy shades (scarcely show ing

any green) such as steel grey to dork oliva gray; eonidia globose;
eolony reverse usually in yellow to deep orange shades.
P. nigricans series .. .. 295
1', Conidiophore walls eoarsely roughened, at least upon malt
extraet agar.
aa. Phialides eonspieuously roughened .
P. radu/atum Smith .... 308
bb. Phialides smooth.
1". Conidia eonspieuously eehinulate .
P. me/inii Thom . . . . 303
2". Conidia smooth or nearly so.
P. raciborskiiZaleski .. .. 306
2'. Conidiophore walls smooth or nearly so on all substrata .
aa. Conidia smooth walled or nearly so.
P. yarmokense Baghdadi . ... 320
bb. Conid ia strongly eehinulate.
1". Conidia eomparatively very large, up to 7 Jlm in
diameter.
P. megasporum Orpurt et Fennell .... 31 6
2". Conidia large, up to 5 Jlm in diameter.
P. a/bidum Sopp .. .. 299
3" . Conidia smaller, up to 3.5 Jlm in diameter to 4.0 Jlm.
P. grana tense Ramrez, Berenguer et Martnez . . . . 310
ee. Conidia slightly roughened, provided with two roughly
parallel thin ridges.
P. inf/atum Stolk et Malla . . . . 312
dd. Conidia eonspieuously and finely spinulose.
1". Colonies growing very restr ietedly upon Czapek's agar,
attaining a diameter of 15 mm; margin abrupto
P. ovetense Ramrez et Martnez . . . . 318
2". Colonies growing less restrietedly, attaining a diameter
of 25-30 mm.
aaa. Colonies heavily sporing in dark brown shades.
P. nigricans (Bainier) Thom .. .. 297
bbb. Colonies slightly sporing, white or nearly so, or in
pale blue-green shades upon malt extraet agar.
P. kapuscinskii Zaleski . . .. 301
e. Colonies developing blue-green or gray-green shades at maturity
of eonidia; eolony reverse typieally in deep purple or blood red
eolours upon some substrata; with similarly eoloured pigment
diffusing into the surrounding agar.
P. atro-sanguineum series . . .. 322

C'OIllIlHJIl ly IIp to 2:">()- 1-00 u n 01' ruorc in lcn gth I with walls ra llging' rom
u ns picou sly cc h in uln tc lo smooth 01' ncurl y so .
Pcn icilli typ ica lly d ivari cat c usyuunctri cul, usua lly co nsisting' 01' a term inal
ve r!icil 01' 2 - 5 m ctu lac, gc nc ra lly som cwhut di vcrgcnt .
C o n id ia glob osc lo cllip tica l , with wa lls gcnerally smooth 0 1' ncarly so.

K cy lo spccies
Paga
A . Colony surfaee velvety or nearly so upon most substrata ; stipes arising
direetly from the substratum or from aerial hyphae.
P. raistrickii series
1. Conidiophore walls eoarsely roughened ; sclerotia well developed,
very hard .
a. Sclerotia very hard , stony, white to pale pink in eolour; vegetat ive
hyphae always eolourless.
P. raistrickiiSmith
b. Sclerotia fairly firm , yellow to ligh t brown in eolour; vegetative
hyphae not always eolourless.
1' . Vegetative hyphae sometimes enerusted with yellow granules;
con id ia subglobose, up to 4.5 JLm in diameter.
P. pulvillorum Turfitt
2'. Vegetative hyphae always enerusted w ith orange granules;
eonid ia elliptiea l, up to 3.5 JLm in the long axis.
P. pedemon tanum Luppi-Mosea et Fontana
2. Conidiophore walls delieately roughened , true sclerotia laeking; small
rounded masses of th iek-walled eells present upon all substrata and
partieularly upon malt extraet agar.
P. soppiiZaleski
3. Condiophore w alls smooth or nearly so.
a. White to pink sclerotia reported .
P. rolfsiiThom
b . Blaek sclerotia reported.
P. mirabile Beljakova et Milko
e. Small masses of thiek-walled eells (as in P. soppii) produeed in
some strains .
P. micz ynskiiZaleski
(see P. janthinellum series)

. . . . 208

. . .. 210

. . . . 212

. .. . 221

. . .. 214

.. . . 217
. . . . 224

.. . . 252

T he seri es is characterized particu larly by th e production of true scleroti a 0 1'

sclerotia-like m a sses of thi ck-walled cclls, th e presen ce of two-stage branched
penicilli with eleme nts ty pica lly d ivergent , a nd a lack of a ny tendency tow ard s

th c lorrnation 01' synne mata. In add ition lo th c our specics cons ide red hcre by
Raper and Thom (191,9), namcly: Pen icillium raistrickii Sm ith , P. puloillorum
Turfitt, P. soppii Zale ski, and P. rolfsii Thorn , th c followin g spccies havc been
included in the seri es in the prescnt study : P. pedemontanum Luppi-Mosca et
Fontana, and P. mirabile Belj akova et Milko . Howevcr , a close relationship
between th ese forms is not presumed here , a nd th e six species are conside red
together principally as a m atter 01' conve n ience.
The sclerotia 01' sclerotia-like cellula r mas ses , differ su bsta ntially, dep ending
upon th e species. In Penicillium raistrickii th ese are globose to su bg lobose , ve ry
hard a nd gritty , be aring a striking resemblance to th e flesh- to pink-coloured
scle ro tia produced by P. thomii Maire , in th e Monov erticillat a Section (colou r
pIate 1). The cellula r masses in P. puloillorum are less regul ar in shape, 01't en
larger, firrn , but not sto ny, com monly su rrou nde d by a loos e network 01'
yellow- to brownish- en crusted veget ati ve hyph ae lending a brownish colouration to the colony su rface . In P. pedemontanum, th e scle ro tia are hard , alw ays
surrou nde d by a loose network 01' fin e orange-enc rus ted hyphae whi ch lends a
deep orange colou ra tion to th e colony . P. roljsii Thom a nd P. mirabile Belj ak ova
et Milko are in cluded in th e series primarily because Thom (1910 ), in his
origin al di agnosis 01' P. roljsii reported th e production 01' white to pink sclero tia,
a nd Belj akova and M ilko (1972 ), in th eir or iginal diagnosis 01' P. mirabile,
rep orted the presen ce 01' hard black sclerotia . On th e othe r hand , the aggrega tes 01' thick-w alled sterile cells ob serve d in P. soppii are qu ite unusu al amo ng
th e Pen icillia , suggesting rudimentary sclerotia, 0 1' possibl y asc omata . T hey
are regul arly produced adjacent to th e su b stratum a nd are generally obsc ured
by the m ycelial ove rgro wth a nd th e heavily spo ri ng a reas.

73 . Penicillium raistrickii Smith : fig . 73, colour plat e 73, and SEM plat e

13 , (t op : left).
Penicil/ium raistrickii Smith - Trans . Brit. Mycol. Soco 18: 90. 1933. See also Raper
and Thorn, 'A Manual of the Penicillia', pp . 275-277. 1949.
Smith's diagnosis is as follows:
'Colonies growing rapidly on most media at temperatures up to 30C, rate of
growth much less above 30C; bluish green, then grayish green , and finally some what brown , velvety; reverse persistently uncoloured, showing masses of sclerotia;
conidiophores arising form the substraturn, up to 250 .tm long, 3.5-4.0 .tm in
diameter, rough; penicilli typically divaricate, comprising 3-5 monoverticillate
heads united into a fairly compact whole; metulae clavare, sometimes very slightly
roughened, 10-13 .tm by 4.0-4.8 .tm; phialides closelv packed on each individual
metula, 7 .5-9.0 .tm by 2.6-3.0 .tm; conidia globose or nearly so, smooth, 2.2-2.7
.tm in diameter, conidial chains packed into diverging salid columns; sclerotia

....

: ~'

'. '

.:.!~
. ...

~~:.'

..
',:

'

r.~

' 0'

v-,

:'.:.
:!"

:oJf?~ \~,)1
:;.

::
'o";

.~.,

:. ...;

'~\

t~~~
~. :

Fig. 73. Camera lucida

draw ings of Penicillium
raistrickii Smith, CBS
261 .33. A , Detailed drawings
of the penicilli. B, Mature
eonidia. C, Habit sketehes
on malt ext raet agar. D,
Habit sketehes on Czapek's
agar. E, Sclerotia.

produced abundantly, hard, feeling like grit under a cover glass, consisting of
masses of irregular-shaped thick-walled cells , white to dirty wh ite, globose or ovoid ,
up to 180 JLm in long axis.'

Our notes follow:

Colonies u pon CA (colour pla te 73, A and a) attain ing a diameter of 45 mm
in 14 days at room temperature (20-25 C), velvety, radiately sulcated wi th
shallow furrows, azo nate or sligh tly za nate; even margin surrounded by a
fringe of submerged hy phae ex tending about 1 mm beyond the ae rial growth;
colony surface ove rlaid by a mycelial overgrowth of white f1occose aerial
h ypha e , lending a lanate appearance to the colony; heavily spo ri ng,
particularly in m arginal areas in blue-green shades (Sguy, PI XXXI, 461);
exudate limi ted in amount, colourless; odou r lackin g; no coloured pigm ent
diffusing into the surrounding agar; colony reverse (colo ur piate 73, a)
colourless to cin namon shades; penicilli as des cr ibed by Smith except conidia
whic h are elliptical (fig 73 , A), 3.0 -4 .0 /lm by 2.0 -3.0 /lm in diameter;

sclcrot iu are ab un dantly prod uced rhroughout thc colony surfacc , often
characterizing ils ap peara nce .
Colonies up on CYA (colour piale 73 , B a nd b) auain ing a diarnetcr 01' 60
mm in 14 days at room temper atu rc (20 - 25C), duplicati ng in tcxtu rc ami
appearance those grown up on C A, but radiatcly sulcat ed by numerou s and
deep furro ws, with cen tre ra ised; margin su rro unde d by a wide fringe 01'
submerged hyphae , ex te nding about 4 m m beyond th e aerial gr owt h ; spor ing
ab undantly in marginal areas in the same shades as described aboye; ex udare
abundantly produce d, crystal c1ear, colou rless; odour lacking ; no coloured
pigme nt diffus ing int o the surroundi ng agar; colony reverse (colour pla te 73,
b) in orange shades; penicilli essential1y as describe d aboye .
Colonies upon MEA (colour plate 73, C and c) attaining a diameter 01' 60
mm in 14 days at roo m temperature (20 -25C), consisting ofa loose-textu red
basal felt overlaid by a Ioosely woven network 01' floccose aeriaI hyphae
in fluencing greatly the colony appearance, plane; heavily sporing in margi na l
areas in blue-green shades (Sguy, PI XXXI, 46 1); exudate and odo ur
lacking ; sclerotia Iess ab undantly produced than upon the precedi ng substra ta;
colony surface appearing light buff from the production 01' sclerotia and the
abundant ovcrgrowth 01' sterile floccose hyphae ; no coloured pigment diffusing
into the surrounding agar; colony reverse (colour plate 73, c) in reddish brown
to deep orange red shades; penicil1i essential1y as described aboye . (Fig. 73, A
and C).
The species description is centered u pon the type strain CBS 26 1.33 = CM l
40 .22 1 = ATCC 10,490 = NRRL 2039 = n-o 6, 104 = QM 1,937 = VKM F-337, isola ted
by George Sm ith fro m mo uldy cotton yarn in 1927, in England .

74 . Penicillium puloillorum Turfitt: fig . 74, colou r plate 74, and SE M

plate 13, (top: right) .

Penicil/ium pulvil/orum Turfitt - Trans. Brit. Mycol. Soc o23: 186-187. 1939. See also
Raper and Thorn, 'A Manual of the Penicillia ' , pp . 277-279 . 1949.
Turfitt's o riginal diagnosis follows, since this adequately presented the species in
question when first isolated :
'Colonies on Czapek's agar at 24C, matted floccose, often radiately wrinkled ,
spreading, becoming 35-40 mm in diameter, 0.5-1 mm deep in 8 days ; marginal
zone raised, white , passing later to brownish shades, 3-5 mm w ide ; conidial areas
at f irst pale green , becoming deeper green , c1early zonate towards t he growing
edge with zanes about 2 mm apart, gradually turning brownish from centre out
wards w ith the development of sclerotia ; reverse colourless at fi rst, then somewhat
zanate in pale yellow shades, becorn inq deeper yellow and brownish in age; odour
none; conidiophores commonly arising as short branches from trailing hyphae,

~ llf lf@'~

' :s-:~
r~
. , A{.
f

e .

100 prn

I~,

~.

Fig. 74. Camera lucida

drawings of Penicillium
pulvilloru m Turfitt, CSS
280.39. A, Detailed drawings
of th e penicill i. S, Mature
conidia . C, Habit sketches.
D, Sclerotium.

1.5-3 ILm in diameter, w ith walls markedly roughened ; penicilli oeeasionally as

sinqle verticils of phialides, usually divarieate with terminal groups of 2-3 metulae ,
and with seeundary penicilli, mos tly monovertieillate, arising from lower nodes of
main axis ; metulae mostly 3-3.5 ILm in diameter, varying greatly in length , 12-25
11m; phialides 8-10 by 2.5 ILm, sharp pointed , with eonidial ehains roughly parallel or
more or less divergent, beeoming tangled in age; eonidia 2.5-3 ILm in diameter,
smoot h, globose; scierotia very abundant, form ing early amongst superficial
growth of trailing and anastomosing hyphae, yellow-brown, irregular in shape , very
variable in size, averag ing 700 ILm in diameter, eonsisting of eompaet hyphal
masses, remaining soft; development of asci not observed .'

Our notes follow :

Colon ies upon C A (colou r pl ate 74 , A and a) attainin g a d iamct er of 50 mm
in 14 d ays at room temperature (20 - 25 C ), co nsisting of a clo se -tex tu red bas al
lclt, dc ep ly radiat ely fur rowcd , in whi ch are em be dd ed irregular, yellow to
ligh t brown sclerotia abou t 100 IJ-m in lon g axis, and with th e colony surface
over la id by a thin , more or less floccose overgrowth of interIacing aerial hyphae

giving' a lan utc appl:a ranee lo 11H: colony, wh ich rc ma ius pcrsistcnrl y whit c I'Osy
in colou r; oven ma rgin su rrou nde d by a frin gc of sub mc rged hyph ae cx ic nd ing

about 1-2 mm bcyond thc aerial growth; slighlly spo ring in palc blu c-gr een
shades (S guy , PI. XXX, 444); odour lacking ; ex udare in large , crys tal-clear ,
un coloured drops ; pal c am ber-colou red pigment diffu sin g in to the su rro unding
ag a r; colony rever se (colour plate 74 , a) in pal e ora nge shade s; peni cilli as
described by Turfitt (fig. 74, A) , with conid iophores cons picuous ly and finc1 y
roug h wall ed , arising mostly from tr ailing hyphae ; co nidia elliptical to
subglobo se (fig . 74 , B, and SEM plate 13, top : right) , with walls smo oth 01'
nearl y so, 3 .0 - 4 .8 /lm by 2.5-4.0 /lm in diamet er. In ou r expe rience , the
scleroti a 01' P. pu loillorum , whil e not hard and gritty , are nevertheless quite firm
and withstand cons ide rable pressure without collapsing .
Colonies up on CYA (colour plat e 74 , B and b) attaining a diamet er 01' 55
mm in 14 days at room temperature (20 - 25 C), duplicating in texture , cultu ra l cha racte ristics, a nd details 01' morpholo gy thos e grown upon CA ; even
margin su rrou nde d by a narrow frin ge 01' submerged hyphae exte nding ab out 1
mm beyond th e aerial gro wth; heavil y spo ring in m arginal areas in gray-gree n
shades (S guy, PI. XXVII, 403); odo ur lacking ; exudate abu ndantly produced
in large , amber- coloured drops ; pale amber-coloured pigm ent diffu sin g into
th e su rrou nding a gar; colony reverse (co lour plat e 74, b) in pal e or ange
shades ; peni cilli and scle ro tia as described aboye .
C olonies up on MEA (colour plate 74 , C a nd c) attaining a dia me ter 01' 45
mm in 14 days at room temperature (20 -2 5C), cons isting 01' a den se layer 01'
larger , yellow to light brown sclerotia , up to 250 /lm in long ax is (fig. 74, D ),
ove rlaid by a loose network 01' interw oven hyphae ; spo ring heavily th rou ghout
in mouse gray shades (Sgu y , PI. XXIX , 430) ; with a n eve n margin
su rr ou nde d by a fringe 01' subme rge d hyphae exte nding abo u t 2 mm beyond
the ae rial growth; odour and exuda te lacking ; on coloured pigm ent diffusing
into th e surrounding agar; colony re ve rse (colou r plat e 74, c) in pale cream
shad es; penicilli esse ntially as described ab oye but with conidiophores arising
mo stly fro m th e substratu m, com mo nly 500 /lm 0 1' m or e in len gth by 3.0- 3.5
/lm in di amet er; phialides in clu sters 01' 3- 8 elements in th e verticil , mo stly
7-11 /lm by 2.5- 3.5 utti, terminating in a conspicu ou sly taper ed , narrowed
ne ck (fig . 74, A); conid ia appearing deli cately roughen ed ; conidial cha ins
tangled 01' formin g divergent colum ns (fig . 74, C) ; sclerotia up to 250 /lm in
diameter.
The sp ecies description is cente red upon th e typ e strains C BS 280. 39 = NR R L
2,026 = IFa 7,763 = IJFM 5118, isolat ed by G .E . Turfitt from ac idic soil
collected in London , U .K.

75. Penicillium soppii Zaleski: fig. 75, colou r plat e 75, and SEM pIate 13
(centre: left) .

WJ..W.u.J

10 jrrn

o~
00~

O 000

O
00

C)

00

80

,,/

~ l?# 1~ 1~/ lR

o
() OoCD ..,.. .

1~ 1
.1lc

Fig. 75. Camera lucida

draw ings of Penicillium
soppii Zaleski, CBS 225.28.
A, Detailed drawings of the
penicilli. B, Mature conidia.
e, Habit sketches.

Penicillium soppii Zaleski - Bull. Acad. Polonaise Sci. Math. Nat. Sr. B, pp.
476-477.1927. See also Ch. Thom, "The Penicillia ', pp . 344-345.1930; also Raper
and Thom, 'A Manual of The Penicillia', pp. 279-282. 1949.

Synonymy
Penicillium matris-meae Zaleski - Bull. Acad. Polonaise Sci. Math. Nat. Sr. B,
pp. 477-479. 1927.

Colonies upon C A (colour pIate 75, A and a) att aini ng a di ameter of 55 mm

in 14 days a t room temperature (20 - 25C), cons isting of a close- tex tured basal
felt ove rlaid by a loose m yceli al ove rg rowth up to 0.5 mm deep ap pearing
almos t lanate , sulcated by very shallow radial furrow s; with a thin , cven
marg in surrou nde d by a wide fri n ge of sub merged h yphae ex tend ing 4- 5 mm
beyond th e ae rial grow th ; spari ng ly spo ring in pale gray-gree n shades (S guy,
PI. X X IX, 430), not infl uen cing the colony ap peara nce ; odo u r faint,
' mouldy ' ; exu date abundantly produced , as large top az yellow-co lou red
drop s, influencin g greatly th e colony as pec t; a m be r-coloure d pigment diffusing
into the surrou nd ing agar; colony reverse (colour pla te 75, a) develo p ing

rcddish sic n na brown shadcs: with rhick-wullcd, stcrilo ce lls in small m a sses u!'

to 50 -60 J.III in diam etcr, suggcs tiug small sclcro tia , occuring adjaccn t to th e
subs tra turn th rou ghout th e colony but rnos t ab undan tly alo ng thc inter colony
margins, resembling fine white sa nd in appearancc but not firm o r gritty ;
peni cilli (fig. 75, A) borne up on lon g con id iopho res with walls smoo th or finel y
roug he ned, up to 1000 J.m by 2.5-3 .0 uiri , arising mostly from the substratum
along the intercolony margin s, a nd as bran ches of va riable len gth from ae rial
hyph ae in centra l are as; penicilli forming large struc tures usu ally consisting of
a terminal verticil of 2-5 metulae , 12- 25 J.m by 2.5 -3. 0 uu , with ap ices
enla rged in a ves ic1e up to 4.0 J.m in di am ct er; phialidcs in c1us ters of 2-8
closely crowded eleme nts in th e verticil, mostly 9- 15 J.m by 2.5 -3 .0 tui,
terminating in a short-po in tcd ncck ; conidia elliptical to subglob osc (fig. 75, B,
and SE M plat e 13, ce nt re: left), mostl y 3 .5 - 6.5 tut: by 3. 0- 4.0 J.m in
diameter , with su rface smooth or delicatel y ro ug he ned; conidial chains loose to
tan gled .
Colon ies up on C YA (colour plate 75, B a nd b) broadly spreading , attain ing
a di amet er of 60 mm in 14 days at room temperature (20-25C), cons isting of
a c1ose-textured basal felt , ove rla id by a loosely interwoven netw ork of floccose
ae rial hyph ae lending a lanate appeara nce to the colony su rface, which is
deepl y radi at ely furrowed ; heavily sporing particul arl y in m arginal areas in
blue-green shades (Sgu y , PI. X X X, 438) ; thin , eve n marg in surrounde d by a
wid c frin ge of submerged hyphae extend ing 4 -5 mm beyond th e ae rial growth;
odo u r lacking; exudate ab undantly produced in pale yellow drop s; very pale
am ber pigment diffu sin g into the su rrou nding agar; colony revers e (colour
plat e 75, b ) in cinnamon shades; penicilli esse ntia lly as described aboye , with
the conidiopho res arising m ostl y from tr ailing ae rial hyphae .
Co lon ies up on M EA (colour plat e 75, C and c) at ta in ing a diamet er of 60
mm in 14 days at room temperatu re (20-25C ), cons isti ng of a deepl y velvety
to subla nate felt , slightly zanat e in central arcas, pla ne , ab undantly spo ring
throu ghou t in pale blu e- green colours (Sguy, PI. XXVIII , 4 15); thin m argin
surrou nde d by a frin ge of submerged hyph ae exte nd ing 4-5 mm beyond th e
ae rial growth; exuda te a nd odo ur lacking ; no coloured pigm ent diffu sin g into
the su rro u nd ing agar; colony reverse (colou r plate 75, e) in pale yellow sha des;
peni cilli esse ntially as described aboye, but with the stipes of the conidiophores
arising mostly fro m th e su bs tratum an d from ae rial hyphae .
The spec ies description is cen tered upon th e type strain CBS 226. 28 = C M I
40,2 17 = ATCC 10,496 = NR RL 2, 023 = I Fa 7,766 = QM 1,964 = IJ FM 5126,
isolat ed by K . Zaleski from soil in Poland , 1927 , and up on strain NR RL 9 12 =
C BS 225.28 isolated fro m th e sa me sou rce by Zaleski in the sa me year , and
m aintained in artificial cultu re since 1928 in th e Northern R egional R esearch
Laborat ory , Bureau of Agricultural and Industrial Chemistry, U.S . D epart
m ent of Agriculture , Peoria , Illinois. However , although it duplicates th e

preccdin g cultu re in cultural cha racteristics a nd in details 01' morphology, thus

far, it failed to produce the typ ical masses 01' thick-walled cclls.

76. Penicillium rolfsii Thom : fig. 76, colou r plate 76, a nd SEM plate 13,
(centre: right).

~50 ~
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O O

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Fig. 76. Camera lucida

drawings of Penicillium ro/fsii
Thom, ces 368.48. A,
Detailed drawings of the
penicilli. B, Mature conidia.
e, Habit sketches.

Penici//ium rolfsii Thom - Bul!. Bur. Anim. Ind. U.S. Dep. Agric. 118: 80-81. 1910.
See also Ch. Thom, 'The Penicillia', pp. 489-490.1930; also Raper and Thom, 'A
Manual of the Penicillia', pp . 282-283. 1949.
Author's original diagnosis follows:
'Colonies upon milk -sugar-gelatin and potato or bean agars gray-green;
floccose, but with aerial part mostly long conidiophores and few vegetative hyphae,
slightly yellowish to pronounced salmon colour below; broadly spreading;
developing elliptical to globose scierotia 150-200 p'm in diameter at the surface of
the substratum in 2-3 weeks, conidiophores 200-500 p'm by 3-4 p'm; penicilli in
verticils of 3-5 branches Irnetulae) 10-17 p'm by 2-3 p'm, rarely secondary verticils

each bearing a dense vert icil of sterigmata (phialid es), 8-10 by 2 J-lm , produclng
long , parallel, or slightly divergent chains of conidia; conidia elliptical or fusiform,
3.5-6 J-lm and producing one to several germ tubas . Colonies slowly Iiquefy
milk-sugar gelatin and produce purple to neutral colours in Iitmus media.
Sent by Prof. P.H. Rolfs from Miami, Florida, on portion of pineapple , March,
1905.
Colour gray-green, with scattered white to pink sclerotia, reverse sulphur
yellowish to pronounced salmon; colour in media reddish or yellowish in special
cases, others none. Odour none.'

O u r notes follow :
Colon ies upon C A (co lour plat e 76, A and a) a tt a ining a di ameter of 65 mm
in 14 days at room temperature (20- 25 C ), cons isting of a close- textured basal
felt , overla id by a den se network of floccose wh ite ae rial hyph ae , lendin g the
colony a lanate appeara nce; th in, eve n m argin su rrou nde d by a wide fringe 01'
submerged hyphae exte ndi ng abo u t 3-4 mm beyond th e aeri al gr owth ; heavily
sporing throu ghout in gray-green to blu e-gr een shades (Sguy, PI. XXX , 449,
442), with con id ial areas partially obsc ured by the den se overgrowth of floccose
ae rial hyphae , espec ially in sub-ce ntra l a nd m arginal a reas, show ing whitc
rosy sha des in the colony su rface ; odo u r a nd exudate lacking; coloure d
pigm ent d iffus ing into th c su rrou nd ing aga r not produced ; colony revers e
developing salmon pink colours (colou r plat e 76, a); peni cilli are irregular in
pattern and essentially as described by Thom ( 19 10), but showing also sim ple
pen icilli (fig . 76, A) , with th e stipes 01' thc conidiopho re s u su ally smooth
walled , but som etimes show ing some what rou gh walls; phialides terminating
in a sho rt, tap er ed neck ; conidia strongly ellip itical, with walls smooth or
nearl y so wh en observed under the light mi cro scop e (fig . 76, B), but delicately
gra nular wh en ob serv ed under th e SEM (pl a te 13, cen tre : rig ht), mo stly
3.0 - 4.0 p.m by 2.0 -2 .8 p.m in di am et er; conidial chains loosely parallel (fig.
76, C ) . Scle ro tia wer e not obse rve d .
Colonies up on C YA (colour plate 76, B and b) broadl y spreading, attain ing
a di amet er of 65 mm in 14 days at room temperaturc (20 -2 5C) , duplicating
in cultu ral charac teristics a nd details of m orpho logy those gro wn up on C A, but
with colony su rface deepl y furrowed in a radial pa tte rn; spor ing sim ilarly as
abo ye in th e sa me shades; exudate and odour lacking; no colou red pigm ent
diffu sin g into the surrou nding medium ; colony reverse (colour plate 76, b) in
deep er salmon pink shades; peni cilli essentially as described aboye but with
conid ial chains tan gled. Scle ro tia wer e not observed .
C oloni es upon M EA (colour plate 76, C and e), attaining a diameter of 65
mm in 14 days at room temperature (20 - 25OC ) , consisting of a close-t extured
basal felt overl aid by a den se netw ork of whi te rosy floccose aerial hyphae ,
obscu ring th e otherwis e heavily sporing areas, particul arl y in central and
subce ntral zon es, influ en cing greatly th e colony appearance, wh ich devel op s

whit c rosy shadcs in ce ntral and subccnt ral arcas ; hcavily sporing arca s in
margin dcvcloping decp blu c-grccn sha des (Sguy, PI. XXX, 441); odour a nd
cxudate lacking; no coloured pigment dilTusing into th e surrounding aga r ;
colony revcrse (colour pIate 76 , e) developing yellowish to pale salm on pink
shades; peni cilli essentially as described aboye . Sclerotia were not observe d.
Colonies grawn upon th e three standard su bs trata us ed in th e curre nt study
conforrn fairly closely with Thoms (1910) original des cription but differ in its
Iailure to produce sclerotia .
W e a gr ee with R aper a nd Thom (1949) in ass ign ing thi s species to th e
Penicillium raistrickii series in th e Asymmetrica-divaritaca su b-sec tion , on
account of th e reported presen ce ol' white to pink sclerotia 150- 200 p.m in
di ameter in th e type strain when originally described.
The species description is ce nte red upon th e typ e strain CB S 368 .48 = C M I
40, 02 9 = ATC C 10,491 = NR R L 1,078 = IFa 7,785 = Q M 1,961 = IJFM 5119 ,
isolat ed by Ch . Thom from a portion of A nanas sativus sent by Prof. P .H . R olfs
frorn Miami , Florida , M arch, 1905 .

77 . Penicillium rolfsii Thom , var. sclerotiale No vobranova: fig. 77 .colour

plat e 77 , and SEM plat e 13 , (bottom: left).

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Fig. 77. Camera lucida

drawings of Penicillium rolfsii
Thom varo sclerotiale
Novobranova, CBS 752.74.
A, Detailed drawings of the
penicilli . B, Mature conidia .
C, Habit sketches. D,
Sclerotium.

Penicillium ro/fsii Thom, varo sclerotiole Novobronova - Nov. Svst, Nizhnlkh Rast.
11: 230-232. 1974.
Novobranova's original diagnosis follows:
'Colonies upon Czapek's agar attaining a diameter of 25 mm-30 mm in 10 davs,
with undulating margin; close textured , with the colony centre somewhat heaped;
furrowed; grayish violet-coloured sclerotia abundantly produced throughout the
colony surface in 34 days, elliptical to subglobose or irregularly shaped, mostly
200-350 .tm or more, up to 400 .tm in diameter. Conidial structures appearing slowly
and sparingly, particularly in submarginal zones, developing gray-green shades;
exudate abundantly produced, principally in submarginal areas, as crystal c1ear,
hyaline droplets. Colony reverse wrinkled in the centre, whitish, with centre in
marble pink colours; strongly splitting the agar underneath. Conidiophores with
walls delicately roughened, 200-500 .tm by 3.0-4.0 .tm. Penicilli borne on c1usters
of metulae, numbering 3-5 elements in the verticil, mostly 10.0-17.0 .tm by 2.0-3.0
.tm; phialides measuring 8.0-10.0 .tm by 2.0 .tm; conid ial chains long, either parallel
or divergent . Conidia smooth walled, elliptical, mostly 3.5-4.0 .tm by 2.0-3.0 .tm in
diameter. The species has been assigned to the Penici/lium raistrickii series, in the
Asymmetrica-divaricata sub-section. It is closelv related to P. ro/tsii Thom, from
which it differs particularly by its morphological characteristics and by the abundant
production of sclerotia upon Czapek's agar.' (Translated from the Russian by Prof.
L. Vilas).

Our not es follow :

C olonies up on C A (colou r plat e 77 , A and a) grow ing rathe r slowly ,
atta ining a diameter of 35 mm in 14 days at room temperature (20 -25 C ),
consisting of a close-tex tu re d basal felt , deepl y velve ty to su bla nate , overl aid
by a thin, closely woven network of floccose aeri al hyphae, influeneing the
colony appeara nce; strong ly raised in th e cen tre in a crate riform pattern and
dee ply radiately wrinkled in m arginal area s; thin , eve n m argin surrounded by
a narrow frin ge of subme rge d hyphae exte nding abo u t 1 m m beyond th e ae rial
gro wth ; hea vily spo ri ng throu ghout in light blue-gr een shades (Sguy , PI.
XXX, 438), some what obsc ured in th e cen tre by th e thin ove rgro wth of ae rial
hyphae ; exudate and odour lacking; no coloured pigm ent diffusin g int o th e
surro un d ing agar; colony reverse (colou r plat e 77, a) strong ly pulling aw ay the
agar underneath , developing ora nge colou rs. Scleroti a hav e not been obs erved
as originally described , but very spa ringly produced up on CA (fig. 77, D) ;
penicilli (fig. 77, A) essentially as described originally by N ovobranova (1974) ,
but with th e conidiopho res appearing smooth wall ed .
Colon ies upon CY A (colour plat e 77 , B and b) , atta ining a diameter of 55
mm in 14 days a t room temperature, consis ting of a close-t extured basal felt,
deepl y velvety, with m argin plan e, su rro unde d by a frin ge of sub me rge d

hyph ae extc nd ing abou t 2 -:~ 111111 beyond the acrial growth ; ce ntral and
subcc ntral arcas deep ly radiately lurrowcd: heavily sporing throughout in
blu e-grecn shades (Sgu y, PI. XXXI, 463) ; exu date and odour lacking; no
colourcd pigment diffusing into the surrounding agar; colo ny re vers e (colour
pial e 77, b) strongly pu lling the agar away , developing brown colours : penicilli
esscntially as described aboye; sclerotia not observed upon this substratum .
C olonies u pon MEA (colour plat e 77 , C and c) growing rather restrictedly,
consisting 01' a looser textured basal felt, vel vet y in appearance, attaining a
diameter 01' 45 mm in 14 days at room temperature (20-25 C) , plane,
sornewha t zonat e , wit h margin irregu larly disse cted surrounded by a fringe 01'
su bmerged hyphae extending about 1- 2 mm beyond the aerial growth; heavily
sporing throughout in a rte m isia gr een shad es (Sguy , PI. XXIX, 430) ;
exudate and od our lackin g; no coloured pigment diffusing into th e
surrounding agar; colony reverse (colour plate 77 , e) in yellow-orange shades ;
pcnicilli esscntially as described aboye; sclerotia have not been observed upon
this substratum .
The spec ies des cription is centered upon the type strain C BS 752 .74 = VKM
F- 1815 = ATCC 24 ,724 = IJFM 7038, isolated from the surface 01' fruit 01' Pirus
malus L. variety 'Renet Burchardt' in the Alm a-Ata region 0 1' the S .S .R. 01'
Kazachstan, USSR , by T .I. Novobranova, April, 1968. It is known on ly from
the typ e spe cies .

78 . Penicillium pedemontanum L u ppi-Mosca et Fontana: fig . 78, colo u r

p late 78 , and SEM plate 13 (bottom : r ight).
Penicillium pedemontanum Lupp i-Mosca et Fontana - Allionia. 9: 35-41. 1963. See
also M.M. Kulik, 'A Compilation of Descriptions of New Penicillium Speci es' .
Ag ric. Handb ook No . 351. Agric . Res. Ser. U.S. Dep. Agri c. pp. 63-65. 1968.
Luppi-Mosca and Fontana's origina l diagnosis follows, since this adequately
presents the species in question:
'Cult ures upon Czapek's agar growing fai rly rapidly, attaining a diameter of 25
mm in 12 days, and about 40 mm in 20 days.
Consisting initially of a basal mycel ium , at first colou red in straw yellow shades,
becomi ng then immediately sulp hur yellow in colou r; afte r 4-5 days, the fi rst
conidial structures develop ing gray-blue shades appea r in the colony centre, whilst
the entire colony surface is overlaid by an abundant production of cream to pale
lemon-coloured sclero tia.
At complete development , t he conidial areas are mostly reduced to the colony
cent re, appearing powdery, in gray-blue shades, while t he abundantly prod uced
sclerotia all over the colony surface develop gray-avellaneaous colours.
A comparative premature exudate is produced as hyaline droplets, increasing
w ith the aging of the colony.

a d iamctcr 01' 55 m m in 14 days at room tempcnuurc (20 - 25 C) , co nsisting 01'

a close -tcxturcd basal Ielt of ycllow-encrusted hyph ac , ap peari ng vclvety 1
cons picuo us ly zona tc, with sulph ur ycllow colours a Iternating with th e graygreen (Sguy, PI. XXIX, 428) shades of the spo rulati ng areas; exudare
ab undantly produced as m in ute drop s; odou r lacking; orange-coloured
pigment diffusing in to the surroun ding agar; eve n margi n surrounded by a
narrow fringe 01' submerged hyphae ex tending abou t 1 mm beyond the ae rial
growth; colony reverse (colour plat e 78, b) in dee p orange shades; colony
surface deeply radiatcly furrowed; pen icilli esse ntially as descri bed aboye;
sclerotia ab undantly prod uced in ora nge colo urs, partially submerged in the
substratum .
Colon ies upon MEA (colour plate 78, C and c) attaining a dia me ter ranging
from 35 to 50 mm in 14 days at room temperatu re, consisti ng 01' a loosetextured felt , appearing velvety, conspicuously zonate , with thin , undul ating
margin su rrounded by a thin fringe of submerged hyph ae extending less than 1
m m beyond the aerial growth; heavily spo ring th ro ug hout in pale blue-green
shades (Sguy , PI. XXX, 444, 438); orange sclerotia submerged in the
substratu m; exudate a nd odour lacking; deep orange -red-colou red pigmen t
diffusing in to the surrounding agar; colony reverse (colour plate 78, e)
deve loping similar colour; penicilli essentially as described aboye but with
conidial chains tangled .
The specics description is centered upon the type strain CBS 265.65 = CM I
143,926 = ATCC 18 ,334 = IJFM 7045, isolated by A .M . L uppi- Mosca and A.
Fontana from Fagus silvatica L. mycorrhizae, Torino, Italy , in 1963; strain CBS
343 .52, isolated from soil in Norway, and strain IJFM 3782, isola ted by D I' 1.
Moriyn from soil in Navarra , Spain, in 1977 .
U pon the basis 01' cu ltural characte ristics and the deta ils 01' mo rp ho logy 01' its
penicilli, the spec ies is bel ieved to be more prop erly ass ignable to the
Asymmetrica -divaricata sub-section than to any other. The presence 01'
orange, more 01' less firm sclerotia, further indicates a relationship to the
Penicillium raistrickii series, hence , the present assignment.

79. Penicilli um mirabile Belj a ko va el Milko : fig. 79 , co lour p late 79 , and

SEM p late 14 , (to p : le ft).
Penicilliummirabile Beljakova et Milko - Mikologia i Fitopatologia . 6: 145-146. 1972.
Beljakova and M ilko's original diagnosis is presented here, since this adequately
presents the species in question when first isolated:
'Colonies upon Czapek's agar attaining a diameter of 40-45 mm in 13 days at
25C, with shallow radial furrows, depressed in the centre, velvety, fimbriate,
zonate , heavily sporing, at first (after 13 davs) colourless in central zone, olive gray

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Fig. 79. Camera lucida

drawings of Penicillium
mirabile Beljakova et Milko,
eBS 624.72. A, Detailed
drawings of the penicilli. B,
Mature conidia. e, Habit
sketches.

to deep olive, copper yellow in marginal areas, then (after 3 weeks) becoming deep
brown coloured, with growing margin narrow and uncoloured; exudate Iimited in
amount, deeply enmeshed in the mycelial felt in central zone, colourless; colon y
reverse strongly wrinkled in the centre and slightly furrowed in marginal area,
developing bright chamo is shades, then becoming black with the production of
sclerotia in central area, (atter 4 weeks) becoming black in the centre and in some
marginal spots. The conidiophores are sparsely produced, septate, up to 500 ~m in
length by 2.3-3.5 ~m in diameter, with apices enlarging in a vesicle up to 4.0-4.5
~m in diameter, arising either from the substratum or from aerial hyphae as short
branches, mostly 10-40 ~m up to 150 ~m in length by 1.5-2.5 ~m in diameter, with
walls smooth or roughened. The penicilli are one- or two-stage branched (one-stage
branched in those arising from aerial hyphae), 100 ~m long by 130 ~m in width.
Metulae divergent, 6.5-10.5 ~m (up to 13 ~m) by 2.5 ~m, with apices inflated in a
vesicle up to 4.0-4.5 ~m (up to 5.0 ~m) in diameter, in c1usters of 4.6-9 elements in
the verticil. Phialides bottle shaped, 8-1 O ~m by 2.0-2.5 ~m, terminating in a long,
tapered neck, in clusters of 5-8 (up to 11) elements in the verticil, seldom appearing
monoverticillate. Conidia are ovoid, 3.3-4.0 ~m by 2.5-3.3 ~m, echinulate, with

eonid ial chalns show lng conspl cuou s conneetlv es between them, at first
uncoloured, then developing deep olive shades. Sclerotia are black and submerged
in the substratum.
Colonies on grape must agar growing rapidly, attaining a diameter of 75 mm in 13
days; radiately wrinkled, with centre raised, velvety, azonate, with a leathery clcse
textured basal felt, heavily sporing, at first in olve-slate gray shades, with margin
copper yellow, developing deeper slate gray -brown shades (after 3 weeks) without
marginal copper yellow colours, with a narrow growing margin (1-2 mm) colour
less; colony reverse radially wrinkled, at first showing leather brown shades, then
(after 3 weeks). developing dirty violet-brown shades, blackening in central areas
(owing to the production of sclerotia). The sclerotia are abundantly produced,
either partially submerged or completely in the substratum. The remaining charac
teristics duplicate the above description .
Upon potato-dextrose agar the sclerotia are abundantly produced, leathery, at
first colourless, then becoming deep brown to black coloured, with walls consisting
of polyhedral cells, 9-15 .tm by 5-10 .tm in diameter, being produced either super
ficially or submerged in the substratum.'
(Translated from the Russian by Prof. L. Vilasl.

Our not es follow :

C olonies upon C A (co lour plate 79 , A and a) gro wing restri ctedl y , attaining
a di amet er of 25 mm in 14 days at ro om temperature (20 - 25C), deeply
velvct y to lanate , cons isting of a close- tex tured basal felt , plane in marginal
are as and slightly raised in th e ce ntre; ea lony surface overla id by a m ycelial
overgrowth cons isting of a network of flocea se hyphae influen cing th e colony
appearancc; thin , eve n m argin surro u nded by a frin ge of su bme rged h yphae
exte nd ing abo u t 1- 2 mm beyond th e ae rial gro wt h; m edium spo ring in dark
brown shades, with conid ial areas partially obsc u re d by th e floccose
ove rg row th of aerial hyphae ; odou r and exu date lacking; coloured pigment
di ffusin g int o the surro u nd ing agar abse nt ; colony reverse (colou r plat e 79, a)
in cin namo n shad es; peni cilli typica lly bi verticillat e asy m metrical (fig. 79 , A),
on e- or two-stage branch ed , with occasi on al single or monoverticillat e penicilli
present ; stipes of th e conid iopho res septate , sm oo th wall ed , 3. 0 p.m in dia
m eter, terminating in clusters of 2-5 wedge-shaped m etulae , m ostly 10-12
psi: by 3- 5 iun , definitely div erge nt, su ppo rting clus ters of bottl e-shaped
phialides, 8 - 13 p.m by 3. 0-3.5 tun, terminating in a conspicuo us ly lon g,
narrowed neck ; conidia ellip tica l wh en you ng, becoming subglobose to globo se
at m aturity , strong ly sp in ulose wh en obse rve d under th e light m icroscope ,
suggesti ng th e chestn ut fru it (fig. 79 , B) a nd strongly echin ula te whe n obse rved
under th e SE M (pl at e 14, top : left) , m ostly 3 .3 -4.5 p.m in long axis. Sclero tia
have not been obse rve d on thi s su bstra tu m . C onidial chains loosely parallel
(fig. 79 , C) .

olon ies upon C YA (COIOllr plat o 79 , B a m i b) growing mo re ra pid ly than

u po n C A , atta ini ng a diam crer 01' 4J -.'"() IIlJ1l in 11 days a l roo m tc ru pern tu re ,
consisting 01' a close- tcxtu rcd basal felr, decpl y velv ct y lo su bla natc in
a ppcanmc c, dccpl y radi a tely wrinkled, with ce ntre irregul arl y fold cd in a
cra teriform pattern , with undulating, thin m argin , surro u nded by a fringe 01'
su bm ergc d h yphae exte nd ing a bo u t 1 m m beyond the ae rial gro wth; colony
surface ove rlaid by a thin rn ycelial ove rgrow th cons isting of a den se network of
fine floccose hyphae influen cin g th e colony a p peara nce; heavily spo ring
throu gh out in brown shades, but with conidial areas obsc ure d by th e floccose
ove rg ro wth , lending cin namon shade s to th e colony su rface ; exudate limited in
a mo unt; odo u r faint , indefinite ; am be r-coloured pigm ent diffu sin g int o th e
surrounding agar; colony reverse (colour pIate 79 , b) in reddish brown shades;
pcni cilli esse ntially as describe d aboye; sclerotia no t ob served upon this
su bstr atum.
Colonies upon M EA (colou r plat e 79, C a nd c) a ttaining a d iam et er of 45
mm in 14 days at room ternperature (20 -25 C), cons isting of a looser textured
bas al felt , velve ty to gra n ula r in a ppe rance, plane , with tu fts of floccose , white
aeria l hyph ae in th e colony centre; thin , even m argin surro unde d by a fringe of
sub merge d hyphae extend ing abo u t 3 mm beyond th e ae rial growth; heavil y
spo ring thro u gho u t in deep brown sha des (S guy , PI. XLVI , 68 1); ex udate
and odou r lacking ; no coloure d pigm en t di ffusing into the surrounding ag ar;
colony reverse (colour plat e 79, e) in pale reddish brown sha des . Sclerotia not
observed on th is substra tu m . Penicilli essentially as descr ibed aboye; conid ial
chains ta ngled .
Penicill ium mirabile as originally describ ed and a s observed in ou r cu rrent
study di ffers only in its failu re to produce sclero tia up on all su bstrata tested .
O n th e basis of cultural an d m orphological characteristics, P. mirabile is
believe d to be m or e prop erl y a ssignable to th e Asymmetri ca-divaricata su bsection th an to any other sec tion , a nd th e reported presen ce ofblack sclerotia in
th e typ e strain , as originally described , fur th er indica tes a close relations hip to
the P. raistrickii series; hen ce , th e present ass ignme nt.
The spec ies description is centered up on th e typ e stra in CBS 624 .72 = V KM
F- l ,328 = C M I 167 ,383 = IJ FM 7101 , isolat ed from soil in C rimea, U SSR , by
L.A. Belj akova a nd A .A Milko , in 1967 .
No additional strai ns cIearly representing thi s species ha ve been encou ntered so far.
Penicillium raistrickii and allied spec ies are encou ntered occasionally as
isolat es from soil 0 1' from soil contam inate d m at eri als.

Penicillium lilacinum sc rres

O U TSTANDlNG FEAT U R ES

Co lon ies typically loose text ured lo lan at c , growing rapidly , slightly to
heavily sporing , in lilac , ochre or avc lla ncous (never grcen) shades.
Conidiopho res va rying gre a tly in len gth , arising from the su bs trat um or as
branches from asce nd ing hyphae , with walls smooth or finely roug hened .
Pen icilli typi cally divaricate , va riable, usu all y cons isting of terminal clusters
of d ivergent metulae, or a ppearing as sim ple structures when borne on short
branches fro m asc end in g stipes .
P hialid es typically termin at ing in a lon g, ta pered , narrow neck.
Conid ia gen erally strongly ellipt ical to lemon sha ped, com mo nly a ppea ring
apiculate , smooth walled or nearly so .
K ey to sp cci cs
Paga
1. Colonies not producing sclerotia or masses of thick-walled cells.
Colonies not showing green, gray -green, or blue-g reen colours with the
ripening of conidia. Colonies deeply velvety or lanate, developing lilac,
ochre, brown, or avellaneous shades .
P. Iilacinum series
a. Colonies tanate. developing Iilacinum shades, with reverse similarly
coloured.
P. lilacinum Thom
(Paecilomyces lilacinus Samson)
b. Colonies velvety, comparatively thin, not developing lilacinum
shades.
1'. Colonies spreading broadly upon most substrata , developing
pinkish buff to avellaneous shades.
P. humulivan Beyma
2' . Colonies growing very restrictedly upon Czapek's yeast extract
agar.
aa. Colon ies developing avellaneous shades upon malt extract agar.
P. argillaceum Stolk, Evans et Nilsson
bb. Colonies developing brown to ochraceous shades upon malt
extract agar.
P. cylindrosporum Smith

. .. . 228

.... 229

. . . . 231

... . 233

.. .. 236

R ap er a nd T hom (1949) erec ted th is series to accomodate th ose spec ies of

Penicillium which did not develop any gr een or blue-green shades wit h the
ripen in g of conidia . Strictly speaking, th e series included Penicillium lilacinum
Thom a nd two addition al species th at wer e conside red with it as a matter of

co nvcn icncc, na mcly P. humuli van Bcy m a a m i 8/Jicaria uiolacea Abbou

(consid ered by Brown a nd Sm ith (19:>7) as a synon YIll 01' Paecilomyces marquandii
(Massec) Hu gh es. For similar rca sons , we have included in ou r cur re nt study
two additional species , name! y P. argillaceum Stolk, Eva ns el Nil sson, and P.
cylindrosporum Smith , both charac te rized by th eir d ivarica te penicilli , and by
th e lack of gr een and blue-gr een colou rs in their colon ies .
Penicillium lilacinum T ho m, altho ug h recently described in th e gen us Paecilomyces (Samson, 1974) , as Paecilomyces lilacinus Sa mson, is still included here
primarily as a m a tter 01' conven ience, and also be cau se we fee! that its
Penicillium-like m orphological cha rac teristics justify its in clu sion in any
lr eat m ent al' the gen us Penicillium.
H owever , a close rclationship betw een the m is not presumed here .

SO. Penicill ium lilacinum Thom : fig. 80, colou r plate 80, and SE M piate
14 , (top: right).

11

O 10 O \J
0IJO o\:)
o

~ \J~

10 prn

, c~

~.~
'~ "

eR

~J/

;:/

Fig. 80. Camera lucida

drawings of Penicillium
lilacinum Thom
(Paecilomyces lilacinus
Sarnson) , CBS 284.36. A.
Detailed drawings of the
penicilli. B, Matu re conidia.
C. Habit sketches.

Penicillium Ii/acinum Thom - Bull , Bur. Anim. Ind . U.S . Dep. Agric. 118: 73-75.
1910. See also Ch. Thom, 'The Penicillia', pp. 331-334. 1930.

Combinatlo nova
Paecylomyces Iilacinus (Thom) Samsom - Studies in Mycology. 6: 58-62. 1974.

Synonymies
? Penicillium amethystinum Wehmer sensu Biourge - in Biourge's Monographie,
La Cellule. 33: 221. 1923.
Graphium cicadicicola Spegazzini - An. Mus. Nac. Hist. Nat. Buenos Aires. 20:
476. 1911.
Penicillium (Scopulariopsisl rubellum Bainier sensu Biourge - in Biourge's
Monographie, La Cellule. 33: 221. 1923.
Spicaria violacea Petch - Trans. Brit. Mycol. Soco 16: 239. 1931. (non Abbott,
1926).
Spicaria rubidopurpurea Aoki - Bull. Ser. Exp. Stn Jpn. 10: 440. 1941.

Colonies upon C A (colour plat e 80, A and a) a ttain ing a diameter of 35 m m

in 14- days at room temper ature (20 -25 C) , co ns ist ing of a loose-textu red basal
felt , 1- 2 mm deep , lanate , with centra l area ra ised , not furrowed , azo natc ;
white, thin, eve n m argin su rro u nde d by a frin ge of su b me rged hyphac ,
extend ing about 1-2 mm beyond th e ae rial gro wt h; colony surface at first
white , gradu ally devel oping lilac to vinaceous shades with th e rip ening of
con id ia ; hea vily sporing throu gh out ; exudate limited in amo un t , as colou rle ss
d ropl ets en meshe d in th e myceli al felt ; odo ur lacking; pa le amber-coloured
pigm ent d iffusing int o th e su rro u nd ing agar; colony rcver se (colour plate 80, a)
in pale ora nge sha de s; peni cil1i va ryi ng in size and complex ity (fig . 80, A),
cons isting of whorl s of m ctulae arrange d at different levels or nades; th e stipe s
of th e conid iopho res m ay va ry greatly in dimen sions, dc pe nd ing upon its
origin; whe n a rising from th e su bs tra tum at the colony margin , th ey range
from 400 to 600 tut: in len gth by 3.0 -4 .0 /lm in di amet er: wh en borne from
ae rial hyphae th ey m ay ran ge fro m ve ry sho rt, up to 100 - 200 /lm in length by
2.0- 3.0 /lm in di ameter , with wa l1s smoo th or delicately rou gh en ed; metulae
in small who rls of 2-3 eleme nts in th e ve rticil, m ostl y 6- 10 /lm by 2.0-2 .5
uitv ; phialides bottle shaped , u su al1y 7- 9 /lm , occasionally up to 11 /lm by
2. 5- 3.0 tun , abru ptly ta pe ring to a com paratively long, narrowed neck ,
conid ia elliptica l to fusiform , m ostly 2.5 -3.5 /lm by 1.5-2 .0 tui , smooth
wall ed or slightly ro ug he ne d (fig . 80 , H and SEM plat e 14, top : right) ; conid ial
cha ins loo sely parallel or tangled (fig. 80 , C ).
C oloni es upon C YA (colou r plat e 80, B a nd b) a tta ining a diameter of 35-40
mm in 14 d ays at roo m tem pe ra tu re (20 - 25 C), cons isting of a closer textured
bas al felt , ove rlaid by a close ly woven net work of white ae rial hyphae influen cin g th e colony appeara nce, deepl y velve ty to lanat e , deepl y radiately wrinkled ,
with cen tre rai sed and depressed in a crateri form pattern ; heavily sporing in
th e same shades as aboye, bu t partially obsc u re d by th e dense ove r gro wth of

stc r ilc aeria l h yph ac ; cvc n m a r gin surroundcrl by a Iringc 01' sub mc rgcd
hyph ac , cx tc ncliru; about 1 111111 bcyond thc ae ria l grow th; cx ucla tc and odo ur
lackin g: a m be r-co lourc d pi gm cnt diffusi ng inr o t.hc surro u nd ing aga r; colony
reverso (colou r plat e 80, b) in rc dd ish brown to ci n na rnon shadcs ; pcni cilli
csscntiall y as dc scrib cd above.
Colon ics upon M EA (colour platc 80 , C and c) attaining a d iamcter 01' 40
IlIlll in 14 days a t room te mperature (20-25C), cons isting of a loose-t extured
basal Ielt , ovc rlaid by a looscly wove n net work 01' floccose hyphae , len ding a
lan ate appea rance to th e colony surface; heavily sp oring th rough out in lilac
sha des; colon y surface not wrinkled, slightly raised in th e cent re, with thin
margin surrou nde d by a fringe 01' sub me rge d hyphae exte nding abo u t 2 mm
bcyond th e ae r ial growth; odo u r a nd cxudate lackin g; no coloured pigm ent
c1i ffusin g into th e surro u nd ing agar; colony reverse (colour plate 80 , e) in
sienna brown sha des; peni cilli essentially as described aboye .
The spec ies description is ce ntered up on th e typ e stra in CBS 284 .36 = CM !
27,83 0 = ATCC 10,11 4 = ATCC 1,11 23 = N R RL 895 = Q M 7,592 = IJ FM 5 116,
isolat cd from soil by C h . T hom in th e USA, a nd m any add itional cultures
isolat ed by the author from Span ish so ils. Amo n g th ese strain IJ FM 5 150,
isolat ed as a cultu re conta m ina n t in M ay 1978 , m ay be regarded as
rep rescntativ e.

8 1. Penicillium humuli van Be ym a: fig. 8 1, colo u r platc 81 , a n d S EM

pl at e 14 , (ce n t re: left )
Penicillium humu/i van Beyma - Z. Bakt. Parasitenk. Inf. Hyg. 11 Abt. 99: 392-394.
1939. See also Raper and Thom, 'A Manual of the Penicillia', pp. 291-294.1949.

Colonies upon CA (colour plate 8 1, A and a) spread ing broadl y, a ttaining a

dia meter 01' 50 mm in 14 days at room temperature (20- 25 C ), cons isting 01' a
close -tex tured basal felt , tough , ove rlaid by a close ly int erl aced ove rg ro wth 01'
trailing hyphae (desc ribe d by va n Beym a as ' powdery ' ), velve ty, slightly
zonate, with thin , even m argin surro u nde d by a frin ge 01' su bme rge d hyphae
extending abo u t 2 mm beyond th e ae rial growt h; spo ring sparingly in cream
shades; colony su rface stro ngly folded in a radi al pa ttern ; odo ur and exudate
lacking; no di ffusin g coloured pigm en t into the surrou nd ing ag ar; colony
rever se (co lour plat e 8 1, a) in pale cream rosy shades; penicilli m ay appear as
sim ple 0 1' m onoverticillat e struc tu res borne on short stipes, 50 - 100 .tm by
2. 0-2 .5 uit , whi ch arise as branch es from trailing hyphae , 0 1' more com plex,
as cons picuo us ly div aricat e structures irregularly b ranch ed (fig. 81 , A), consisting 01' a whorl 01' 2-3 m etulae, ranging from 20 to 35 .tm in length by
2.0 -2 .5 .tm in di amet er, with apices slightly enla rge d in a ves icle up to 4 .0 .tm
in diamet er, borne on longer stipe s, up to 250 uit: in len gth , typi call y arisin g

o0
O

0o 0
0 0 11
0

~O O

00

0 00

00

"",Ve-"":.}
.

----==---

100 J.lm

..

1~

f..

Fig. 81. Camera lucida

drawings of Penicillium
humuli van Beyma, CBS
231.38. A, Detailed drawings
of the penicilli . B, Mature
conidia. C, Habit sketches.

from th e basal felt , sm ooth wall ed ; phialides bottle shaped , usually in clusters
of 2- 8 elem ents in th e verticil, mostIy 9-15 p.m by 2. 5-3 .0 uitv, terminating in
a com pa ra tively long, narrowed taper ed neck ; conid ia elliptica l to su bg lobose,
sm oo th walled , m ostIy 3 .0- 4 .0 p.m by 2. 8 -3 .5 p.m (fig. 8 1, B, a nd SEM plate
14, centre: left) ; conid ial chains loosely tangled (fig . 81, C) .
Colon ies upon C YA (col our piate 81 , R and b) attaining a diameter of 60
mm in 14 days at room temper ature (20 - 25 C) , consisting of a tou gh , close textured bas al feIt , strong ly furrowed in a radial pattern , rais ed gra dually
tow ar ds th e ce ntral arc as, wh ere th e m yceli al felt is ofte n cra cked on upper
surface s of folds, and strongly and irregula rly fold ed ; heavily spo ring
throu gh out in cinna mo n shades , partially obsc u re d in ce ntral a nd subcentral
arcas by a thin and dense ove rg ro wth of close ly int erl acin g trailin g white
hyphae; eve n thin m argin largely surro u nde d by a fringe of su bme rge d hyphae
exte ndi ng more tha n 5 mm beyond th e ae rial gro wth; ex udate and odo u r
lacking; coloured pigment diffusing int o th e surrou nd ing agar not produced ;
colony reverse (colour plat e 8 1, b) stro ng ly pulling away th e agar underneath,
developing cin namo n shades ; peni cilli esse ntially as described aboye.

olon ies up on M EA (colour pla te ni, e a nd e) spread ing broadly , uu a ining

diamet er 0 1' 65 - 70 mm in 14 days at room tempcraturc (20 - 25C) , vclvcty ,
planc, with vcry thin margin su rrou nde d by a wid c I'ringe 01' submerged
hyphae cxtcnding ab out 5 mm bcyond th e ae rial growth ; heavily sporing in
cinnamon shades (Sguy , PI. XIV , 204), lending a powdery appearance to the
colony surface ; odour and exu date lacking; colou red pigm ent diffu sin g into the
surrou nd ing a gar not produced ; colony reverse (colour plat e 81 , c) in pale
crcarn shad es; peni cilli essent ially as described aboye, but with con idiopho res
arising mostly from the substratum.
The spec ies description is cente red up on the typ e strain C BS 213. 38 = C M I
39, 8 17 = AT CC 10,452 = NR RL 872 = IF 7,726 = IJFM 5115, isolat ed from
Lupulus humulus L. by Dr H . Schnegg in W eih en stephan in 1938.
N o add itional strains clearl y representing this spec ies have been encountered so fa r.
11

82 0 Penicillium argillaceum StoIk , Evans el NiIsson : fig. 82 , coIour pIat e

82 , and SEM pIat e 14, (centre: right) .

"':/i Ov
",
'V/'

"

:;;

r,

O~~

o,
' 0

!!:

0 0

{/

\)("\OO

'. BJJ=

'. 0

~: .

1::.:::

00'

: -,

'i,.

\:'' t
.
;

~m

r ,

~; l

"~~
~ ~ lC

L.-...-l

100

jrrn

eRo

Fig. 82. Camera lucida

drawings of Penicillium
argillaceum Stolk, Evans et
Nilsson , CBS407.73. A,
Detailed drawings of the
penicilli. B, Mature con idia .
Habit sketches,

e,

Penicillium orgillocoum Stolk, Evans e t NlIsson - Trans. Brlt. Mycol. Soco 53:
307-310. 1969.

Synonym
Geosmithia argillacea (Stolk, Evans et Nilsson) Pitt - Can. J. Bot. 57: 2021-2030.
1979.
The author's original diagnosis follows, since this adequately presents the species
in question:
'Colonies upon Czapek's agar attaining a diameter of 4-4.5 cm within 2 weeks at
30C; with surface growth almost velvety; somewhat loose textured, consisting of a
very thin network of hyphae either above or below the agar surface from which
conidial structures arise, in central areas overlaid by a thin overgrowth of loosely
interwoven hyphae bearing also penicilli; azonate; with margin irregularly outlined,
feathery; sporulating somewhat irregularly, showing brownish colours which range
from chamois and cinnamon-buff to avellaneous (Ridgway, PI. 30, 29, 40); exudate
lacking, odour reminiscent of pine resin; reverse of colonies uncoloured at first,
becoming yellowish or pinkish with age.
Colonies on malt extract agar spreading broadly, attaining a diameter of 6-6.5 cm
within 2 weeks at 30C, in texture and colour as on Czapek's agar but sporulating
more abundantly and with margin even , circular in outline; reverse uncoloured to
avellaneous.
Vegetative hyphae hyaline, smooth walled, 1.5-3 .tm in diameter. Conidiophores
arising either directly from the substratum or as branches from aerial hyphae,
varying greatly in size, ranging from 60 to 400 .tm in length by 2-3 (4) .tm in diameter, hyaline, septate, with walls usually roughened, but occasionally smooth
walled. Penicilli biverticillate-asymmetric, varying in complexity , mostly consisting
of somewhat appressed verticils of two to five metulae, bearing c1usters of
phialides, but occasionally developing larger penicilli showing one or two rami in
addition to the main axis . Monoverticillate heads are also presento Rami and
metulae, as well as metulae and phialides, may occur in the same verticil. AII
elements of the penicillus are hyaline, usually rough walled, but occasionally
smooth walled . Apices of conidiophores in monoverticillate heads are mostly somewhat inflated . Rami 12-35 by 1.5-3 .tm . Metulae 12-25 .tm by 1.5-3 .tm enlarged at
the apex to 4 .tm in diameter. Phialides 9-14 .tm by 1.5-2 .tm, typically occurring in
dense c1usters up to 10 in number, but in reduced smooth-walled penicilli mostly
few in number; consisting of a narrow cylindrical base, tapering more or less
abruptly to a narrowed conidium-bearing tube (neck), about 1-2 .tm by 1 .tm.
Conidia (2.5) 3-4 (4.5) .tm by 1.2-2 .tm, varying from cylindrical to ellipsoid and
ovoid, hyaline, smooth walled. Conidial chains tangled or adhering in loose , twisted
columns.
P. argillaceum is thermotolerant, having its minimum temperature at about 15C,

It9 optlmum at about 36C and ita maxlmum at about 6O oC . At high temperatures
(40-S0 0Cl the cultures are predominantly floccose, sporulating only poorly.'

O ur not es follow :
Colon ies upon CA (colour plat e , 82 , A and a) attaining a diamet er of 30- 35
mm in 14 days at room temperature (20- 25C), consisting of an extremely
loose-textured basal fe1t, mostly submerged, from which arise very sparingly
conid ial str uc ture s, lending a powdery appe a ran ce to th e colony, and a
cinna mo n coloured surface ; exu da te lacking; odour faint, aromatic; colony
rcverse (colour piate 82, a) colou rle ss; penicilli essentially as originally
dcscribed, with th e conid iopho re s arising from th e basal fe1t and from
uniculose hyphae (fig . 82 , A), strongly verrucose including th e phialides;
conid ia mostly cylindrical , 3 .5-5 p-m by 1.2-2 .0 p-m in diamet er, smooth
wall ed (fig . 82, B, and SEM plate 14, ce ntre : right); conidial ch a in s loose1y
tangled (fig . 82 , C) .
Colonies upon CYA (colour piat e 82 , B and b) growing restrictedly ,
atta ini ng a diameter of 25 mm in 14 days at room temperature (20-25 C) ,
consisting of a close-textured bas al felt overlaid by a white mycelial overgrowth
consisting of a closely interwoven network of ae rial hyphae influencing th e
colony appearance ; colony surface appea ring sublanate , slightly raised in the
centre; very poorly sporing, remaining persistently white ; exudate lacking;
odour faint , indefinite; no coloure d pigment diffusing into the surrounding
agar; colony rev ers e (colour piate 82 , b) colourless with cinnamon sh ades
showing through; penicilli essen tially as described aboye .
Colonies upon MEA (colour plate 82, C and e) growing more rapidly, attaining a di ameter of 40 mm in 14 days at room temperature (20-25 C) , consisting of a dense basal fe1t overlaid by a close1y interwoven network of floccose
ae rial hyphae , lending a lanate appearance to th e colony ; m argin irregularly
di ssected , surrounded by a fringe of submerged hyphae ext ending about 3 mm
beyond th e aerial growth; he avily sporing throughout in ave llaneou s shades
(Sguy, PI. XVII, 249); not furrowed , with tufts of white floccose a erial
hyphae in cen tral areas ; exudate lacking; odour faint, a rom atic ; no coloured
pigment diffusing into the surrounding agar; colony reverse (colour plate 82 , e)
in yellow to ochraceous shades; penicilli essentially as described aboye .
Penicillium argillaceum is be1ieved to be close1y re1ated to P. cylindrosporum
Smith, on account of th e av ellaneous to ochraceous colours of th e colonies, the
morphological details of its penicilli and the shape of th e conidia . The two
species, however, differ in th eir phialides, which are conspicuously roughened
in P. argillaceum and smooth wall ed in P. cylindrosporum , Moreov er, th e two
species ca n easily be distinguished from one another by their cultu ral aspe cto
Colonies upon MEA of P. cylindrosporum have a very dark gr eenish, almost
black rev erse (colour plate 83, e), while this pigment is lacking in P. argillaceum .

In add itio n , th e th e rmot olc ran t P. argillaceum gro ws a l ternperatures up lo

SooC, whil c P. cylindrosporum docs nol gro w a l 1S oC , growing vcry re strictcdly
a t a H tcmpcraturcs upon C A (co lour plate 83 , A a nd B).
T he spec ies description is centered upon the typ c strain css 101.69 , isolatcd
fro m a mine tip with a ve ry high surfac e tcmper ature in Sta ffords h ire , UK by
Dr H. C . Evans, M ay 1967 , a n d upon th e strain css 407.73 = IJ FM 1405 ,
isolated by the author fro rn Piper nigrum in 197 3 , Madrid , Spain .
Four ad di tio nal strai ns wer e isolat ed by H .C . E vans a nd Dr T. Nil sson,
Skogs h kola n , St ockh olm , Sweden : on e frorn th e a ir, b y H. C . Evans , two
fro m sp ruce ch ips a nd fro m pine ch ips by T . Nil sson . The Sw edi sh strains
di ffer slig h tly from th e type st rain in producing upon Czape k 's agar thinner,
almost colou rIess colonies, ve ry poorIy spo ri ng .

83 . P enicillium cylindrosporum Sm ith : fig. 83, colour pl ate 83, a n d SEM

plate 14 , (b ottom : Ieft).

<::::J

OOO '::

c-..... B~ e'!
"V

c::::J""

O~

~o IJ
Cl ~

'~1-::,,~
~;j (t~

JI<

y
{-

~
L..----l

100J.lm

Fig. 83. Camera lucida

drawings of Penicillium
cylindrosporum Smith , eBS
275.58. A, Detailed drawings
of the penicill i. B, Mature
con idia. e, Habit sketches .

Penicillium cy/indrosporum Smith - Trans. Brit. Mycol. Soco 40: 483-484. 1957. See
also M. M. Kulik, 'A Compilation of Descriptions of New Penicil/ium species' .
Agric. Handbook No. 351. Agric. Res. Ser. U.S. Dep. Agric. p. 34. 1968.

Synonym
Geosmhie cvlindrospore (Smith) Pitt - Can. J . Bot. 57: 2021-2030. 1979.
Smith's original diagnosis follows:
' Colonias on Czapek agar thin velvety, inclined to be patchy, dull avellaneous,
with reverse uncoloured; on malt agar growing fairly well , velvety, dull avellaneous
wth sordid yellow margin and with reverse dark dirty greenish brown; on potato
ogar similar except that the reverse is somewhat more greenish; stated by Mrs J.
Robson (frorn whom the original culture was received) that green perspiration drops
ore produced in old cultures, but these have not been observed in subcultures;
penicillus sometimes with a more or less divergent cluster of metulae (the P.
citrinum tvpe) . but more often irregular, with a branch or branches from lower
nodes, bearing phialides alone or metulae and phialides mixed in the same verticil ,
somet imes showing the same mixed verticils in the terminal part of the structure;
eonidiophores long and sinuous, mostly arising from submerged mycelium, rough ,
2.5-4 .tm in diameter; rarni, when present, rough, 16-25 .tm by 2.5-4 .tm; metulae
usually 12-20 .tm long, but shorter, 9-10 .tm, in the more complieated heads, and
2-4 .tm in diameter; phialides almost cylindrical , tapering abruptly at the tip, very
regular in length, 11 .tm by 2.5-3 .tm ; eonidia usually cylindrical (hence the specific
epit het ), 4.2-5.5 .tm by 1.8-2.2 .tm, mostly 5 by 2 .tm, but oecasionally ovate to
ellipsoidal, 3.2-3.5 .tm by 2.2-2.5 .tm; eonidial chains divergent then tangled, or
sometimes in several loose, twisted columnar rnasses.'

Our not es follow :

Colon ies upon CA (colour plat e 83, A a nd a) gro wi ng extre me ly rest rictedl y ,
atta in ing a di ameter of 10 mm in 14 days at room te m pe ratu re (20 -25 C),
cons isting of a loose-textured b asal felt , velvety, pla ne, with m argin irregul arl y
di ssected su rro u nde d by a n arrow fring e of sub me rge d hyph ae exte nd ing less
than 1 mm beyond th e ae rial growth; spo ring through out in avella neo us
shades ; no exu date produced; odo ur lacking ; no coloured pi gm ent d iffus ing
int o th e su rro u nd ing agar ; colony reverse (colou r plat e 83, a) colou rless;
penicilli (fig . 83, A) esse ntially as or iginally descr ibed ; conid ia stro ngly
cylind rica l, smooth wa lled , up to 6 tut: in len gth by 1.5-1.9 .tm in d iameter
(fig. 83, B, and SE M plate 14, bottom : left) .
Colonies upon CYA (colour plate 83, B a nd b) atta in in g a di ameter of
7-8 mm in 14 day s at roo m temperature (20- 25 C), consist ing of a closetex ture d ba sal felt , velvcty , conve x, almos t ste rile, rem aining persistently
white; exudate and odo ur lacki n g; coloured pigment diffu sing in to th e surrounding a ga r not produced ; colony revers e (colou r plate 83, b) in cinnamo n
shades .
C oloni es upon M EA (colour pIate 83, C and e) , gro wing more rapidly th an
upon th e preced in g su bstrata, a tta ining a diameter of 20- 25 mm in 14 days at

room tcmper at u rc (20- 25C), typically vclv cty, pla no, with m argin il'rcgularly outlincd surrounded by l IHU 'I' OW Irin ge 01' sub me rged dark brown
hyphac ex ten d ing about 1 mm bcy ond th e acrial gro wth; hcavily sporing
throughout in dark avellaneous to ochraccou s shadcs; brown-coloured pigment
di ffu sing into th e surrounding ag a r ; colony rev erse (colour platc 83 , e) in d ark
green ish brown , alm ost black shades; penicilli csse n tia lly as originally
des cribed , with th e stipes 01' th e con id iop hores co nsp icuously warty (fig . 83, A)
a n d metulae usu all y smooth walled ; phi alides m ay range frorn 11 to 15 Jlm in
length by 2.5 - 3 .5 Jlm in di amet er.
The species descr iption is ce n te red upon th e typ e strain C BS 275.58 = C M I
71 ,62 3 = AT CC 18 ,223 = I FO 8 , 110 = Q M 7, 524 = IJF M 704 3, isolat ed as a
cu lture contaminant by Mrs J. Robson , 0 1' the At omic Energy Es ta blishmen t,
Harwell , UK , in 1957.
This species is to be elas sified in th e Asymmetrica -divaricat a su b-sec tion,
within th e Penicillium lilacinum series on accou n t 0 1' th e tot al la ck al' green 0 1'
blue colou rs in its colo n ies .

Penicillum Janthinellum series

UTSTA NDING F EAT UR ES

C olonies sp read in g braadly , deve!oping gray, gray -gree n colours with th e

rip ening 01' co n id ia; vegetative hyph ae becoming orange , ora nge-re d 01' reddish
purple in sorn e stra ins; colony surface varying from alm ost velv ety to definite!y
lanat e , a n d occasionally d ev e!oping funi cles; colo ny reverse a t first colou rless,
a nd in sorne strains remaining so, in othe rs deve!oping colo u rs ranging frorn
ye llo w 0 1' ye llo w-gree n throu gh ora n ge, 01' orange-red , to reddish 01' vinaceous
purple.
Stipes 0 1' th e con id iop hores a rising directl y frorn th e subs tratu m 0 1' as
branch es frorn tr ailing h yphae , ranging from very short up to 1000 Jlm in
len gth, with walls usuall y granul ar 01' raughened.
Penicilli variously ramified a nd ranging frorn simple a p ical wh orl s 01'
phialides (appearing m on ov erti cillat e) to a sym metric struc tu res co nsistin g 0 1'
a n indefinite number 0 1' strongly di varicat e branch es a n d/or m etulae bearing
ve rtic ils 01' phi alides.
Phialides few in th e verticil , slen de r, a n d typicall y terminating in a n
abruptly to co nspic uo us ly tapered , lon g , n arrow ed neck .
C onidia a t first definite!y ellip tica l, u su ally b ecoming ovate to su bglobose in
age, with one 01' both e nds 01't en pointed , a nd with wall s smooth 01' de!icate!y
rou gh ened .

Key to spccies
Page

1. Colonies not produciny sclerotia or masses of thick-walled cells.

Colonies showing some shades of green, gray, gray-green, or blue-green
with the ripening of conidia. Penicilli with the divaricate character well
marked; colonies growing rather restrictedly, at least upon some substrata; with metulae scattered on the conidiophore, or usually partly
aggregated into true verticils. Colonies developing typically pale bluegreen or gray-green shades with the ripening of conidia; colony reverse
often brightly coloured. Conidial chains strongly divergent and/or
becoming tangled in age, not tending to form compact columns.
Phialides abruptly tapering to a long, narrowed neck; colonies usually
not funiculose .
P. janthinellum series
A. Conidia conspicuously roughened, with roughness typically arranged
in spiral or transverse bands.
1. Conidia globose to subglobose.
P. daleae Zaleski
2. Conidia conspicuously apiculate.
P. asturianum Ramrez et Martnez
B. Conidia usually roughened, with roughness not arranged in spiral or
transversal bands.
1. Vegetative hyphae and colony reverse strongly coloured (orangered, reddish purple, etc.) in recent isolates.
P. janthinellum Biourge
2. Vegetative hyphae uncoloured or in slightly buff to peach shades;
colonies sporing sparingly or tardily; colony reverse colourless or in
yellow or orange shades.
a. Conidiophores delicately roughened; penicilli usually consisting
of terminal verticils of divergent metulae; conidia ovoid to subglobose, delicately roughened; colony reverse yellow to colourless.
P. simplicissimum (Oudernans) Thom
b. Conidiophore conspicuously granular or warty; irregular; conidia
elliptical, smooth walled or delicately roughened; colony reverse
developing orange colours.
P. ochro-chloron Biourge
c. Conidiophores smooth walled or nearly so; penicilli commonly
irregular; conidia mostly subglobose, conspicuously echinulate;
colony reverse in cream to light shades.
P. piscarium Westling

.. . . 238

. .. . 243
. . . . 254

. . . . 240

.. .. 246

... . 248

... . 250

d. Conidiophores smooth wall ed or nearly so; penlcllll Irregular;

conidia elliptical to subglobose , smooth walled or naarly so.
1'. Conidia comparatively larga, up to 6.5 ,m in long axis;
phialides comparatively large, up to 19 ,m in length.
P. aragonense Harnrez et Martnez .... 266
2'. Conidia comparatively small, up to 2.5 ,m in long axis;
phialides comparatively small, up to 10 ,m in length.
P. miczynskiiZaleski . . . . 252

The Penicillium j anthinellum series erec ted by Ch . Thom (1 930) to accomo datc
an unusualIy variable, individuall y and collectiv eIy, group of Penicillia with
colony texture rangin g from com pa ra tiveIy thin, veIvety, loose textured to
lanat e . They a re cha rac terize d by th eir m or e or less fast gro wing , more or lcss
bri ghtl y coloured colonies, by th eir irregular and stro ngly divergent penicilli ,
and particul arl y by thei r phial ides characteristicalIy tapering to a long and
narrow neck . Eight spec ies a re recognized in th e cu rre nt stud y , as follows:
Penicillium daleae Zal eski , P. asturianum Ramrez et M artnez , P. Janthinellum
Biourge , P. simp licissimu m (O ude mans) Thom , P. ochro-chloron Biourge , P.
piscarium W estling, P. aragonense Ramrez et Martnez , and P. miczinskii
Zaleskii. P. godleswski i Zaleski , characterize d by colon ies gene ra lIy more or less
funi cul ose , included by R aper a nd T ho m (1949) in thi s seri es, has be cn
exclude d from here to design a te th e newl y erec ted Penicillium godlewskii seri es .
Penicillium daleae and P . as tu r ia n um are distinguished particularly by th c
spira l ban ding of conid ia . Penicillium janthinellum is by far the mo st abundant
a nd most va riable. When first isolated , strains are com monly brightly
coloured, but u sualI y becom e less colou rful with con tin ue d labor atory
cultivation . Penicill ium simp licissimum is usualIy a heavil y spo ri ng form upon alI
su bstrata a nd less colou re d, eithe r in th e vegetative hyphae or in th e colony
rever se. Penicillium ochro-chloron is not eworthy for its conspicuo u sly warty
con id iopho re s and its tolerance to coppe r and high acidities. Penicillium
piscarium is a com pa ra tiveIy loose textured , lanat e form with su bg lobo se and
stro ngly echi n ulate con id ia . Penicillium aragonense, is a close textured, veIvety
form , cha racte rized by its com pa ra tively large , eIliptical, smooth walIed
conid ia. Penicillium miczynskii is di stinguished by its colonies at first white ,
velve ty, becoming blue-gr een with th e ripe n ing of conid ia, with yelIow
vegeta tive hyphae abu ndantly produced.

84 . Penicillium janthinellum Biourge: fig . 84 , colour plate 84, and SEM

plat e 14, (botto m : ri ght).
Penicillium janthinellum Biourge - Monographie, La Cellule. 33: 258-260. 1923. See
also Thom, 'The Penicillia', pp . 283-241.1930; also Thom in Pratt, J. Agre. Res.

L..-.........J

100}Jm

Fig. 84. Camera lucida

drawings of Penicillium
janthinellum Biourge, CBS
340.48. A, Detailed drawings
of the penicilli. B, Mature
conidia. C, Habit sketches.

13: 94-95. 1918; also Raper and Thom, 'A Manual of the Penicillia', pp. 299-303.
1949.

Synonymies
Penicillium echinulonalgiovense Abe - J . Gen. Appl. Microbiol., Tokyo. 2: 81.
1956.

Penicillium guttulosum Gilman et Abbott - lowa State Col. J. Sci. 1: 298. 1927.
Penicillium glauco-roseum Demelius - Verhand. Zool. Bot. Gesellsch. Wien . 72:
72 . 1923.
Penicillium meleagrinum, varo viridiflavum Abe - J. Gen. Appl. Microbiol. Tokyo.
2: 92. 1956.

Penicillium rivoliZaleski - Bull. Acad. Polonaise Sci. Math. Nat. Sr. B. pp. 471473. 1927.

Penicillium propium Morotchkovsky - Bull. Sci. Recueil Biol. Univ. Kiev. 2: 78 .

1936.

oloni es upon C A (eolour plat o B4, A and a) attu in ing 11 di amct cr 01' :~ 0 -:~5
mm in sorn e stra ins, while 60 - 70 mm in o thc rs, in 14 da ys al room
ternperature (2 - 25 C) , cons isting of a lou gh , closc-texturcd basal 1'c11
overlaid by a closely interwoven nctwork ol' fine hyphae lending a dclicat ely
lanate ap peara nce to th e colony su rface, whi ch is unevenl y tuftcd, abruptly
rai sed , or in sorne strains somewhat fu n iculose, irregul arl y wrinkled in th e
centre and radially furrowed in m arginal areas ; with an cve n margin
surrounded by a fringe 01' submerged hyphae extending about 1 mm beyond
th e ae rial gro wth ; at first white , but in recent isolat es becomin g va rious ly
colou re d from th e tardily and irregul arly sporing areas, mostly in pale gray
(Sgu y , PI. XXIX, 430) to glauc us gray (Sguy , PI. XXX, 444 ), and th e
sim ultaneous shadi ng 01' non-sporulating areas to dull bu 1'1', ora nge -red , or in
sorne strains purple vinaceous shad es; azonate to broadl y zonate; exudate
lacking or limited in a m ou nt; odo ur strong , typi cally suggesting a roma 01' red
beet; no colou re d pigment diffusing into the surroundi ng a gar; colony reverse
(colou r plat e 84 , a) colou rless in old isolat es a l'ter m an y tr ansfers, but usually
in bright shades in recent isolates, com monly ye llow-g ree n to orange at first ,
changing to ora nge -red, m aroon or purpl e-red colou rs; penicilli typi cally
asy m me tric, strong ly di varicat e (fig. 84, A), abu ndantly produccd in sorne
strains , less in othe rs, sometimes eve nly distributed throu gh out th e colony bu t
usuall y m ore abundant in submarginal areas, borne apically on asce nd ing
con id iopho res up to 500 tm in len gth by 2. 0 -3. 0 tm in diamet er , with wall s
smooth or fin e1 y rou gh en ed , or on sho rt branc hes from ae rial hyphae , m ostl y
10- 50 tut: by 2.0 -2.8 tut , va rying greatly in com plexity, rangin g from simple
verticils 01' phialides (appearing monov erticillate) to verticils 01' m etulae 01'
unequal length , or ve rticils of both m etulae and phialides ; branch es va ria ble,
rangin g from 10 to 25 tm by 2. 0- 2 .3 tu ; metulae , m ostl y 10-1 5 uu: or
lon ger, up to 20 tm by 2 .0-2.2 tun, with a pices slightly enla rged in a vesicle up
to 3.0 tm in diameter; phialides divergent , va ria ble in dimensions, ranging
from 9 to 20 tm by 2.5-4.0 tm in the sim ple pen icilli to 7-11 tm by 2. 5-3 .0
tm in th e biverticillate asymmetrical str uc tu res , abru ptly tapering in a long,
narrowed ne ck , characte ristic 01' the series; con id ia strongly ellipt ical wh en
formed , u su all y rem aining so , but in sorne strains becoming ovate to
su bglobose with ends 01't en apicul at e and with walls more or less rou gh en ed
(fig. 84 , B , a nd SEM plat e 14, bottom : ri ght) , m ostly 3- 4 tm by 2.8 -3.2 tm
in di amet er; conid ial chains 100se1y divergent (fig. 84, C ).
Colonies upon C YA (colour plat e 84, B and b ) attaining a di ameter 01' 50
mm in 14 days at roo m temperatu re (20 -25 C), cons isting 01' a tou gh , closetextured ba sal felt , overl aid by a closely int erwoven network of fine aerial
hyphae lending a su bla nate a ppearance to th e colo ny surface, dee ply ra d ia tely
furrowed in su b marg ina l and m argin al arcas, with cent re rai sed and depressed
in a crateriform pattern ; zonate; thin , eve n m argin with a fringe 01' su bmerged

hyphae cx tc nd ing ab out 2 mm bcy ond th e Heria) gro wth ; heavily sporin g
thr ou ghout in bro wn ish {!'aY shade s, partially obscurcd by the dense network
1I1' fin e acrial hyphac d cli cat cly tintcd in lilac shadcs , cspcc ially in cen tra l a nd
su hcc n tral arcas; ycl low-co lo urcd pigmcu t di llusing into the surrounding agar;
odour strong, typically suggest ing red bce ts; ex udate lacking; colo ny reve rso
(cnlour plate 84 , b) in violet to ora nge sh ade s ; pcnicil1i essential1y as descri bed
ubove .
C olon ies u po n MEA (colour plate 84, C and e) attaining a diameter 01' 60
1I11ll in 14 days at room temperature (20-25C), consisting 01' a th in felt but 01'
a tcx ture looser than upon CA and CYA, con spicuousIy zonate, at first whit c,
(h cn becoming pale yellow; co lon y surface de ep ly la n a te , with thin margin
su rrou n ded by a wide fringe 01' submerged hyph ae , extending 10 mm 01' more
hcyond the aerial growth ; not furrowed, slightly convex; h eavily sporing in
g"l'a y shades ; exudate la cking; odour less pronounced but similar as des cribed
ah oye; n o colou red pigment diffusing into the surrounding agar; colony
rc ve rse (colour plate 84, c) in yellow -orange sh ades; penicilli com m only la r ger
Ib an upon CA and CYA but essen tially similar in pattern .
The sp ecies description is centered on strain CBS 340. 48 = C M I 40,238 =
A'I'CC 10,455 = NRRL 2,0 16 = QM 6 ,865 = IJFM 3863, isolated by K .B . Raper
rorn Ni caragua soil in 1945, a nd n umerous other strains isolated by us in
Spain .
T his species is probably the most abundant Penicillium found in soi ls, being
cosmopolitan in origin o It is al so us ually isola ted from decaying vegetation
undergoing th e final stages 01' de composition.

B5. Penicillium daleae Zaleski: fig. 85, colour plate 85, and SEM plate 15,
(top : left) .
Penici/lium daleae Zaleski - Bul!. Acad . Polonaise Sci. Math . Nat. Sr. B, pp. 495496. 1927. See also Ch. Thom , 'The Penicillia' , pp. 360-361. 1930; also Raper and
Thorn, 'A Manual of the Penicillia ', pp. 296-299. 1949.

Synonymy
Penici/lium krzemieniewskii Zaleski - Bul!. Acad. Polonaise Sci. Math. Nat. Sr.
B, pp. 490-492. 1927.

Zaleski's original diagnosis follows, since this adequately presented the species in
question when first isolated :
'Colonies in neutra l Raulin with 10% gelatin in Petri dishes, growing slowly,
attaining a diameter of 24-26 mm in 12 days, liquefying the gelatin tardily but
completely, velvety or somewhat subfloccose, zonate but indistinctly and in
marginal area, with t he whole central area broadly radiately fu rrowed , w ith centre

JI

....

",..,

ilii1>~f~ua;~~ , ~
'l J \~
100,..m

Fig. 85. Camera lucida

drawings of Penicillium
daleae Zaleski, CBS 211.28.
A, Detailed drawings of the
penicilli. B, Mature conidia .
C, Habit sketches.

somewhat depressed and showing few colourless drops of exudate; white margin

2-3 mm wide; conidial areas at first developing blue-green shades (Klincksiek and
Valette's Code des Couleurs, Paris, 1908, No. 371, 372), becoming dark yellowgreen (Klincksiek & Valette's Code No. 273), and later dark orange-brown such as
(168, 164, 198, 139); colony reverse at first in orange yellow shades such as (171,
166, 157, to 133, 138), later becoming red-orange (84, 88, 92, 97); odour lacking or
faint; conidiophores 10-200 or 300 Jlm by 2-2.5 Jlm, with apices more or less
enlarged in a vesicle, commonly unbranched, occasionally with short branches,
flexuous, varying greatly in length, erect or ascending; penicilli mostly 10-20 .tm,
less frequently 25-30 Jlm or 40 Jlm long, with walls smooth; metulae 8, 10 Jlm-20 or
24 Jlm in length by 2.5-3.0 Jlm in diameter, in groups of 2 or 3, commonly unequal in
length and irregularly arranged, with apices commonly inflated; phialides about
9-10 Jlm by 2.5-3.0 Jlm, commonly in verticils of 3, 5-10 or 12 elernents, sornetimes occurring singly; conidia 2.5-4.0 Jlm by 2.5-3.0 Jlm, varying considerably in
size, coarsely echinulate, ovate elongated or subglobose .
Habitat: Strain isolated from pine forest soil near Poznan, Poland.'

The type st ru in 01' Penicillium daleae Za lcski exa m incd 1'01' th c pr cscnt study,
scnt by th c C en tra albureau voor Schimmclculturcs , Baa rn , Thc Nctherlands,
as stra in c ns 211 .28 , no longer produces colonies confo rm ing with Zaleskis
(1 927) dcscription , nor does it produ ce well -developcd penicilli on any su bstra tu rn tested . The conidia , how ev er, are entirely cha rac teristic of the species .
O u r not es follow :
Colonies upon CA (colour plat e 85, A and a) attaining a di amet er 01' 43 mm
in 14 da ys at room temperature , con sisting 01' a e1ose-t extured ba sal felt ,
uppea ring velvety 01' lanate owing to th e thin overgrowth 01' close ly interwoven
ac rial hyphae and ropes 01' hyphae ; ra d iately furrow ed , alm ost azonate, with
marginal are a becoming slightly gray colou re d with th e sparse devel opment 01'
ro nid ial area s; exuda te a nd odour lacking; no coloured pigm ent di ffusin g into
lil e su rro u nd ing agar ; colony reverse (colou r plate 85, a) colourless to pale
rrca rn ; penicilli very sim ple , consisting 01' single phialides 01' very small el ust er s
of phi alides (fig. 85, A) , 8-12 p,m by 2.0-2 .5 tun , te rm inating in an a bru ptly
narrowed neck; structures suggesting true peni cilli a re rare ; co nid ia are
cllip tical to su b -globo se, 3.0 - 3 .5 p,m by 2.5 -3. 0 p,m , coarse ly roughen ed , with
rou ghness a rrange d in spirally bands 01' bars (fig . 85, B, a nd SEM plat e 15,
rop : left) ; conid ial chains loosel y parallel (fig . 85, C) .
Colonies upon CYA (colour plate 85 B and b) attaining a di amet er 01' 50 mm
in 14 days at room temperature (20 -25C) , duplicating in texture a nd colony
a ppea ra nce those gro wn upon CA , but spo ring better in m arginal area s in gray
xha des (Sg uy , PI. XXXV , 522) ; thin , eve n margin su rrou nde d by a frin ge 01'
subrnerge d hyphae exte nd ing 2- 3 mm beyond th e ae rial gro wth; am be rr oloure d pigment diffusin g into th e surro u ndin g agar; exudate and odo u r
lacking; colony surface ove rlaid by a wh ite network 01' closely woven ae rial
hyph ae obsc u rin g greatly th e con id ial area s; lending a lan at e appearance to th e
colonies; colony revers e (colour pIat e 85, b) in brownish to orange shades;
pcnicilli esse ntially as described aboye.
Colon ies upon M EA (colour pIat e 85, C , a nd c) atta ining a di am eter al' 45
mm in 14 days at room temperature (20- 25 C ), sim ila r in texture a s described
ab oye , but not furrowed; heavil y sporing in brownish gr ay shade s (Sguy , PI.
XVI , 23 1); velv et y to lanat e , with thin m argin surrounded by a frin ge 01'
su bmerged hyphae exte nd ing 2- 3 mm beyond th e aeri al grow th ; pen icilli
cssentially a s described , mostly appearing sim ple; ex udate and odo u r lacking;
brown-coloured pigm ent di ffusing into th e su rro u nd ing agar; colony revers e
(colou r plat e 85, e) in ora nge shade s.
The species description is ce nte re d upon th e type strain CBS 211.28 = CMI
34.9 10 = ATcc l 0,435 = IF06,087 = NRRL2 ,025 = IJFM 5117 ,isolatedbyK.
Zaleski from a pine forest soil near Poznan, Poland in 1927 . (C olour pIat es 228
and 229 present two representative strains 01' P. daleae, isolat ed from an andosol
in the province 01' Navarra , Spain , 1976, by DI'. A .T. Martnez.)

HG. Penicilliu m simplicissim um (O udcmans) T ho m: lig. HG, colou r plute

86 , and SE M plat e 15, (top: right ).

lw.J.W.W l

O
v,

~":s .:?

01O)Jm

O O O
O O BO O

r.:

O O \\__-,

1"j -. )

';..~
;;" ..... (((
,\)

000
O. O. O'

'L

'<~~,...

~\'"~:J) r~\~ ~,;.. -:-.

1
,

'------J

100)Jm

(~f1:~'

R.

Fig. 86. Camera lucida

drawings of Penicillium
simplicissimum (Oudemans)
Thom, CBS 372.48. A,
Detailed drawings of the
penicilli . B, Mature conidia .
C, Habit sketches.

Penicilliumsimplicissimum (Oudemansl Thom. - "The Penicillia', pp . 335-336.1930.

See also Raper and Thom, 'A Manual of the Penicillia', pp. 304-305. 1949.

Synonym
Spicaria simplicissima Oudemans - Nederl. Kruidk. Arch. Ser. 3, 2: 763. 1903.
See also C.N. Jensen, Bull. Cornell Univ . Agric. Exp. Sta. 315: 493. 1912.

C oloni es upon C A (colour plat e 86, A and a) attaini ng a diameter of 45 mm

in 14 days at roo m temperature (2 -25C), cons isting of a tou gh , closetex ture d basal felt , appe aring deepl y velvety, deepl y radiat ely furrowed , with
central areas somewhat ra ised in a craterifor m pattern , azo nate or slightly
azo nate in sorne strains ; thi n , even m argin surrounded by a fringe of yellow
submerged hyphae ex tending ab ou t 4 m m beyond the ae rial growth; heavil y
sporing thro ugho ut in blue-green shades, with th e blue colour dominat in g

(SI: gUY, PI. XXXIII , 4112 , 4IU); exuda re limit cd lo abundaut , cry sta l clca r ,
rulou rlcs s; odo u r lackiu g ; no co lo urcc l pigm ent dilfusin g inr o th c su rro u nd ing
:I~a r ; colon y reve rso (colour plat o 1I6, a) in bul' shadc s with margin in ycllowol'ang"e colours ; pcnicilli typi call y asy rn mct r ic (fig . 86 , A) and divaricate ,
usually consisting 01' terminal clus te rs al' 2 -4 divergent metulae bearing
vc rt icils al' phialides (appcarin g esse ntially simple) , borne upon ascending
xtipos arisin g m ostly from th e substratum, al' from veg etative aerial hyphae
Ilpon sho rte r stipes; con id iophores with walls fin ely roughened , varying greatly
iu dimensions , rangin g from 200 to 800 11m al' more in length b y 2.5 -4 .0 11m in
d i.unc te r in larger structures, to very short, usuall y less th an 50 11m by 2.5 11m,
whc n arising as lat er al branch es frorn ve getative hyphae; metulae va riable,
Illoslly 12- 20 11m b y 2. 5- 3 .5 11m ; phialides in whorl s al' 3 - 10 elements in the
ve rticil , mostly 10-1 3 11m by 3 .0 -3.5 11m , abruptly tap ering to a consp icu ou s,
n.urowed , lon g neck ; con id ia a t fir st ellip tical , usuall y remaining so but
Irc que ntly appearing subglobose, mostly 3.0-4.0 11m in long axis , with walls
Iin cly roughened wh en observed under the light mi cr oscope (fig . 86 , B) but
with wall ornamentati on a p pear ing cere b rifor m when observe d under th e
SEM (plate 15 , top : right) ; con id ia l chains in long loosel y parallel colu m ns (li g .

11<;, C) .

Colon ies upon C YA (colour plate 86, B a n d b) a ttain ing a di amet er a l' 55
in 14 d ays at room temperature (20 -25 C) , sim ila r in texture to those
g-rown upon CA, but more d eeply su lca ted with numerou s radial furrow s, a nd
hcavily sporin g in blue-gr een to artemisia gree n shades, with the gr een colou r
prcd ominant (Sgu y , PI. XXVII , 403 , 40 4) ; thin, even margin surrounded by
a wide frin ge al' su b me rge d h yph ae exte nd ing about 4 mm beyond th e ae rial
g-rowth; ex u date limited , as crystal c1ear , uncoloured droplets ; odour lacking;
110 coloured pigment diffu sin g into th e su r rou n d ing agar; colony reverse
(colou r pl at e 86, b) in buff to cin namo n shades; penicilli essen tially as
dcscribed a bo ye, b u t with con id ia l chains tangled.
Colo n ies upon MEA (col our pIat e 86, C a n d c) attaining a diamet er al' 60
mm in 14 d a ys at ro om temperature (20- 25 C) , cons istin g a l' a looser textured
basal felt , plane , not furrowed , with thin m argin surrounded by a frin ge a l'
sub me rge d hyphae ext ending ab ou t 3 - 4 mm beyond th e ae r ial gro wth; heavily
spor ing throu gh ou t in blue- green shades (Sgu y, PI. XXXIII , 488); odour a nd
exudare lacking ; no coloured pigment di ffusing into th e su rro un d ing agar;
colony reverse (colour plat e 86 , c) colou rless with a green ca st showing
through ; penicilli essen tially as d escribed a boye, but with th e con id iopho re s
a rising primaril y from th e su bs tratu m, a nd with the conidial chains tangled.
The sp ecies description is ce n te red upon strain C B S 372.48 = C M I 39, 8 16 =
ATCC 10,495 = NRRL 902 = IF O 5,762 = QM 1,0 39 = IJFM 704 8 , isolat ed in 1940
Irom a flannel b a g by Mrs E .M . Laughton , Cape Provin ce, South Africa, and
add ition al cu ltures isolated from soil in Spain by th e author al' th e current
stu dy.
IIIIn

B7 . Penicillium ochro -chloron Biou rge : fig. 117 , colour plate B7 , and SE M
platc 15 (centre: len ).

A
~~:.
A

~
. 7
" A'

'"

,,:.,..,. . .';.~, v
::,..
.,

" "

0..00 V
O O_n o
o

B(JcrO

O ()

O ()

0 0

Fig. 87. Camera lucida

drawings of Penicillium
ochro-chloron Biourge, CBS
357.48. A , Detailed drawings
of the penicilli. B, Mature
conidia . C, Habit sketches.

Penicillium ochro-chloron Biourge - Monographie, La Cellule. 33: 269-270. 1923.

See also Ch. Thom, 'The Penicillia', pp. 363-364. 1930; also Raper and Thom , 'A
Manual of the Penicillia' , pp , 305-308. 1949.

Synonymies
Penicillium cuprophilum Sato - J . Agric. Chem. Soco Jpn. 15: 359-369. 1939; and
Bull . Agric. Chem. Soco Jpn . 15: 77. 1939.
Penicillium biforme Thom varo vitriolum Sato - J. Agr ic. Chem. Soc oJpn . 15:
359-369. 1939; and Bul!. Agric. Chem. Soco Jpn . 15: 76-77. 1939.

Co lonies upon CA (colour plate 87, A a nd a) growing rather rapidly,

attaini ng a dia meter of 55 mm in 14 days at room temperature (2-25C),
consisting of a close-textured basal felt, wit h surface somewhat lanat e or

cou ony owing lo thc closely int erw ovcn , fine , veget ativo, aerial hyphae
overlying th e co lony su rfac e , partially obs curing the otherwise heavil y sporing
a rcas with radial tufts uf white f1occose hyphae ; colony surface radially
furrowed and raised in central arca, azonate 0 1' in some cases be coming zonate
with the development 01' conidial structures ; thin, even margin surrounded by
a frin ge 01' submerged hyphae exte nding about 5 mm beyond the aerial growth;
hcavily sporing throughout in deep olive gre en shades (Sguy, PI. XXIX,
11 ~ 6) ; exudat e abundantly produced in some stra in s, not in others; odour
lacking ; no coloured pigm ent diffusing into the surrounding ag ar; colony
rcverse (colour plate 87, a) in buff to flesh shade s, with margin developing
or an ge colou rs; penicilli usually more con centrated in m argin al areas , variable
in size and in com plexity, strongly divaricate (fig. 87, A); conidiophores
va riable in length, 01'ten a risin g as branch es from aerial hyphae a nd rarely
cxceeding 100 p.m in length by 2.0-2 .5 p.m in di amet er, less com monly arising
lrorn th e substratum , up to 300 - 500 p.m in length by 2.0-2 .5 01' 3. 0 p.m in
c1iamet er , with walls conspicu ou sly warty ; penicilli appearing variously sim ple,
irr egul arly ramified, 01' occasionally with fairly well defined ve rtic ils 0 1' metulae
IIr br an ches, variabl e in dimensions, ran ging from 10 to 20 p.m by 2 .5 - 3 .0 usn ,
with walls com mo nly warty; phialides in whorls 01' 3-1 O elements in th e
vcrticil, divergent , mo stly 10-12 p.m by 2.5- 3.0 uu, abruptly tapering to a
cons picu ously long, narrowed neck; conidia elliptical with on e end com mo nly
upicula te, usuall y 3.0 - 4 .0 p.m by 2.0- 3.0 p.m in diamet er, with wall s sm ooth
or deli cately roughen ed (fig . 87, B, a nd SEM plate 15, centre: left) .
Co lonies upon CYA (colour plate 87, B and b) att a ining a diameter 0 1' 60
mm in 14 days at room temperature (20 - 25 C ) , co nsisting 0 1' a sim ila rly
(cxtured basal feIt as described aboye , deepl y velvety to lanate, radiately
sulcated with numerous fu rro ws; eve n margin surrounded by a wid e fringe
cxtend ing about 5 mm beyond th e ae rial growth ; abu ndantly spo ring
through out in gray-green shad es (Sguy, PI. XXIX , 427) , partially obs cured ,
cspecially in m arginal areas by th e floccose overgrowth of closely interw oven
ae rial hyphae ; exudate abundantly produced as crys tal-clear, uncoloured
drops; odo ur lacking ; no coloured pigment diffusin g into th e surrounding
agar; colony revers e (colour plate 87, b) in dark cinnamo n shad es with margin
tinted in orange colours; penicilli essentia lly as described ab oye , with conidial
cha ins loosely parallel to tangled (fig . 87, C ) .
Colonies upon M EA (colour plat e 87 , C and e) attaining a di amet er of 55
m m in 14 days at room temperature (20 - 25C) , con sisting 01' a looser textured
basal feIt , velvety, plane, not furrowed , with thin , eve n margin su rro u nde d by
a frin ge of sub me rge d hyphae exte nd ing abo u t 1- 2 mm bcyond th e ae rial
growth; heavil y sporing throughout in gray-green shades (Sgu y , PI. XXIX,
429); colony surface ove rla id by a thin netw ork of aeri al hyphae and thin ropes
of hyphae; exu date and odour lacking; no coloured pigm ent diffu sing into th e

surround in g aga r; colony reverso (colour plat e 87, e) in pa le orange shades :

penicilli cssent ially as d escribed a bo ye but with co nid iophore wall s mor"
coa rsely rou ghc ncd .
The spec ies descrip tion is ccnte red up o n strain C HS 357 .4B ~ CMI 39, 806
ATCC 10540 = NRRL 926 = QM 7,604 = IJ FM 7049 , isolated by Professor
H otson , U n ivers ity of W ashington , Seattl e, W ashington , USA, fro rn a 2 %
solution of coppe r sulphate.
Penicillium ochro-chloron has repeat edl y been isolat ed fro m cop pe r solutions ,
from tentage and fabr ics tr eat ed with coppe r n aphthan at es 01' other copperbearing mildewcides, suggesting th at we are dealing with a form characterized
by a n unu su al toler ance of this metal.

88 . Penicillium piscarium Westling: fig. 88 , colou r plat e 88, and SEM

pIate 15 , (centre: ri ght).

Uw.J.=.J

10~m
r.,~

~::~~~~

;~~

.-

'~ y

.:
'--------'

100~m

CK.

Fig. 88. Camera lucida

drawings of Penicillium
piscarium Westling, CBS
220.30. A , Detailed drawings
of the penicilli. B, Mature
conidia . e, Habit sketches .

.. ".
Penicillium piscarium Westling - Ark. Bot. 11: 54. 1911. See also Biourge's
Monographie, La Cellule. 33: 190-191. 1923; also Ch. Thom, 'The Penicillia', pp.
487-488. 1930; and Raper and Thom, 'A Manual of the Penicillia', pp. 308-309.
1949.

C olonies upon CA (colour plate 88, A and a) spreading broadly , attainin g a

diameter of 60 mm in 14 days a t room temperature (20- 25 C) , cons isting of a
c1osc-textured ba sal fclt , fairly tou gh , ovc rla id by a loose floccos e ove rg ro wth
01' int erwoven ae rial hyphae , white , radiatcly fur ro wed , usually rai scd in
central area s; thin m argin su rrou nde d by a fringe of su bme rge d hyphac
extend ing abo u t 1- 2 m m be yond th e ae rial growth ; conid ial structu res ve ry
spa rsely produced , not influencin g th e colony ap pearancc ; exudate lacking or
limit ed in amou nt; odo u r lacking ; no colou re d pi gm ent d iffu sin g in to thc
surrou ndi ng agar ; colony reverse (colour plat e 88, a) in cream to pale salmon
pink shades; penicilli extrem cly ir regula r in pattern (fig . 88 , A) with th e
idcn tit y of th e clements diffi cult to det ermine , rangin g fro m simple , throu gh
vert icils of th ree or four unequal b ranch es or m ctulac to very com plex
st ructures with m etul ae and phialides borne at various nodes on branch es of
un eq ual length , m ctul ac produc ing whorl s of few phialides or grow ing out in to
scptate hyphae (fig . 88, A) and us ually producing a single phialides at th e
apcx; conid iopho res smooth wall ed , usu ally arising as sho rt branch es fro m
acria l hyphae , 50 /lm or less in len gth by 2 .5 -3.0 /lm in d ia me te r, or wit h
pcn icilli borne tcrminally on lon g trailing hyphae ; coni d ia globose to
subglobose (fig . 88, B a nd SE M plat e 15 , centre: ri ght ), conspicuo us ly
cchin ulate, m ostly 3.5 -4.0 /lm in d ia metcr; con id ial cha ins sho rt, looscly
par allel to ta ngled .
Colonics upon C YA (colou r plat e 88, B and b ) attaini ng a d iameter of 60
mm in 14 days at roo m te m pe rature (20-25C), duplicating in texture th ose
grown upon C A, but deeply floccose and sporing heavier in brown colours
(Sguy, Pi. X LVII, 701) , colony surface sulcated by n umerous radial furrows;
thin margin su rroundc d by a fringe of su bme rge d hyphae extendi ng abo ut 3-4
mm beyond th e aerial gro wth; exudate limited in amou nt as small d ro ps
cnmes hed in the m ycelial felt ; odo ur lackin g ; no coloured pi gm ent di ffusin g
int o th e surrou nd ing agar; coni d ial areas partiall y obsc u red by an abu ndant
overgrowth of white floccose , loosely wove n aerial hyp hae ; colony reverse
(colou r plat e 88 , b) in pale orange to cinnamon shades ; penicilli some what m or e
reg ula r th an a boye, but with cle me nt n umbers very variable ; b ra nc hes from 12
to 20 /lm by 3. 0 uu; m etulae 8- 15 /lm by 2.5 -3.0 /lm ; phialides 9 - 12 /lm by
2.3-2.5 tui , abru ptly tap ering to a long, narrowed neck; conid ia as described
aboye .
Colonies upon MEA (colour plat e 88, C and c) attaining a d iameter of 40
mm in 14 days at room tempera ture (20-25 C), cons isting of a looser textured

basal fclt , ov c rlaid by a dee ply Iloccosc ove rgrow th 01' interwo ve n aerial, whi tc
hyph ae , len di ng a lanat e a ppcara ncc to the colony surfacc; spo ring in graygree n shades (S guy, PI. XXIX, 428) , bu t with conid ial arcas mostly obs cu red
by th e floccose white overgrowth , exce pt in th e ce ntra l a rea; exudate and odo ur
Iacki ng ; no coloured pi gm en t di ffusi ng int o th e su rraund ing a gar ; colony
reverse in eream sha des (colour plat e 88, e) ; eonid iopho res u su ally longer th a n
on CA, rangin g from 50 to 100 .tm in len gth wh en borne as branehes and with
tr a ilin g hyphae mor e ofte n terminating in pen ieilli ; penicilli as described
aboye .
The spee ies deseription is een tere d up on th e typ e st ra in CB S 362 .48 = CMl
40 ,032 = ATCC 104,926 = NRRL 1,075 = IFO 8, 111 = V KM F-1 ,82 3 = IJFM
7056, and strain C BS 220 .30 = N R RL 2132 (both presu m abIy de seendin~ from
W estlin g' s type), isoIat ed from eod Iiver oiI by W estling in 1911. OnIy on e
add itionaI strain approx imating Penicillium p iscarium was isolat ed by P .B.
M arsh fro m a pieee of u n treated cotto n due k at Be1tsville , M aryIand , USA , as
NR R L 2022 in 1949.

89 . PeniciLlium miczynskii Zaleski : fig. 89 , colou r plat e 89, and SEM

plate 15 (b ot tom: Ieft).

I
O 00

oC)

O
O o

w.ww....J

1O .... m

OB o

00 O 00 o
o 00

0 O

y r i~
l..--...J

100 prn

fR

Fig. 89. Camera lucida

drawings of Penicillium
miczynskii Zaleski, CBS
220.28. A, Detailed drawings
of the penicilli. B, Mature
conidia . e, Habit sketches.

n;c;//;um miczvnskli Zaleski - Bull. Acad . Polonaise Sci. Math. Nat. Sr. B, pp.
482-484. 1927. See also Ch. Thorn , 'The Penicillia', pp. 488-489.1930; also Raper
and Thom, 'A Manual of the Penicillia', pp. 309-312. 1949.

Synonymies
Peniclllium sulfureum Sopp - in Ch. Thom, 'The Penicillia', p. 451 . 1930.
Penictium menqini Duch et Heim - Trav. Crypt. ddis a L. Mangin . Mus. Nat.
Hist. Nat., Paris, pp. 431-458. 1931.

Colonics upon C A (colou r plat e 89, A and a) a ttaini ng a di amet er of 40 mm

in 14 da ys a t room temperature (20 - 25C), consistin g of a thin a nd brittle
basal felt of white to ye llow m ycelium , close textured , az on at e or becoming
zo na te in marginal a rea with the development of co n id ial stru ctu res ; with
rolony su rface ir regularl y rai sed a n d com monly buckl ed a n d wrinkled in a
cc rcbr iform p attern , espec ia lly in the central a nd su bce n tral a reas, ve lve ty in
nppe a rance ; with a thin, even m argin surrounded by a frin ge of su b merge d
hypha e ex te n d ing abo u t 2 mm beyond th e ae rial gro wth; abundantly spo r ing
in marginal a reas in blue- green shades w ith th e blue colour predominant
(Sguy , PI. XXXIII , 483); exu d a te ab u ndan tly produced or lacking ; odour
Iaint or lacking ; p ale amber-coloured pi gment diffusing into th e surrounding
agar; colony reverse in ye llo w-o range shades (colour pIa te 89, a); penicilli
va riable, typicall y asy m metr ic di varicat e , sometimes appearing sim ple ,
usu all y co nsisting of 3 -5 m etulae ofte n of unequal length a nd som etimes
arising a t diffe rent nod es , with phialides a nd m etulae occasionally borne a t th e
sa rne level ; with conid iop hore s arisin g primarily from th e b as al felt , less
ro m mon ly from ae rial hyphae , up to 300 - 400 JJ-m b y 1.8 -2 .5 JJ-m , with walls
smo oth (fig . 89 , A) ; m etulae 8 -1 5 JJ-m by 1. 5- 2.5 JJ-m ; phialides in com pact
rlu sters of 3 - 8 ele ments in th e ve rticil , m ostl y 7-10 JJ-m by 2.5 - 3.0 JJ-m,
terminatin g to a narrowed , tapering n eck ; con id ia ellip tical, com paratively
srnall, 2.0 -2 .5 JJ-m by 1.5- 2 .0 JJ-m in di amet er , with wall s thin and smooth or
nearly so (fig . 89, B, and SEM pl a te 15 , b ottom : left); con id ial chains tangled
(Iig. 89 , C).
C olo n ies upon C YA (colour pl a te 89 , B a n d b ) a tt ain ing a diamet er of 45
m m in 14 days at room temperature (20- 25 C) , sim ila r in texture to th ose
grown upon CA, but m ore regular in p attern , with numerous radial furrows
a nd ce ntre raised in a cra te riform pattern ; remaining persistently white owing
to the thin , d ense ove rgrow th of closely wov en , ae rial, white hyphae ; slightly
spo ring in the same shad es as aboye, but n ot influencing th e colo ny
a ppe a rance; odour faint , indefinite ; ex udate lacking; yellow-coloured pigment
diffusing into th e surro u n d in g ag a r; colo ny re verse (colour pIat e 89 , b) in
yellow-oran ge sh ade s; penicilli essen tially as described aboy e .
C olonies upon MEA (colour pl a te 89 , C an d e) att ain ing a diamet er of 50

mm in 14 d ay s a l 1'00 m tcm pcratu re (20 -2:>0) , co ns isiing 0 1' a dceply velv ety
felt , with thin rnar gin su rro u nde d by a wid c Irin gc 01' subme rgc d hyph ae
exte nd ing abo u t 5 mm beyond the ae rial gro wt h; colony surfacc plane
somewha t radi ately fur rowed in the centre, a ppear ing slightly floccosc owing to
th e loose-t extured floccose ove rgrowth; heavily sporing throu ghout in bluegreen shade s (Sguy, PI. X X XIII , 482) ; ex udate and odo ur lacking; no
colou red pigment diffusin g into th e su rro u nd ing agar; colony rever se (co lou r
plat e 89, c) in cream shades; penicilli esse nt ially as described aboye, with
conid ial cha ins loosely parallel to tan gled .
T he species description is centered up on th e typ e stra in C BS 220 .28 = CMI
40 ,0 30 = ATCC 10470 = NRRL 1,077 = IFa 7,730 = Q M 1,957 = IJ FM 5125,
isolat ed by K . Zaleski , Pol and , 1927 , from coni fer forest soil.
The presen ce 01' aggregates 01' inflated , heavy-walled cells in strai ns,
othe rw ise closely approx imating Penicillium miczynskii, suggests a relatio ns hip 01'
thi s spec ies to P. soppii in th e P. raistrickii seri es, hen ce its inclu sion in th e key to
th e Penicillium raistrickii series.

90. Penicillium asturtanum R amrez el Martnez : fig. 90, cala u r plat e 90 ,

and SEM plat e 15, (bottam: right).

'--'\

.~:

,
I

,( e.
j

;;:~?: l1
lo o
o

fR..
~

100J.lm

Fig. 90. Camera lucida

drawings of Penicillium
asturianum Ramrez et
Martnez, ATee 42,226. A,
Detailed drawings of the
penicilli. B, Mature con idia .
e, Habit sketches .

nicillium esturianum Ramlrez et Martlnez - Mycopathologia. 74: 34-49, 1981.

C olon ics upon C A (colour plat c 90 , A and a) sprcading broadly, a tt ain ing a
di.un ct cr 01' 45 - 50 mm in 14 days a t room tcmpcrature (20- 25 C), consisting
IIr a close-tcxtu re d basal felt, overlaid by a closely interwoven network 01' fin e
hyphac lending a deeply velvety to lanate appearance to the colony surface;
gro win g margin raised and cen tre plane depressed, developing deep bluegl'ccn shades (Sguy, PI. XXX, 446 , 447) in heavily sporing areas , while
poorly sporing areas remain persistentIy white or tinted in pale yellow colours ;
exuda re limited to a few , uncoloured drops ; odour lacking; no coloured
pigment diffusing into th e surrounding agar; colony rev erse (colour plate 90, a)
dcv cloping cream to pale salmon pink colours ; penicilli typically as ymmetric
.md divaricate (fig. 90 , A), with conidial cha in s in long parallel rows, forming
more or less well-defined compact columns, borne on asc ending conidiophores
IIp (o 300 11m in length by 2.7 -3.0 11m in diamater, with walls smooth, 01' on
xhort branches arising from aerial hyphae, commonly 10-50 11m by 2.5-3 .0
/.tIIl , varying in complexity from simple verticils 01' phialides (appearing then
monoverticillate) to verticils consisting 01' both metulae and phialides, 01'
vcrticils 01' metulae 01' unequallength ; metulae usually in small verticils 01' 2-3
clcme nt s in the verticil, usually 17-27 11m by 2.0-4.0 11m, with apices more or
lcss vesiculose ; phialides comparatively large , diverging, in verticils 01' 3-8
clernents in the verticil, en la rged at the base th en tapering abruptIy to a fairly
long, narrowed neck, cha racter istic 01' the P. janthinellum series, mostIy 12-15
11m by 3.0-4.0 11m; conidia strongly elliptical initially, and remaining so but
with both ends apiculate at maturity, with walls delicately roughened, with
roughness arranged in spirally bands (fig . 90 , B, and SEM plate 15, bottom :
right), commonly 5 .0-6 .5 11m in long axis by 2.5-3 .5 11m in short axis;
con id ial chains in more 01' less long, compact columns (fig . 90, C) .
Colonies upon CYA (colour plate 90 , B and b) spreading less broadly than
aboye , a tt a in in g a diameter 01' 43 mm in 14 days at room ternperature
(20-25 C), deeply velvety , con sistin g 01' a cIose-textured basal felt overlaid by
a thin network 01' dosely interwoven aerial hyphae , deeply radiately furrowed
with centre depressed in a crateriform pattern; even margin with a fringe 01'
sub merged hyphae extending about 1 mm beyond the aerial growth; heavily
sporing in deep blue-green shades (Sguy , PI. XXX , 446), with surface
unevenly tufted, lending a granular appearance to the colony ; exudate and
odour la cking; reverse in cinnamon shades (colour pIate 90, b) ; ambercoloured pigment diffusing into the surrounding agar; penicilli essentially as
described aboye .
Colonies on MEA (colour plate 90, C and c) growing rather restrictedly,
attaining a diameter 01' 35 mm in 14 days at room ternperature (20-25C),
consistin g 01' a loose textured felt, typically velvety, thin; heavier sporing

th rough out in de cp blu e-grccn shadcs (S guy , PI. XXX, 4106); vcry thi n
margin surroundcd by a wid c frin ge 01' sub mc rged hyph nc cxtc nd ing abo ut 1mm beyond th e ae rial grow th; exuda tc and odo ur lackin g ; pale a m bcreoloured pigme n t diffu sin g in to the surrounding agar; colony rcverse (colou r
plat e 90, e) in pale yellow to ora nge sha des ; peni eilli cssc ntially as described
aboye , but with eonidiopho res mostly arising prim aril y from the su bstra tum ;
eonid ial ehains in fairly lon g eompae t eolu m ns .
T he spe eies deseription is based on th e typ e stra in IJ F M 3871 = ATCC 42,2 26,
isolated by th e au thor from th e a ir 01' M adrid , 1977 . It is at present kn own only
from the typ e strain .

91. Penicillium aragonense R amrez el Martnez : fig. 91 , ca la u r plate 91 ,

and SEM plate 16, (ta p : left) .

i'
11. "r
~

100

.'

Fig. 91. Camera lucida

drawings of Penicillium
aragonense Ramrez et
Martnez, ATCC 42,228. A,
Detailed drawings of the
penicilli. B, Mature conidia.
C, Habit sketches.

Penicillium aragonense Ramrez et Martnez - Mycopathologia. 74 : 35-49, 1981.

C oloni es upon C A (eolour plat e 91 , A and a) spre ading mod erately ,

attaining a di ameter 01' 45 mm in 14 days at room temperature (20 -2 5C) ,

velvcty , co nsisting 01' a close-textu rcd basal fclt , com pa ra tivcly thin, with
white , cvcn rnargin surrounded by a Irin ge 01' dccply sub mc rged hyph ac
cxtending 2 -3 mm beyond th e acrial growth; colony su rface sulca tcd hy
shullow radial furrows, moderat ely raiscd towards th e cent re ; sporing
nbundantly throughout in pal e gray-green shades (Sguy , PI. XXIX, 429) lo
pule brownish colours (Sguy , PI. XXIX , 427); exu date a nd od our lacking;
nmbc r-colou red pigment diffusing into the surrounding agar; colony rcvcrsc
(co lou r pIate 91, a) in cream to pea ch shades; penicilli , both strictl y sim ple and
1I1so typically asymmetric and divaricate, with con id iophore s arising primarily
lrom th e substratum and from trailing vegetative hyphae (fig. 91, A), smooth
walled , usually short , less than 100 Lm in length by 2.5- 3 .5 Lm in diameter,
with apices enlarged in a vesicle up to 5 Lm in diameter ; metulae in verticils 01'
" - 3 e1em ents 01' unequal len gth in th e verticil , mostly 18-45 Lm by 2.5- 3 .0
. /un ; phialides comparatively large , cylindrical , abruptIy tapered to a lon g,
narrowed neck, mostly 13-18 Lm by 2.8-4.0 tui, consisting ofwhorls 01' 3 - 8
cleme nts in the verticil, divergent; conidia usually elliptical to subglobo se ,
com paratively large, with walls smooth 0 1' nearly so (fig . 91 , B, and SEM plate
1(>, top: left) , mo stly 4.0-6.0 Lm by 3.2-5 .0 Lm in diameter, forming chains
with conspicuou s connec tives betw een th em ; con idial cha ins at first div ergent,
hecoming tangled in age (fig . 9 1, C) .
Colonies upon CYA (colour plate 91 , B and b) spread in g broadl y, attaining
a diameter 01' 55 mm in 14 days at room temperature (20-25C) , consisting 01'
a close-textu red bas al feIt, velv ety in appearance , deepl y radiatel y furrowed
with centre irregularly folded and raised ; white margin plane, with a fringe 01'
submerged hyphae extend ing about 1 mm beyond th e ae rial growth ; heavil y
sporing in blue-green shades (Sgu y, PI. XXX, 443); colony surface overla id
by a thin, close -textured network 01' fine , aeri al , white hyphae , partially
obscuring, especiall y in central and subcentral areas , th e conidial structu re s;
exudate and odour lacking; arnber-colou red pigment diffu sing into the
su rrou ndin g a ga r ; colony revers e (colour piate 91, b) in cinnamon to
ochrace ou s shades ; penicilli essentially as described aboye.
Colonies upon MEA (colour plat e 91 , C and e) attaining a diameter 01' 50
mm in 14 days at room temperature , quite sim ila r in texture to th ose gr own on
CA, consistin g 01' a thin , close-textured basal felt , showing some zon ation ; thin
margin su rrou nde d by a wid e fringe 01' sub me rge d hyphae , extend ing more
th an 10 mm b eyond th e aerial growth; colony surface deepl y radiately
furrowed , slightI y raised towards the centre; sporing abundantly in light bluegreen shades (Sguy , PI. XXX , 444) to pale brownish colou rs (Seguy , XLVII ,
694), depending on th e rip eness 01' con idia ; exu date a nd odo u r lacking ; colony
reverse in crea m to or ange shades (colour plate 91 , e); pale am be r-co lou red
pigm ent diffusing into the su rrou nding ag ar; penicilli esse ntially as described
aboye.

T he spccies descripti on is ce n tc rc d upon th e type slrain 1.lIIM !'H)72 - .vr cc

42 ,228, isolat ed by th e a ut ho r (C.R.) lrom th e a ir 01' Madrid , Sp a in , in 1978.
It is actually known only Irom th e typ e strain.

Penicillium godlewskii series ser.

DOV .

Coloniae plus minus restrictae crescunt. velutinae, colore virido, griseo-virido, vel
coeruleo-virido ; penicilli divaricati, cum catenis conidiarum diversis, plus minus
compactis columnis formantur, cum phialidibus sine conspicuo collulo; conidia
exigua, non plus ultra 3 ]Lm in diametro; reverso fulgido coloratur.
Typus : Penicillium godlewskii Zaleski.

C olonies showing sorne sha des al' gr een , gray-gree n, 0 1' blue-gr een with th e
rip ening al' con id ia. Penicilli with the di varicat e charac te r well marked , with
conid ial cha ins divergent , usu ally tendin g to form m or e 01' less com pact
colu m ns . C olonies gro wing m ore 0 1' less restrictedl y , a t lea st on sorne
substrata . M etulae usually aggre ga ted into true ve rticils; phialides not
abruptly tapered to a long , narrow ed neck ; conid ia com pa ratively small,
u sually not excee d ing 3 p.rn in di amet er. C olon y reverse brightl y colou re d.

K ey to species

Paga

1. Colonies upon malt extract agar growing restrictedly, not exeeeding 30

mm in diameter in 14 days .
A. Colony diameter upon malt extraet agar not exeeeding 25 mm in 14
days.
1. Colonies predominantly white, funieulose.
P. godlewskii Zaleski .... 259
2. Colonies developing very deep blue-green eolours upon malt
extraet agar with the ripening of eonidia, typieally velvety.
a. Colonies upon malt extraet agar produeing orange coloured
pigment diffusing into the surrounding agar.
P. gorlenkoanum Baghdadi . . . . 269
b. Colonies upon malt extraet agar not produeing sueh pigmento
P. botryosum Batista et Maia . ... 261
B. Colonies upon malt extraet agar attaining a diameter of 30 mm in 14
days.
1. Conidiophore walls smooth .
a. Conidia smooth or nearly so.
P. eben-bitarianum Baghdadi .. . . 266
b. Conidia eonspicuously spinulose.
P. damascenum Baghdadi ... . 268
2. Conidiophore walls coarsely roughened.
P. baradicum Baghdadi .. . . 264

11. Colonies upon malt extract agar growing more rapidly, attaining a diameter exceeding 40 mm in 14 days.
A . Colonias upon malt extract agar conspicuously lanate.
1. Colony reverse developing bright orange -red shades upon malt
extract agar; similarly coloured pigment diffusing into the surroun ding agar.
P. harmonense Baghdadi .. .. 271
2. Colony reverse not as above.
P. sizovae Baghdadi . . . . 273
B. Colonies upon malt extract agar typically velvety.
1. Conidiophore walls smooth; colony reverse showing deep green
colours upon malt extract agar.
P. citreo-virens Abe ex Ramrez . .. . 279
2. Conidiophore walls conspicuously warty; colony reverse showing
typical strawberry red colour.
a. Colonies usually developi ng mouse gray shades with the ripening
of conidia.
P. novae-caledoniae Smith . . . . 275
b. Colonies remaining persistently white.
P. novae-caledoniae Smith, varo album Ramrez et Martnez ... . 278

T he Penicillium godlewskii newly erected series is admittedly artificia l in

concept and ha s been designed to cover certa in forms which cannot otherwise
be satisfactorily placed in any of Raper and Thom 's recognized series (1949) .
T en spec ies and one variety have been incl uded , namely: Penicillium godlewskii
Zaleski , P. botryosum Batista et Maia, P. baradicum Baghdadi, P. ebenbitarianum Baghdadi, P. damascenum Baghdadi, P. gorlenkoanum Baghdadi, P.
harmonense Baghdadi, P. novae-caledoniae Smith , P. novae-caledoniae Smith , var.
album R am rez et Martnez , and P. citreo-virens Abe ex R a m rez . A close
re1ationship be tween the species included in it is not presumed , and th ey are
conside red together primarily a s a matter of convenience. They do, however,
show certain cha rac te ristics in common , pa rti cularly th e strongly divarica te
character of the pe nicil1i, us ually with con idi al cha ins showing a tendency to
produce fairly divergent , wel1-defined , com pac t colu m ns; the com para tive1y
small, globos e to subglobose con id ia , wit h wa lls ge nerally smooth; and the
rather res tricted growth of its colonies upon malt extract agar.

92 . Penicillium godlewskii Zaleski : fig . 92, co lo ur p late 92 and SEM plate

16 , (top: right) .
Penicillium godlewskii Zaleski - Bull, Acad. Polonaise Sci. Math. Nat . Sr. B, pp .
466-467. 1927. See also Ch. Thom, 'The Penicillia', pp, 365-366.1930; also Raper
and Thom , 'A Manual of the Penicillia', pp. 312-314. 1949.

l.l..J.J..WwJ

10 jrm

00

00
00

O 000

g80oo~

OOOB OO

0 0

0 0

r r~r
L--.J

100 ).1m

fl?.

Fig. 92. Camera lucida

drawings of Penicillium
godlewskii Zaleski, CBS
215.28. A, Detailed drawings
of the penicilli. B, Mature
conidia. e, Habit sketches .

Synonym
Penicillium intricatum Thom - Bull. Bur. Anim. Ind. U.S. Dep. Agric. 118: 75-76.
1910.

Colon ies upon C A (colour pla te 92, A and a ) growing restri ctedly , attaining
a d ia meter of2 0 mm in 14 days at room te m perature (20 -25 C), cons isting of
a thin , to ugh, close -tex tured basal felt , developing usu all y wet , sodde n, mo re
or less bristly to funiculose colonies, characterist ica lly producin g a limited cro p
of con idia, rem a in in g persistently wh ite in sorne strains but in others
developing abu ndant conid ial str uc tu res in m arginal a re as , in ligh t blue
green shades (Sg uy , PI. XXX , 445); with m argin plane an d ce ntral and
subcentral a reas strong ly ra diately fu rrowed, with ce ntre folde d in a
crateriform patte rn ; odour indefini te or lackin g ; ex udate limi ted in amo u nt, as
colou rless d ro plet s enmes hed in the m ycelial felt; no coloured pigment
d iffusing in to the surro u nd ing agar; colon y reverse (colour plat e 92, a)
yellow ish; penicilli varia ble in size and pattern bu t typica lly d ivaricat e (fig. 92,
A); conidiop hores va riable in size and origi n, rangin g from 50 to 300 p.m in

"\)1

lcn gth by 1.9 - 2.4 Jtlll , ar ising eithc r di rcctly lrom th o su bs tratu m 01" as shorl
hrun chcs frorn ae rial hyphac , slighlly cn la rgcd a l th e apcx, with wall s smooth;
pcn icilli cons isti ng of singl e ve rticils 01' phial ides (a p pearing rnon overticilIat e) or
vert icils 01' m ctulae of unequal len gth , usu all y 2 - 4 eIeme nts in th e ve rticil,
nbo ut 8 -1 4 Jtm by 1.5- 2.8 u in , with apices enla rged in a ves icIe up to 3.0 ust:
in d iamc ter; ph ialides bottle shape d, in clus te rs of2-8 eIeme nts in the ve rt icil,
mostly 7- 9 Jtm by 2.3 -3.4 utn , abru ptly tapere d to a ve ry sho rt, narrowed
ncck ; conidia globose to subglobose, sm oo th wall ed wh en observed under th e
light microscope (fig. 92, B), a nd deIicat ely ro ughe ned whe n observed under
(h e SE M (plate 16 , top : right) , u su all y 2.3 -3. 0 Jtm by 1.8 - 2.5 Jtm ; conidial
cha ins stro ngly divergent a nd sho rt (fig . 92 , C).
Colon ies upon C YA (colour plate 92, B and b) growing restri ctedl y,
.utaining a di am et er of abou t 30 mm in 14 days at room temperature
(20- 25 C ) , consis ting of a to u gh, cIose-tex tured ba sal feIt, with m argin al and
subce ntral a reas strong ly suleate in a radial patt ern , a nd central areas
irr egul arl y folded and slightly raised in a craterifor m patt ern ; very poorl y
sporing, remaining persistently wh ite ; od our and exudate lacking ; d iffusin g
pigm en t abse nt ; colony reverse (colour plat e 92, b ) stro ng ly pulling apart th e
underl ying agar a nd deveIoping cinna mo n shades; penicilli esse n tially as
desc ribed abo ve .
Colon ies upon MEA (colour pIate 92 , C a nd c) atta in ing a d iamet er ofabo ut
25 mm in 14 d ays at room te m pe ra ture (20 - 25C), sim ila rly textu red as
abo ye , but with less radial fu rrows ; centre irregularl y fold ed in a crateriform
pattern ; very poorl y sporing, re maini ng persistently white; ex udate a nd odo ur
lacking ; diffusin g pi gm en t abse nt; colony reverse (colour plate 92, c)
deveIopi ng yello wish cinnamo n shades; penicilli essent ially as described aboye.
The spec ies description is cente red up on the typ e strain CBS 215.28 = CM I
40,59 1 = ATcc l0, 449 = NRRL 2,111 = IF07 , 724 = QM 7,566 = IJFM7054 ,
isolat ed by K . Zaleski from pine for est soil , Poland , 1927 and upon Thorrr ' s
notes rep or ted in his Monogr aph ( 1930, p . 366), sin ce th e spec ies is di fficult to
mainta in in culture and does no longer represent ade q uate ly the typ e whe n
first isolated, a t least in cult ural appearance.

93 . Penicillium botryosum Batista el M aia : fig. 93, coIour pIat e 93, a nd

SEM pIat e 16, (centre : Ieft) .

Penicillium botryosum Batista et Maia - An. Soc oBiol. Pernambuco . 15: 157. 1957.
See also M. M. Kulik , 'A Compilation of Descriptions of New Penicillium Species' .
Agric . Handbook No. 351. Agric . Res. Ser. U.S . Dep. Agric. pp. 20-21. 1968.
Author's diagnosis follows:

' Colonies on neutral Czapek's agar characteristically depressed in the centre and

4 \J'"

eR.

vYlir il
100)Jm

Fig. 93. Camera lucida

drawings of Penicillium
botryosum Batista et Maia,
CBS 252.55. A, Detailed
drawings of the penicilli. B,
Mature canidia. C, Habit
sketches .

undulating and flat in margin, conspicuously zonate, floccose in appearance, 1-2

mm deep, consisting of a basal felt overlaid by a loose network of aerial mycelium,
consisting of fine ash-green, white, interwoven aerial hyphae, attaining a diameter
of 4-5 cm in 10-12 days at room temperature (24-26C) ; colony surface radiately
furrowed, not extending to the margin in younger colonies; margin white, 0.5-0.8
mm in diameter; exudate limited , as yellow drops, or lacking; odor strong,
suggesting 'mouldy'; colony reverse in bright yellow to brick-red shades.
Conidiophores arising from submerged hyphae or from aerial hyphae, with walls
smooth or slightly roughened, septate, hyaline or in yellow-green shades,
67.5-137.5 .tm by 2.5-3.5 .tm, showing typically monoverticillate penicilli on
inflated vesicles similar to Aspergil/us sydowii (Bainier et Sartory) Thom et Church,
but biverticillate and polyverticillate penicilli are more abundant.
The vesicular branches with or without inflated apices numbering 2-3 per
conidiophore, ranging from 11 to 13.5 .tm by 4.5-5.5 .tm.
The normal branches, 2 per conidiophore, 20-25 .tm by 2.5-3.5 .tm, bear verticils
of 3-5 clvate. diverging metulae, measuring 10-17.5 .tm by 2.5-3.5 .tm. The
phialides are bottle shaped, 4-10 or more elements in the verticil, 3.5-4.5 .tm by
1.0-2.0 .tm, with a short neck .

Conldla globose, thlck - and smoot h w allod, yollow , 2.2tm In dlameter, separated
or formi ng short chains.
Cleistot hecia and sclerotia lack ing.'
(Translated fr om the Portuguese by the authorl .

Our notes follow:

Colon ies upon C A (colou r plat e 93 , A a nd a) gro wing rathcr restrictedly ,
att aining a di ameter 01' 40 mm in 14 days at room temperature (20-2 5C),
consisting 01' a c1ose-t extured b asal felt , typi call y velvety in appearance , with
i hin , undul ating margin , [lat su rro u nde d by a fringe 01' su b me rge d hyphae
cx tcnd ing abo u t 3- 4 mm beyond th e ae rial gro wth, conspicuo us ly zonate,
sorn ewh at lan at e in centra l area owing to th e thin network ofloosely interl acin g
ac rial hyph ae ov erl ayin g th e colony surface; heavil y spo ring throu gh out in
hlu e-gre en shades with th e blue pred ominant (Sgu y, PI. XXXII , 472 );
exud a re ab undantly produced as crys ta l c1ear , yellowish, large drops; odou r
laintly mouldy 01' lacking; pale a mbe r-coloure d pigm ent diffusin g into th e
surro und ing agar ; colony reverse (colour plate 93, a) colou rless, bu t with an
orange to brick-red cast showing throu gh ; pen icilli sho wing typic ally simple
peni cilli , but com monly showing di varicate conid ial struc tu res cons isting 01'
vcrticils 01' 2 or 3 m etulae with apices enlarge d in a vesic1e up to 4.5 p,m in
di ameter; m etulae , mostly 12-20 p,m by 2.3 - 2.8 p,m; phialides bottle shape d,
mostly 9 .0 -10 p,m by 2.5-4 .0 p,m, terminating in a short , tapered , narrowed
neck (fig. 93, A); conid iopho res smooth wall ed, ari sing di rectly from th e
su bstratu m or from trailing aeri al hyphae, rangin g from 25 to 100 tut: in length
by 1.8 -2 .5 p,m in di amet er ; con id ia comparatively sm all , globose to
su bglobose, 2 .0- 3.0 p,m in di amet er , sm ooth walled or nearly so wh en
obse rved under the light microscope (fig . 93, B) but delicately roughen ed wh en
obse rve d under the SEM (pIate 16, centre : left) ; conid ial chains forming
definitely div ergent colu m ns (fig . 93 , C) .
C olonies upon CYA (colour plate 93, B and b) sp reading broadly , att a in ing
a diameter 01' 55 mm in 14 days at room temperature (20-25C) , cons isting 01'
a c1ose-textured basal felt, typi call y velve ty , with a flat margin , white ,
su rrou nde d by a fringe 01' submerged hyphae ext ending about 4 mm be yond
th e ae rial growth ; colony su rface in centra l a nd su bce ntral areas deep ly
sulcated by numerous radial furrows a nd gr adually rai sed tow ards th e ce ntre,
which is overlaid by a thin network al' loosely interwoven aeri al hyphae ,
lending it a sublanat e appeara nce; heavily sporing through out in deep blue
green shades (Sgu y , XXXIII , 486); exudate abundantly produced , as large
yellowish drops; am be r-coloure d pigment di ffusin g into th e su rrou nd ingaga r ;
odour lacking; colony reverse (colour plat e 93, b) in reddish brown or yellowi sh
brown sha de s; penicilli essentially as described aboye .
Colonies upon MEA (colour plat e 93 , C and e) gr owing restrictedly,

a uain ing a d iumctcr 0 1' 20 IlIIlI in 11' days a t roo m tc m pernturc (20- 25C) ,
cons isting 01' a close-tcxtu rcd basal rei r, dcnsely vclvct y , plano, not u rrowcd,
with a few sca ttered tu fts 01' white sterile loccose hyphae in the centre; heavily
spo ri ng thro ughou t in very dark blu e- green shades (Sguy, PI. XXX, 491);
exudate and odour lacking ; amber-coloured pigment slowly diffusin g into thc
su rro u ndi ng agar; colony reverse (colour plate 93, c) in yellow-orange shades;
peni cilli esse nt ially as des cr ibed aboye, with conid iopho res arising primarily
from the subst ratum.
T he species description is centered u pon the type strain CBS 252.55 = ATce
12,068 = QM 6,946 = IJFM 7059, isolat ed as a culture contam inant in 1954 by
A . Batista and H . Maia, Institute 01' Mycology, U niversity 01' R ecife ,
Pern ambuco , Brazil . It is act ually kn own only from th e type strain .

94. Penicillium baradicum Baghdadi: fig . 94, coIour pIate 94 , a nd SEM

p iate 16 , (centre: r ight) .

.'
'' ..

"

::

' r

~ '.

.:::.:" f to um

-:

;..: Oc?B9J
O 1
-:':

/1

: :.;

A ,. .
"

1. . . .

'::

do ooo

1::

000

00
O

100pm

: ;.
l

-:

.'

-:
-:

ff..

Fig. 94. Camera lucida

drawings of Penicil!ium
baradicum Baghdadi, CBS
416.69. A , Detailed drawings
of the penicilli. B, Mature
conidia . e, Habit sketches,

Penicillium baradicum Baghdadi - Nov. Syst. Niz. Rast. pp. 107-108. 1968.

olonies upon CA (colour plate 94, A and a) auainin g a diamct er 01' 40 mm

in 14 days al room tcmpcraturc (20 -25C), con sisting of a close-tcxturcd basal
felt, ovcrlaid by a loosely woven network of fine floeeose hyphae 1- 2 mm deep,
inllucneing greatly (he eolony appearanee and obscuring the otherwise
abundantly sporulating conidial arcas; radiately wrinkled and raised, with
thin, even margin surrounded by a narrow fringe of submerged hyphae
cxtending about 1 mm beyond the aerial growth; heavily sporing underneath
(he deep floccose network of aerial hyphae in pale blue-green shades (Sguy,
PI. XXX, 439); exudate abundantly produced as large topaz yellow drops;
odour lacking; amber-coloured pigment diffusing into the surrounding agar;
colony reverse (colour plate 94 , a) in cinnamon shades with sectors in reddish
brown colours; penicilli typically divarieate (fig. 94, A) with conidiophores
arising primarily frorn the substratum 600 J-m or more in length by 2.5-3.0
J-m, conspicuously warty, or as branches from trailing aerial hyphae,
consisting of verticils of 2-4 metulae 01' unequallength, mostly 12-20 J-m by
2.8-3.0 J-m, with apices slightly enlarged in a vesicle up to 4.0 J-m in diameter;
phialides bottle shaped , abruptly tapered in a very short narrowed neck,
mostly 9-11 J-m by 2.5-4.0 J-m ; conidia com paratively small, globose to
subglobose, smooth walled when observed under the light microseope (fig . 94,
B) or delicately roughened when observed under the SEM (plate 16, centre :
right), mostly 2.5-3 .5 J-m by 2.2-3.0 J-m in diameter; conidial chains forming
divergent, loosely knit columns (fig. 94, C) .
Colonies upon CYA (eolour plate 94 , B and b) spreading broadly, attaining
a diameter 01' 55 mm in 14 days at room temperature (2-25C), similarly
textured as aboye , with numerous radial furrows ; thin, even margin
surrounded by a fringe 01' submerged hyphae extending about 2 mm beyond
the aerial growth; heavily sporing throughout in grayish brown shades (Sguy,
PI. XXIX, 428) underneath th e deep floccose network 01' fine aerial hyphae
overlaying the colony surface; exudate abundantly produced as large topaz
yellow drops; odour lacking ; amber-coloured pigment diffusing into the surrounding agar; colony reverse (colour plate 94, b) in reddish brown to sienna
shades; penicilli essentially as described aboye .
Colonies upon MEA (colour plate 94, C and c) growing very restrictedl y,
attaining a diameter of 35 mm in 14 days at room temperature (2-25 C),
consistin g 01' a close-textured basal felt , deeply velv ety to sublanate, plane with
few shallow radial furrows; thin , even margin surrounded by a fringe 01'
submerged hyphae extending about 2 mm beyond the aerial growth; colony
surface partially overlaid by a thin network 01' floceose aerial hyphae not
influencing th e colony appearance; heavily sporing throughout in deep bluegray shades (Sguy , PI. XXXIV , 499), with sorne floccosity in colony centre ;
exudate and odour lacking; no coloured pigment diffusing into the
surrounding agar; colony reverse (eolour plate 94, e) yellowish with a brownish

cast showing th rou gh in th e centre ; pcni cilli essc ntiully as dcscribcd , hut with
lon ger con id iophores and with con id ial cha ins formin g well -d efincd di ve rgen r
colum n s.
The species description is centered up on th e type strai n C BS 416 .69 = V K M
F-1,069 = C M I 140 ,336 = IJ FM 7106, isolat ed by V . C h. Baghdadi from Syri an
desert soil near Irna , in th e Damascu s region , Sy ria, M arch , 1964 . lt is known
only from th e type strain.

95. Penicillium eben-bitarianum Baghdadi : fig. 95 , colour plate 95 , and

SEM plate 16, (bottom: left).

1~~~

L-----I

~ 100}Jm

R.

Fig. 95. Camera lucida

drawings of Penicillium
eben-bitarianum Saghdadi,
CSS 415.69. A, Detailed
drawings of the penicilli . S,
Mature canidia . e, Habit
sketches.

Penicil/ium eben-bitarianum Baghdadi - Nov. Syst. Niz. Rast. pp . 106-107. 1968.

Colonies up on C A (co lour pla te 95, A and a) growing restrictedl y , attain ing
a diamet er al' 20 mm in 14 days at room te m pe ratu re (20-25C), con sisting 01'
a to ug h, close -tex tured basal felt, abru ptly rai sed , ove rlaid by a den se ne twork
01' ste rile white hyphae influen cin g greatly the colony a ppearance, lending it a
lanat e as pect; undulating m argin surro unde d by a narrow fringe 01' submerged

hyphac extc nd ing about 1 mm bcyond th c ae rial growth ; colony surf acc
irrcgula rly wrinklcd a nd rais cd; medium sp oring underncath the network 01'
rcri nl hyphae in blue-grc cn shades (Sguy, PI. XXX, 437); exudate and odour
lackin g; pal e amber-coloured pigment diffusing into the surro u nd ing agar;
colony rcv erse (colour plate 95, a) in cinnamon shadcs; pcnicilli typi cally
sim ple (appearing monoverticillate) and strongly divaricatc (fig . 95, A),
cunsisting al' verticils 01' branches and/or metulae al' unequal length, strongly
d ivcrgent , with conidiophores smooth wallcd, septate, arising primarily from
1he su bstra tu rn or from trailing hyphae, ranging from 50 to 300 Lm in lcngth
by 2. 5- 3.0 Lm in diameter, with apices (in simple penicilli) cnlarged in a
vcs icle up to 6 Lm in diameter; branche s up to 100 Lm in length by 2.0 -2.5 Lnl
in diameter; metulae, 30-60 tun by 2.0- 3.0 uu, with apices enlarged in a
vcs icle up to 6 .0 Lm in diamet er; phialides bottle shaped, tapering to a short,
narrow neck, mostly 13-15 Lm by 3. 0 - 4.0 Lm; con id ia globose to su bglobose ,
xmooth walled or nearly so wh en observed under th e light mi croscope (Iig. 95 ,
B) but appearing delicately granular when obs erved under the SE M (plate 16,
hottorn : left) , mostly 2.9-4 .0 ust: by 2.5-4 .0 Lm ; conidial cha ins forming loose
div ergent columns (fig . 95, C).
Colonies upon CYA (col our plate 95, B and b) atta ining a diamet er 01' 30
111m in 14 days a t room temperature , consisting 01' a tou gh, close-tex ture d basal
k it, appearing de eply velvety, radiatly wrinkled a nd rais ed , with undulating
margin entire; at first white, th en be coming coloure d in blue-gr een sh ades
sta rting from the centre (Sguy, PI. XXX , 437 , 438) with th e rip ening al'
conidia ; exu date and odour lacking; pale am ber -coloured pigment diffu sin g
i nto th e surrounding agar; colony revers e (colour piat e 95, b) in pale yellowor ange shades ; penicilli essentia lly as de scribed aboye, with conidiophores
a rising mo stly from trailing hyphae.
Co lon ies upon MEA (colour plat e 95, C and c) attaining a di amet er 01' 30
mm in 14 days a t room tem peratu re (20-25 C) , co nsistin g 01' a looser textured
ba sal felt, velvet y , slightly zonate , plane with a thin margin surrounded by a
fringe 01' submerged hyphae ex tend ing ab out 2 mm be yond the aerial growth ;
slightly rai sed in the centre ; heavily spor ing throughout in deep blue-green
sha des (Sguy , PI. XXX, 438) except in th e centre, which is overlaid by a thin
network 01' floccose aerial hyphae ; odour faint , indefinite ; ex uda te lacking; no
coloure d pigment diffusing into th e surrounding agar; colony reverse (colour
plat e 95, e) in oran ge shade s with a gr een cas t showing through in th e centre;
peni cilli essentially as des cribed aboye but with conidial cha ins forming divergent compact columns.
The species de scription is ce nte re d on the type strain C B S 415.69 = VKM F1075 = CMI 140 ,338 = IJFM 710 5, isolated by V .Ch . Baghdadi from Syrian
desert soil near Irna , in th e D amascu s region , Syria , March, 1964 . It is known
only from th e typ e strain.

96. Penicillium damascenum Baghdadi: lig . 96, colour pla to 96, and SEM
plate 16 , (bott om: rig ht).

w.w.w..u

/'(', O

\1)

.10

\V a jrrn

C\

o Oo~ ():

()~ )'O

00
O

~/

eR .
e

~~ .L..-

100 urn

Fig. 96. Camera lucida

drawings of Penicillium
damascenum Baghdad i, CBS
411.69. A , Detailed drawings
of the penicilli. B, Mature
conid ia. e, Habit sketches.

Penicil!ium damascenum Baghdadi - Nov . Syst. Niz. Rast. pp . 101 -102, 1968.

Colonies u pon CA (colou r plate 96, A a nd a) attaining a d iame ter of 40 mm

in 14 days at room temperature (20 -25 C), co ns isting of a tou gh , closetextu red basal felt, ove rlaid by a closely woven network of ae ria l floccose white
hyphae lend ing a la nat e to granular appearance to th e otherwise ve lvety colony
surface, whic h is sulcated by radial and circular fu rrows, with a thin marg in
plan e surrounded by a fringe of su b me rged hyphae extending a bo ut 1- 2 m m
beyond the aerial growth; wit h ce nt re umbona te ; heavily sporing througho u t
in pale blue-green shades with the blue predom inant (Sguy, PI. X XXI, 458),
especially conspicuous in m a rginal areas but obscured in ot he r zones by th e
network of floccose whi te aerial hyph ae ; odour faint, ' mo uldy'; ex udate
abu ndantly p roduced as yellow drops; pale yellow pigment diffus ing into ihe
surrounding agar; colony reverse (colour plate 96, a) yellow in th e centre and
in redd ish orange shades in su bmarginal a re as, almost co lou rless in the

margin ; pen icilli are e itber str ict ly sim ple (appeurin g m un ov ert icillat e) a m i
01' 2 - 5 m etulac 01' cq u al or un equal
Icn gth, typically di vergent (fig . 96 , A) ; m on overticillate penicilli are usu ally
p rcd ominant in th e margin; co n id iopho re s a r ising eithe r from th e su bs tratu m
a nd frorn trailing hyphae, smooth wall ed in our obse rvat ion (rough walled in
Baghdadi's original d iagnos is) , with a pice s e nla rge d in sim ple struc tures up to
:1. 5 /lm in di amet er, m ostl y 200 -250 /lm b y 1.8- 2.5 /lm when borne fro rn th e
su bstratu m , and shorter, up to 50 /lm in length when borne from aerial
hyphae ; m etulae usu all y 10 -20 /lm by 2.0 -2 .5 tuu , with a pices e nla rged in a
vesicle up to 3. 0 /lm in diamet er; phialides b ottle shape d, a b r up tly tapered to a
sho rt n eck , co m m only 7-10 /lm b y 2. 5 -3 .0 usu , in compact whorls 01' 2 -8
c lerne n ts in th e ve rtici l; coni d ia glob ose , cons p icuous ly sp inu lose (fig . 96, B,
a nd SEM plate 16, bottom: right), 2. 8 -3.5 /lm in diamet er; co n id ia l chains
lorming loo se di vergent columns (fig . 96 , C ) .
Colon ies upon CYA (colou r pl a te 96, B a n d b ) a tt a in ing a di amet er 01' 40
mm in 14 days a t ro om temperature , (20- 25C) , co ns isting 01' a close-textured
basal felt , velve ty in a p pearance, d eeply radiately furro wed a nd rais ed in th e
ce n tral a rea, with a wh ite undulating m argin surrou nde d by a frin ge 01'
sub merged h yphae ex te n d ing a bou t 1 mm beyond th e ae r ia l growth; heav ily
spor ing through out , a t fir st in brownish green colours (Sguy, PI. XLVII ,
702 ), turning to blue- gr een shade s (Sgu y, PI. XXX , 443) with th e r ipen in g 01'
co n id ia; odou r faint , ' mou ldy'; exuda te a b u n dan tly produ ced , as ye lloworan ge dro ps; a m be r pigment di ffu sing in to th e su r ro u n d ing agar; co lony
re ve rse (colou r plate 96 , b) in cin namo n to oran ge sh ades; penicilli essen tially
as de scribed a boye .
Colonies upon M EA (colour plate 96 , C a nd c) a tta in ing a di amet er 01' 25
m m in 14 days a t ro om temperature (20 - 25 C), co ns isti ng 01' a looser textured
felt , velve ty in a ppearance, not furrowed , plane , w ith even , thin m argin su rro u nded by a wide frin ge 01' submerged h yphae extending abo u t 2 mm beyond
th e ae ria l gr owth ; hea vily spo r in g throughout in blue-gray sha des (Sguy, PI.
X X X I, 463 , 464) ; ex u date a nd od our lacking ; amber-coloured pi gment di ffus ing int o the su r rou nd in g a gar; colony reve rse (co lou r plat e 96 , c) ye llow ish
with a green ish cast sho wing th ro ugh; penicilli essen tia lly as described a boye
but with con idiop hores mostly borne primarily from the su bs tra te, a nd sim ple
p enicilli predominantly encou n te re d over th e divaricat e ; di varicat e peni cilli
usu all y a r ising fro m trailing h yphae .
The species de scr ip tion is ce n te re d upon th e type strain C BS 411 .69 = V K M
F-1070 = C M I 140,337 = IJ F M 7104 , isola te d b y V. C h . Ba ghdadi fro m Sy r ia n
d esert soil n ear Irna , in the D amascu s regi on , Syria , March , 1964. It is kn own
on ly from th e type strain .
t ypically diva ricatc , co ns isting uf verticils

97. Penicillium gorlenkoanum Baghdadi : fig. 97 , colour plat e 97 , and SEM

plate 17, (top: left).

08

O O

0 00

SOO
0 0

00
00

e
.8

r
A

=-

'1~ 100~m
L....-....l

l?

Fig. 97. Camera luc ida

drawings of Penicillium
gorlenkoanum Baghdadi ,
CBS 408.69. A, Detailed
drawings of the penicilli. B,
Mature conidia. C, Habit
sketches.

Penicillium gorlenkoanum Baghdadi - Nov. Syst . Niz. Rast. pp. 97-98. 1968.

Colon ies upon CA (colour pla te 97, A and a) growing rather restrictedly ,
attaining a diameter of 25 mm in 14 days at roo m temperature (20 -25 C),
co nsisting of a tough , close -text ured basal felt , ty pically velve ty , radiately
sulcated by shallow furrows and raised towards the centre whi ch is umbonate ;
thin, u ndula tin g margin surrounded by a narrow fringe of submerged hyphae
extending less than 1 mm beyon d the aerial growth; heavily sporing
througho u t in blue-green shades (Sguy, PI. XXXIII , 483 , 484) , especially in
the margin: exudate abundantIy produced as large orange-coloured drops,
obs curing in central and subcentral area s th e con idial structures; odo ur
lacking ; ve ry pale a mber-coloured pigm ent diffu sing slowly into the surround ing
agar; colony reve rse (colour pIate 97, a) strong ly pulling a nd cracking the agar
away, in pale burnt sienna brow n shades (Sguy , PI. X LVII , 694, 695);
pen icilli ty pically diva ricate (fig. 97, A) , consisting of ve rti cils of 2- 3 metula e
of uneq ual len gth fairly divergen t ; coni dio phore s smooth wa lled in our
observation (in Baghdadi' s orig inal descrip tion rou gh wa lled conidiopho res
were also enco untered), mos tIy 150-1 000 .tm in len gth by 2 .5 -3 .5 tut: in

rluunctcr, with apices usu ully cnlargcd in a vcsiclc up to 4. 5 ,.u n in di nmctcr ,

co m mo nly arising' prim arily ro m the subst r. u u m in a d en se st a nd, scp uuc;
mct ulac in fairly regula r vc rt icils 01' 2 - :i clcments in (he verticil, sugges ting the
Hivurficillat a c- syuu netrica in patt crn , usu ally l :i - 20 un by 2.8 -3 .5 11m , with
ip iccs cn la rge cl in a ves icle u p to 6. 0 11m in di a mct er; ph ia lides bottle shaped ,
nhru p tly tapered to a very sho r t nec k , cornrno nly 8- 10 11m , by 2.8 -3.3 11m , in
wh orls 01' 6 - 10 elerne n ts in th e ve rticil; conidia globose to subglobose, with
wu lls delicately ro ughe ne d (fi g. 9 7, B a nd S EM pla te 17, top : left ) , co mmonly
" .5- 3.0 11m in lon g ax is; co nid ia l cha ins for mi ng fairly compact di vergent
rn lu m ns (fi g. 9 7, C .).
C olonies upon C YA (co lou r plat e 97, B and b) grow ing more ra pidly,
nua ini ng a di ameter of 40 m m in 14 d ays at room temper ature (20 -25 C),
s uuila r in textu re to th ose grown upon C A, but deepe r ra d ia tely furrow ed in
su hma rginal a reas and circ ula rly sulcated by shallow wrinkles in th e ce nt re ,
I ypically velve ty; heavil y sporing throu ghout in deep blue-green shades, with
hc blue colo u r pr edom inant (Sguy, PI. XXX , 448 , 447); odo ur lacking;
r-xudate a bu ndan tly produ ced in large orange d rop s; yellow colou re d pi gm ent
dilTusing in to the surro u ndi ng agar; colony reverse (co lour plate 97, b) in
ycllow-orange shades; penicil1i cssc ntial1y as dcscribed aboye .
Colon ies upon M EA (co lour pla te 97, C a nd c) growi ng restrictedl y , atta ining" a di ameter of 20 mm in 14 d ays a t roo m temperature (20 - 25 C ),
onsisting of a close-tex tured ba sal felt , typi cal1y velve ty, not furrowed , plane ,
wi t h centre d epressed in a crate riform pa ttern ; white narrow m argin , irn:gul arl y di ssect ed in a denticulat e pattern ; heavily spo r ing through ou t in deep
hlue-gray shades (Sgu y , PI. X XXV , 514) ; exu date a nd odo u r lacking ; no
ro lou red pi gment d iffusin g into th e su rro u n d ing agar; co lony reve rse (co lour
plate 97, e) in reddish b ro wn shades ; penicil1i as described a boye.
T he species descr ipti on is ce n te re d upon our observations of th e typ e strain
<:BS 408 .69 = V K M F-1 ,079 = C M ! 140 ,339 = !J FM 7102, isolated b y V. C h .
Baghdadi fro m Syri an deser t soil near Nabi-Ba ra de, in th e Damascu s region ,
Syria, O ctob er, 1963. It is kn own only from the typ e spec ies.

98 . P enicillium harmonense Ba ghd adi : fig. 98, colour plate 98, and SEM

plate 17 (top : right ).

Penicillium harmonense Baghdadi - Nov. Syst. Niz. Rast. pp. 102-103. 1968.
Colon ies upon C A (colour pla te 98, A a nd a) gr owing fa irly rapidly , a tta ining a di amet er of 55 mm in 14 d ays a t room temperature (20 -25 C) ,
consisting of a loose- textured ba sal felt , ove rlaid by a deep m ycelial network of
closely in te rwoven floccose ae ria l h yphae lending a lanat e a p peara nce to th e
co lony su rface , 2- 3 m m de ep , co nspicuo us ly zo na te, radiately a nd co nce n tri-

u.w..U.w.J

10 jirn

O O

00

0 0

O BO O

0O0

\lllL
L.--J

100 prn

Fig. 98. Camera lucida

drawings of Penicillium
harmonense Baghdadi , CBS
412.69. A , Detailed drawings
of the penicilli. B, Mature
conidia. e, Habit sketches .

cally sulca te d by shallow furrows; thin , eve n margin surrounde d by a wid c

frin ge of sub me rge d hypha e extend ing abo ut 3 -4 mm beyond th e aerial
growth ; heavily sporing throu ghou t in deep blue-green shades (Sguy , PI.
XXX , 448) , with con id ial areas partially obscure d by th e deep floccosc
network of aerial hyphae ; od our stro ng ' m ouldy ' ; exudate abu nda nt1y
produced to limited, collect ing as bright yellow -coloured drops; deep yellowcoloured p igm en t diffusing into the surrounding agar; colony reverse (colour
piate 98, a) in bright ora nge shades; penicilli typi cally sym metr ic and
div ari cat e, althou gh fairly regular verticils of metulae are not infrequently
encounte red , su ggesting th e Biverticillate- symmetrica in pattern (fig . 98 , A),
with con id iophores , according with our obs ervations , smooth walled (Baghdadi 's original description a nd illustrations show rough-walled con id iophores),
commonl y arising from the substrat e a nd from aerial vegetative hyphae,
200- 300 t.m by 2.0-2.5 tui, enl arged in the apex in a vesicle up to 3.0 t.m in
diameter; m etulae mo stly in regular verticils of 2-4 eleme nts in the cluster,
fairl y equ al in len gth , mostI y 10-12 t.m by 2.3- 3.0 tui, with apices enlarged
in a vesicle up to 4.0 t.m in diameter; phialides bottle shape d, abrupt1y tapered
to a very short neck, ranging from 8 to 10 t.m by 2.8- 3.0 tui, forming whorls
of 5- 8 elements in the ve rtic il; conidia globose to subglobose , appearing

mooth wall ed or nearly so wh en obs crv ed undcr th e light mi croscope (Iig. 98,
B) but delicatc1y roughcncd wh en obs ervcd undcr th c SEM (platc 17, top:
1'Khl), ranging from 3.0 to 4 .5 p.m by 2.3 -3.5 p.m, in diameter; conidial chains
Iunnin g Iairly well-defined divergcnt colum ns (fig. 98, C) .
C olonics u pon CYA (colour plate 98, B and b) attaining a diameter 01' about
"o mm in 14 days at room temperature (20-25 C), consisting 01' a closei cxturcd basal felt, de epIy velvety to lanate , sulcated by numerous radial
lu rrows, gradually raised towards the centre; white, wide undulating margin
surrounded by a fringe ofsubmerged hyphae extending about 2-3 mm beyond
1he acrial growth; heavily sporing throughout in deep blue-green shades
(S(~ g u y , PI. XXX, 447); exudate abundantly produced, collecting as largc,
bri ght yellow-coloured drops ; odour strong, 'mouldy'; bright yellow-colourcd
pigm cnt diffusing into the surrounding agar; colony reverse (colour plate 98,
1 in bright orange shades; penicilli essentially as described aboye.
C olonies upon MEA (colour plate 98, C and c) spreading broadly, attaining
I diameter of 55 mm in 14 days at room temperature (20 -25C), consisting of
a looser textured basal felt overlaid by a loosely woven network of floccose
ic rial hyphae , lending a lanate appearance to the colony surface; wide , thin,
whit e margin surrounded by a fringe of submerged hyphae extending up to 5
111111 01' more beyond the aerial growth; colony surface plane , not furrowed or
very slightly radiately wrinkled; heavily sporing throughout in blue-green
shades (Sguy, PI. XXX, 443 , 444); conidial areas partially obscured by deep
scc iors and tufts of floccose aerial hyphae; exuda te and odour lacking; no
coloured pigment diffusing into th e surrounding agar; colony reverse (colour
plate 98, e) developing red shades; penicilli essentially as described aboye,
commonly arising from conidiophores usually borne upon aerial hyphae.
The species description is centered upon the type strain CBS 412 .69 = VKM
10'-1,078 = CMI 140,340 = IJFM 7103 , isolated from Syrian desert soil near
lrna , in the Damascus region, Syria, by V .Ch. Baghdadi, March, 1964. It is
known only from the type strain .

99. Penicillium sizovae Baghdadi: fig. 99, colour plate 99, and SEM
plate 17, centre: left).
Penicillium sizovae Baghdadi - Nov. Syst. Niz. Rast. pp. 103-104. 1968.

Colonies upon CA (colour plate 99, A and a) attaining a diameter of 40 mm

in 14 days at room temperature (20-25C) , consisting of a close-textured basal
Ielt, deeply velvety in appearance, overlaid by a loosely woven network of floccose hyphae lending a lanate and granular appearance to the colony surface;
thin, white margin surrounded by a wide fringe of submerged hyphae
cxtendin g about 3-4 mm beyond the aerial growth; heavily sporing

00

8 00
0 0 0
000 O

00
00

0
l..w.i.l=J
10 urn

CJ~[)
Iioo~l

fR .

Fig. 99. Camera lucida

drawings of Penicillium
sizovae Baghdadi , CBS
413.69. A, Detailed drawings
of the penicilli . B, Mature
conidia. e, Habit sketches .

throughout in blue-gray shades (Sgu y , PI. XXXIII , 493) with conid ial areas
partiall y ob scured by th e m ycelial tloccose overgro wth; colony surface
rad iately fu rro wed a nd slightly raised tow ards th e centre ; ex uda te a bundantly
produced, as crys ta l clear, uncoloured drops ; odour lacking ; no coloure d
pi gm ent diffu sin g into th e surrounding a ga r ; colony reverse (colour plat e 99, a)
in light cin na mon shades; penicilli cha racte ristically divaricat e (fig. 99 , A) ,
co ns isting of verticils of fairly equal metulae , conspicuo us ly divergent , but
suggesting th e Biverti cillata- symmetrica in pa ttern, owing to th e regular
a rrange ment of metul a e at th e tips of th e cond iophores; sim ple penicilli borne
on lat er al branches from conidiophor es are not in freguently enco u ntere d;
co nid iophores smoo th walled , mostly arising fro m the ba sal felt , com mo nly
ra n ging from 50 to 200 IJ-m by 1.5- 2.8 IJ-m with a pices enlarge d in a ves icle up
to 5.0 IJ-m in di ameter ; m etulae in whorls of 2- 4 eleme nts in th e ver ticil ,
rangin g from 11 to 25 IJ-m by 1.5- 3.5 IJ-m , some times of eq ual len gth but
u suall y of unequal size; phialides bottle shape d, abru ptly tapering to a sho rt,
narrowed n eck , commonly 9- 14 IJ-m by 3 .0 -4 .0 IJ-m , in crow de d clus ters of
4-12 elements in the ve rticil; conid ia globose, com pa ratively sm all , sm ooth
walled whe n obse rve d under th e ligh t mi croscop e (fig. 99, B) but delicately

ro ughcncd wh en observed un de r the SE M (plat e 17, centre : left), usually

" .5-:LO tlTl in di arn eter ; con id ia l cha ins formin g lon g, well-defin ed divergcnt
1'0 111 IJIns ,

up to 200 tm in Icn gth (fig. 99, C ).

olon ies upon C YA (colour plate 99, B and b) spread ing broadly, a ttain ing
'1 dia metc r of 55 mm in 14 days at room temperature (20- 25C), cons isting of
I closc-textured basal felt , typi cally velvety , overla id in central and subce ntra l
ll't:as by a fin e network of closely woven ae rial hyphae influen cing th e colony
lppca ra nce; thin , evenly margin surro unde d by a frin ge of subme rged hyphae
cxrcnding a bo u t 4- 5 mm beyond the aeri al growth ; colony sur face deepl y
xu lca ted by numerous radial furrow s and by conce ntr ic shallow wrinkles;
hcnvily sporing throughout in blue-gr ay shad es (Sguy , PI. XXXIII , 493) with
r onid ia l areas cons picuo us in m argin al zon es but obscure d in central and
subcentral a reas by the fin e ove rg rowth of white ae rial hyphae ; ex udate
ulnmd antl y produced, as yellow ish-co lou red large drop s; odo ur lacking ; no
ro lou red pigm ent diffu sin g into th e surrou nding agar; colony reverse (colour
plat o 99, b) in brown shades with margin in pale cre am shade s; penicilli
rs scntially as described aboye .
Co lonies up on M EA (co lour plat e 99, C and c) a tta in ing a diameter of 55
111111 in 14 days a t roo m temperature (20- 25C), cons istin g of a looser textured
hnsa l felt, ove rlaid by sector s of my celial , closely interwoven , floccose ae rial
hyph ae, partiall y obscuring th e othe rwise heav ily sporing a reas; conidial are as
in dcep gray-g ree n shades (Sguy, PI. XXVII , 402); thin margin surro unde d
11Y a frin ge of subme rged hyphae extending about 4 mm beyond the aerial
.;rowth; odo u r faint ' mo uldy'; exudate lacking ; no coloured p igm en t diffusin g
iur o the su rr ou ndi ng agar ; colony reve rse (colou r pla te 99, c) in cinnamon
xhad es with sector s of a gree n ish cas t showing throu gh ; penicilli essentia lly as
dcscribed aboye.
T he spec ies description is centered up on th e typ e stra in CBS 413.69 = C M I
110 ,344 = VK M F-1,073 = IJFM 7108 , isolat ed by V . Ch. Ba ghdadi fro m
Syrian desert soil near Mysalum , in the D amascu s region , Syria, M arch ,
1963. Two addition al strain s duplicat in g th e cha rac ter istics of morphology a nd
cu ltu ra l as pec t we re isolat ed by th e sa me autho r from Syr ian desert soil near
Bluda n a nd Berza respectively, in th e same region, M arch , 1963.
Penicillium sizovae wa s tentatively placed with spec ies of Penicillium inc1uded
wit hin th e P. chrysogenum series (Baghdadi, 1968) .

100 . Penicillium nooae-caledoniae Sm ith : fig. 100, colour plate 100, and
SEM plate 17, (centre right ).
Penlcillium novee-celedoniee Smith - Trans. Brit. Mycol. Soc o48: 273. 1965. See
also M. M . Kulik, 'A Compilation of Descriptions of New Penicillium Species'.
Agric . Handbook No. 351. Agric. Res. Ser. U.S. Dep. Agric. p. 58-59. 1968.

.".

"
",

..x:
'

:::

:::
~

10 jrrn

v-

,:,'

'::

0 00 0

i-:
O

O 00 O
O B O 000
O O O 00 0
0 0 00

r.~C(

fR. .

!,-tl t
100jJm

Fig. 100, Camera lucida

drawings of Penicillium
novae-caledoniae Smith,
CMI 140.441. A, Detailed
drawings of the penicilli . B,
Mature conidia. C, Habit
sketches.

Smith's original diagnosis follows, since this adequately presented the species in
question when first isolated:
'Colonies on Czapek's agar growing rapidly, bluish gray-green at first, becoming
grayer and duller, with narrow white margin during the growing period, thick
velvety to matted floccose, with reverse at first buff to dull pink, becoming pinkish
brown; penicilli usually with terminal verticil of metulae, these being commonly of
different lengths in the same verticil, sometimes with a secondary branch from the
next lower node of the conidiophore, typically divaricate; conidiophores rough,
mostly 3 tm in diameter; metulae 10 to 15 tm by 3 tm, often widening somewhat
towards the summit; phialides short-pointed, 8 to 10 tm by 2.2 to 2.5 tm; conidia
slightly rough, subglobose, 2,5 to 3.0 tm in diameter; conidial chains at first more or
less parallel, soon becoming tangled.
Isolated from soil, New Caledonia, by B. Huguenin, 1964.'

O ur n ot es follow :
Colon ies upon C A (co lour pla te 100 , A a nd a) a ttain ing a di amet er o f 25 - 30
mm in 14 da ys a t room te mperatu re (20 - 25 C), cons ist ing of tou gh , close-

tcxtu rcd basal Ielt., vc1vcty to sublan at c in a ppcara ncc , with whitc dcnticulatc
m urgin , surrounded by a frin gc 01' su bme rgcd hyphac cxtcncl ing ab out 2 mm
bcyond thc acrial growth ; colony su rface slightly su lca ted by shallow radial
Iurrows; hcavily sporing throughout in blu e-gray sha de s (Sg uy, PI. XXI ,
~1 ()1I ) ; cxud a te and odour lacking ; pal e vinace ous coloured pigm ent diffusing
Il lo the surrou nd ing ag ar; colony reverse (colour plat e 100, a) in purple-brown
xhudcs ; penicilli confor m with Smith's origin al diagnosis (fig . 100 , A) ,
CllIlsisting 01' con idiophores co nspicuously wa rty, ranging from 50 to 300 .tm in
lcn gth by 2.8-3 .5 .tm in diamet er, septa te, with terminal verticils ofmetulae 01'
u ncqual len gth , com mo nly 18- 25 iut: by 2. 5- 3. 5 uru , with a pices enla rge d in
1 vcsicle up to 4.0 tui: in diamet er; phialides bottle shaped, tap ering abruptly
lo a short narrowed neck , 10-14 .tm by 3 .0 - 4 .0 tui ; con id ia globose to
Il lll>globose, appe a ring smooth 0 1' nearIy so when observed under th e light
iu ic roscope (fig . 100, B) but delicat ely rou ghen ed when obse rved under the
SJo: M (pl at e 17, cen tre: right); co n id ial chains at first loosely divergent , soon
hccorning tangled (fig . 100, C ) .
Co lon ies up on CYA (colour plat e 100, B and b) spreading broadl y , att a in ing
I d iameter 01' 50 mm in 14 days a t room temperature (20 -2 5 C ), co ns isting 01'
a close-textured basal felt, deeply velvet y , ove rIa id by a th in m ycelial
uvcrgrowth cons isting 01' a closely woven network 01' fin e aerial hyphae
iu fl uenc ing gre a tly the colony a ppearance , lending to it a subla na te aspec t;
c: olony surface radiat ely furrowed , with white margin surrounded by a narrow
Iringe 01' submerge d hyphae ex tend ing abo ut 1 mm bey ond th e aerial grow th;
iu cdium sporing in pale brownish gree n shade s (Sguy , PI. XLVII , 693);
exudare and odour lacking ; reddish brown- coloured pigm ent diffu sin g into th e
xurrou nding agar; colony revers e in sienna reddish brown colou rs (co lour plat e
100 , b); penicilli esse ntially as described above .
C olonies upon MEA (co lour plate 100 , C and c) attaining a diamet er 01' 35
111111 in 14 day s at room temperature (20-25 C), cons isting 01' a close-t extured
basal felt , typi cally velvety to su bla na te , owin g to a loose ove rgrowth 01' floccose ae rial hyphae overIaying the colony surface; margin entire ; heavily
sporing throughout in mou se gray shades (Sgu y , PI. XXXV , 523); exu da te
nnd od our lacking; colony reverse (colour plat e 100 , c) developing typi cal
stra wbe rry red sha de s; sim ila rIy coloured pigm ent slowly diffu sin g into th e
sur rounding aga r ; penicilli esse ntia lly as described aboye, with con id iophores
a rising as short bran ch es fro m th e netw ork 01' tr ailing hyphae.
The spec ies description is centered upon the stra in IMI 140 ,441 = IJFM 7196,
rcceived in 1979 from th e Com m onwealth M ycological Institute, K ew , Surrey ,
England , and isolat ed by MI' Ian H . Pallet from de cayin g Lycopersicon sculentum
roots in soil near Betchw orth, Su rrey , U.K . The typ e strain IMI 104 ,1 56 ,
isolated from N ew Caled oni an soil by B. Hugu enin in 1964 , unfortunately lost
Irorn th e CMI cu ltu re collection, could not be exam ined by the au thor 01' th e

.urrcnt study, because a ll attempts to obtain it Irom ot hcr ln rernation a l

C ult ure C ollcc tions fail cd . Howevcr, in o ur opin ion, strain IM I 140 ,441
reprcscnts qui te satisfactorily the spe cies in question as des cribcd by G . Srnit h.

101. Penicillium nooae-caledoniae Smith, varo album R amrez et M artn ez:

fig. 101 , colour plate 101 , an d SEM plate 17, (bottom : left).

,
"

"

,e
Ll..W..W.J.JJ

10jJm

"
"

'.

000
00 0 B
O

0 00 0
O O 0 0

Fig. 101 . Camera lucida

drawings of Penicillium
nove e-celedoniee Smith varo
album Ramrez et Martnez,
IJFM 7184. A , Detailed
drawings of the penicilli. B,
Mature conidia . e, Habit
sketches.

00

tt

100jJm

Penicillium novae-caledoniae Smith , varo

album

Ramrez et

Martnez

Mycopathologia. 74: 35-49, 1981.

Co lon ies u po n CA (colour plate 1 1, A and a) attaining a diameter 01' 45 - 50

m m in 14 days at room tempe rature (2-25 C), consis ting 01' a close-textured
basal felt appearing sub la nate, with eve n m argin in pale pink ish cream
shades, surro un de d by a fringe 01' submerged hyphae extend ing abo ut 2-3 mm
beyon d the aerial growth; colony surface radiately furrowed, slightly ra ised
towards the centre, ove rlaid by a de nse network 01' closely interwoven ae r ial
hyphae rem aining persiste nt ly white; poo rly spo ring; exudate limi ted in

IrI101lnt, as min u te d roplct s c n mcs hc d in the m ycc lia l 1'1.:11 ; no co lou rcd pigrncnt dil'l'lIs in g into th e su rr ou nd ing agu r : colo ny re ve rso (co lou r plate 101, a)
11 palo cin namo n sha de s with rna rgin yellow ; pcni cilli typically divaricate (li g .
101 , A) com monly cons isting 01' verticils 01' 2 -:~ me tu lae of unequal len gth ,
umc timcs with a seco ndary branch , ran gin g fr orn 16 to 20 /lm by 2.8 - 3 .5 uiu ;
cu uid iop horcs with wall s co ns p icuo us ly warty, a risin g frorn th e b a sal felt,
n lllslly 50 - 150 /lm by 3. 0 - 4 .0 urtv; phialides bottle sh ap ed , tapering to a sho rtpui utcd ncck , in clus te rs 01' 3 -9 eleme nts in the ve rt icil, co m mo nly 10- 12 /lm
liy :1.0- 4.0 psi ; co n id ia globose to sub globose , mostly 2. 5 - 3. 5 tut , smo oth
wullcd 0 1' nearly so wh en observe d under th e light mi croscope (fig . 101 , B) bu t
k-lica tely rough ened whe n obse rve d under the SEM (pla te 17, bottom : left);
ru uid ial chains at fir st more 01' less div ergent , becoming tangled in age (fig.
lo 1, C) .
Colon ies upon C YA (co lou r plate 101 , B a nd b ) sp read ing broadly , attaining
I d ia rn et er of 55 mm in 14 d a ys a t room temperature (2 0 -2 5C), co ns ist ing 01'
I rlosc -textu re d ba sal felt, ove rlaid by a dense m ycelial network 01' ae r ia l
hyphac, with shallow rad ial fu r rows n ear th e m argin ; gro w in g m argin
MlII'l'Ou nd ed b y a fr inge 01' sub merged hyphae , exte ndi ng a bo ut 5 mm beyond
111(' ac ria! gro w th; ve ry poorly sp oring, rema ining persistently white ; odo ur
,ll'O ng , suggesti ng mushrooms; ex u date lacking; pale a m be r-colou re d p igment
d illu sin g int o th e su rro u n d ing agar; colony reverse in reddish brown shades
(I'lllou r pla te 101 , b); penicil!i esse n tially as described a bo ve .
Colon ies upon MEA (co lou r plat e ,101 C a nd e) gro win g rather restrict edly,
II tta ining a di amet er 01' 35 mm in 14 da ys a t room temperatu re (20 -25C),
l'lIlIsist ing 01' a close- tex tu re d b asal felt , de eply lanat e , with ce n tre rai sed , w ith
c vc n margin su r ro u nde d by a frin ge 01' sub mer ged hyphae exte nd ing a bo u t
1-4 m m beyond th e aeri al growth; colony surface not furrowed , rem aining
pc rsistently white ; sparsely sporing , with con id ial structures a r ising as shor t
h run ches from ae r ial h yphae ; ex u da te and odou r lack in g ; colony reverse
('olour plat e 101 , c) at fir st in cin na mon sh ades, slowly developing the typi cal
sI ra wbe rry red co lou r characte r istic 01' th e spec ies.
T he va r iety duplicat es the typ e spec ies in colony texture , rat e 01' growth a nd
in the det ails 01' m orphology , but differs from it in th e persistently white obve rse
lll' (he colon ies upon al! subs tra ta tested .
The description is ce n tere d upon strain IJ FM 7184, isolat ed by th e a u tho r 01'
the curre n t stu dy in 1979 fro m the a ir 01' M adrid , Spain . It is at present kn own
nnly from th e type strain .

102. Penicillium citreo-otrens Abe ex R amrez: fig. 102, colour plate 102,
and SEM plat e 17, (b ottom : right).
Penicil/ium citreo-virens Abe - J. Gen. Appl. Microbiol. Tokyo. 2: 87-88. 1956. See

o
-

00
O

000
-:.n 8 0
OUO O
O

10jJm

y~

Fig. 102. Camera lucida

drawings of Penici//ium
citreo-virens Abe ex
Ramrez, ATCC 18,231. A ,
Detailed drawings of the
penicilli . B, Mature conidia.
C, Habit sketches.

also G. Smith, Trans. Brit. Mycol. Soco 46: 332. 1963; also M .M. Kulik, 'A
Compilation of Descriptions of New Penici/lium species'. Agric. Handbook No.
351. Agric. Res. Ser. U.S. Dep. Agric. pp. 28-29. 1968.

Penicillium citreo-oirens was isolat ed fromjapanese soi! by S . Abe in 1952 who

proposed the name P. citreo-oirens and described the species without providing a
L atin di agnosis (1956) . G . Smith (196 3) , examining 26 cultures of Ab e's newly
des cribed spe cies (1956), sent by the au thor to th e Commonwealth
Mycological Institute, K ew, Surrey, UK , stated that P. citreo-uirens Abe had to
be regarded as a synonym of P. sp inulosum Thom in the Monoverticillata
section .
The author of the cu rre nt study, after examining two subcultures of Abe 's
type strain of P. citrio-oirens received from the C entraalbureau voor
Schimmelcultures , Baarn, The Netherl ands, and from the Am erican Type
Culture Collection , Rockville , Maryland, USA , and CBS 320.59 and ATCC
18,231 , re spectively, ca rne to the conclu sion that Pen icillium citreo-oirens could

be rega rded as quite a distinctive species devcloping typical divari ca te

structurcs in sufficicnt number as to necessita te placing the species in the
Asyrnmetrica -v d iva ricata sub-sec tion of the Biver ticillata. Therefore , in our
llpin ion the ori ginal generic name should be maintained.
'1'0 validate the species a Latin diagnosis of it is given below ; the name of th e
or iginal author bein g maintained under R ecommendati on 46 C of th e
l nternational C ode of Botanical Nom encl ature (Stafleu (ed .) 1972).
Latin diagnosis of Penicillium citreo -uirens Abe ex R amrez:
Coloniae in agaro Czapekii post duas hebdomadas ad 25C, 35 mm in diametro
ott ingunt , laxo texto basali coacto mycelio, velutinae vel subfloccosae, cum
margine plano , aequaliter, fimbriato e hyphis submersis (2 mm) formatur, cum
cent ro fortiter plicato et crateriforme, parum radiatibus sulcis; parum sporulantes,
cum areis fructificationibus conid ialibus pallido viridibus; exsudatio parco; odor
debile; reverso pallido aurantio; sine colore diffluente in medio . Penicilli plerumque
fo rtiter asymmetrici divaricati vel etiam interdum monoverticillati, ergo in subsectio
Asymmetrica- divaricata inclusus est; conidiophora laevia, e substrato, 100-300 p,m
longa, vel etiam e hyphis reptabundis, 25-50 p,m longa, exoriuntur, per 2.5-3.0 p,m
in diametro, cum apicibus inflatibus tenus 5 p,m in diametro; metulae tenus 2-3 per
verticillum, plerumque 15-30 p,m per 2.5-3.0 p,m, cum apicibus inflatibus tenus 4.0
/tm in diametro; phialides lagaeniformae, ad basem dilatae sunt et ad apicem
abrupte angustate ad tubulum exiguum conidiophorum terminantur, 6-10 per
verticillum, pleraque 7-12 p,m per 3-4 p,m; con idia globosa vel subglobosa,
plerumque 3-4 p,m in diametro, laevia; densis diversis columnis formantur.
Coloniae in agaro Czapekii cum extracto levedinis, post duas hebdomadas ad
25 C, 40 mm in diametro attingunt, velutinae, plicatae, cum radiatibus sulcis;
fortiter sporulantes; cum areis fructificationibus conidialibus pallido griseo-coeruleis
colorantur; exsudatio abundantis, flavido coloratur; odor parco; reverso aurant io;
sine colore diffluente in medio .
Coloniae in agaro hordeaceo celeriter crescunt, post duas hebdomadas ad 25C,
55 mm in diametro attingunt, tenues , velutinae, planae, laxo texto basali coacto
mycelio; margine effusa, fimbriata e hyph is submersis (5 mm) formatur;
fructificationibus con idialibus abundantibus, coe ruleo-viridibus colo rantu r;
exsudatio et odor absunt; reverso virid i in centrali area, sine colore in marginali area;
sine colore diffluente in medio.
Typus cultura IFO 6030 e solo Japonico a S. Abe, anno 1952 isolatus esto
Abe's original diagnosis follows:
'Colonies on Czapek's agar grow rather rapidly , attaining a diameter of 47-49 mm
in 10- 12 days at 25C, consisting of a somewhat compact basal felt, appearing
typically velvety, with a w hite margin about 1.0 mm wide, smooth, sporulating
abundantly in bright yellow-green shades near empire green, becom ing cress green
with age; exudate lacking; odou r limited or indefinte, mouldy; reverse usually in

cream shades; becoming typically green-ollve In the central ares and sepia in the
marginal areas; surrounding agar usually colourless; conidiophores arising mostly
from the substratum or from the basal felt, mostly 60-250 p.m in length by 2.5-2.7
p'm in diameter, with apices 2.8-4.7 p'm in diameter and with walls smooth or nearly
so, usually unbranched; penicilli typically biverticillate and asymmetrical, consisting
of a terminal group of 3-5 compact or slightly divergent metulae that measure
about 12.5-25.0 p'm by 2.5-3.7 p'm (apices commonly enlarged to 2.6-4.7 p.m),
each supporting a cluster of 3-7 compact or parallel sterigmata (phialides)
measuring about 7.5-11.9 p'm by 1.7-2.5 p'm, and bearing conidia in columns or
loosely tangled chains up to 60-120 p.m in length; conidia globose to subglobose,
mostly 2.3-3.3 p'm, smooth walled or nearly so; and conidia walls delicately
roughened (spines less than 0.1 p.m) as shown in electron microscopy.'

Our notes follow:

Colon ies upon CA (colour plat e 102 , A a nd a) attaining a diameter of3 5 mm
in 14 days at room temperature (20- 25C) , cons isting of a rather loosetextured basal felt, appearing velvety or slightI y lanat e , with m argin plane
surro un de d by a frin ge of submerged hyphae exten d ing abo ut 3- 4 mm beyond
th e aeri al growth, with colony centre deeply radiately furrowed a nd raised up
to 1-2 mm deep in a crateriform pattern ; slightIy spor ing, especially in
m arginal areas in light gre en shades (Sgu y, PI. XXVII , 404) ; ex udate limited
in am ount, as small dropl ets enmeshe d in th e mycelial felt; odour faint,
indefinite; no coloure d pigment diffu sin g into the surrounding a ga r; colony
reverse (colour plate 120 , a) developing oran ge shades in the centre and pale
cream in the margin ; penicilli esse ntially conform to the original descr iption ;
the con id iopho res are variabl e in len gth , dep ending up on their origin, usual1y
fairl y long, ranging from 100 to 300 p.m long by 2. 5-3.0 p.m in d iamet er wh en
arising directIy from the su bs tra tum, to quite sho rt, less than 50 p.m in length
when borne as branch es from tr ailing hyphae , with walls smo ot h, with a pices
enlarged in a ves icle up to 5.0 p.m in diameter ; the penicilli cons ist usu ally of
wh orl s of metul ae of unequal length , with 2- 3 elements in th e ve rticil,
typi cally div er gent, ranging from 15 to 30 p.m by 2.5 -3 .0 tuu , with apices
enlarge d in a vesicle up to 4 .0 p.m in di ameter; phialides bottl e sha ped,
ab ru ptIy tapering to a short na rrowed neck , in clus ters of 6-10 eleme nts in th e
verticil, m ostIy 7-12 p.m by 3. 0- 4. 0 p.m; conid ia globose to subglobose,
usually 3 - 4 p.m in diamet er, appearing smooth wall ed wh en observe d under
th e ligh t mi croscope (fig. 102 , B) and very delicately roughened when obse rve d
under the SEM (pl at e 17, bottom : ri ght); con id ial chains in fairly well-defined
di vergent colu m ns (fig . 102 , C ). Simple penicil1i (appearing m onoverticillate)
are not infrequentI y obse rved, usually borne from short conid iophorcs ar ising
directIy from tr ailing hyphae .
C olonies up on CYA (colour plate 102 , B and b) atta ining a diameter of 40
mm in 14 days at room temperature (20- 25 C), typi cally velvety , with margin

"
pluu c surrou nded by a fringe 01' submcrgcd hyph ae extcn d ing abo u t 1 mm
licyond thc acria l growt h; su bmarg ina l a nd ce n tra l a rcas dccply radi atcly
1'1I1'I'Owcd and de p res scd in th c ce n tre , co lon y su rfacc co ns pic uo usly zanate ;
hunvi ly spo rin g throughout in d ccp blu e-grecn sha des (Sguy, PI. XXXI,
;1(11); cxud a te abundantly produccd , cs pec ially in th e ce n tre as large, cr yst al
C1l:11 l', colou rless drops; odou r la ckin g 0 1' indefinitc ; nu colou re d pigment
d lllu sing into the surrounding ag ar ; colony revcrse (col our plate 102 , b)
d evc lop in g ora n ge sh ades ; penicilli essen tia lly as described .
:o\on ies upon MEA (col our plate 102 , C and c) sp reading broadly ,
lll /lilling a di ameter 01' 55 mm in 14 days a t room temperature (20- 25C),
c't1l1 sis ting of a loo ser textured basal felt, slightly zona te, typically velvety , with
1II Ill'g in thin , su r ro u nde d by a wide frin ge 01' su b merg ed hyphae ex te nd ing
rho u t 5 mm beyond th e aerial grow th ; colony su rface not furrowed , plane w ith
(' (' 111 re sligh tly umbon ate; hea vily spo r ing thro ugho u t in very dark blue- green
hades (Sguy , PI. XXX , 446) ; ex udate and odo u r lacking; coloured pi gment
d iflus ing into the su r ro u ndi ng aga r is not observed; colony rev erse in crea m
Mhades with a d a rk green cast sho wi ng through in the cen tral area ; penicilli
,'sselltially as described abo ye.
T he species description is ce nte re d upon th e type strain IFO 6 ,0 30 = ATCC
1ll , ~nt = CBS 320.59 = CMI68 ,2 30 = IJFM 7069 , isolated fromJapanese soil
hy S . Ab e in 1952 , a nd many othe r strains isolat ed b y th e a u tho r fro m Sp anish
stli1.
Penicillium citreo-uirens is in clu de d in th e Asymmetrica-divaricata sub-sec tion
s iucc the m aj ority 01' th e penicilli sho w this b asic pattern. M any struc tu res,
huwcver , ap pear clearly monoverticillate w ith individual penicilli borne singly
1111 sho rt branches. While we rega rd th e present placement as co r rec t, th e
p robabl e relationship 01' th e species to th e M onoverticillat a , possibl y as a
t rn nsi tio nal for m, sho u ld not be di sregarded .

Penicillium canescens series

O UTSTANDlNG FEATURES
Colon ies gro w ing some what restrictedly upon m ost subs trata, consisting 01' a
l'airl y de nse basa l felt wi th su r face growth loose textured , m ore or less lanat e ,
ron id ial a reas in ligh t ye llow-green to gr ay- gr een shades, co lony rev erse var ia ble, ra nging fro m dull peach to orange -b rown or orange-red , depend in g on
(h e spec ies.
Conid iopho res va r iable in origin a nd d im ensions , a r ising eithe r fro m th e
su bstratu rn, atta in ing up to 400 - 500 ,m in length , bearing metulae eithe r irrcg ularly arranged or in fa irly well-defined terminal ve rticils; or b orne as sho r t
h ran ch es from ae r ia l hyphae , u su all y much sho rter, a nd bearin g smalle r a n d
mor e irregular penicilli.

Peni cilli cha rac teristically div aricatc , wilh eo nid ial chains typically ad herent
in loase di vergcn t columns .
Con id ia globose to subglobose, with wa lls rang ing from smooth or nearly s
to conspicuo usly roughe ne d, de pe nd ing upon the spe cies.

K ey to species
Paga
1. Penicilli with the divaricate character well marked; growing rather
restrictedly, at least upon some substrata; with metulae scattered on the
conidiophore, or usually aggregated into true verticils. Colonies
develop ing typically pale blue-green or gray-green shades with the
ripening of conidia ; colony reverse often brightly coloured. Colonidial
chains tending to produce compact columns , at least in young cultures;
conidia globose to subglobose.
P. canescens series ... . 283
A. Colony reverse developing deep red or brown shades; penicilli
strongly divaricate , not tending towards ramigenous.
1. Colonies up to 0.5-1 .0 mm deep, with surface loase, more or less
lanate; conidial areas in dull blue-green shades; conidia about
2.0-2.5 Jlm in diameter; colony reverse orange, becoming brown in
age.
P. canescens Sopp .. . . 285
2. Colonies deeply lanate, up to 2.0-3.0 mm deep; conidial areas in
brighter greenish glaucus shades; conidia about 3.2-3.6 Jlm in diameter; colon y reverse in deep red shades near maroon.
P. nalgiovense Laxa .. .. 287
3. Colonies appearing typically velvety.
a. Conidiophores
smooth
walled;
conid ia
conspicuously
roughened.
P. turolense Martnez et Ramrez . . . . 293
b. Conidiophores delicately roughened; con idia strongly echinulate .
P. murcianum Ramrez et Martnez . ... 291
B. Colon y reverse orange , in dull peach shades or colourless, not
developing dark colou rs; penicilli often appearing somewhat
ramigenous.
P. jensenii Zaleski . .. . 289

The Penicillium canescens series was erected by R ap er an d T ho m (1949)

principally to cover species in which the divaricate character was well marked,
b ut which did not fit very satisfactorily into any recognized series. It is the n
probably artificial in character. H owever , a certain degree of re lationship
see med to exist between the spe cies incl uded in it , namely P. canescens Sopp , P.

Ifllgiovtmse Lax a , and P. [ensenii Zaleski, AlI 01' thcm tcnd to produ ce conicl ial
chuins in loose colu mns, all produce lon g conidiophores, often with I'a irly wellurgunized terminal peni cilli , a nd all prod uce globoso to subglobo se conid ia .
'I' wo add itional species have been inclu dcd hcre in the cur rent stud y, namcly
l'enicillium turolense M art nez et Ram rez , an d P. murcianum R arnfrez el
M ur tncz .

10:1. Penicillium canescens Sopp : fig. 103, colour plate 103, and SE M
plate 18, (top : left ).

..:.~:

.:.

:::;

(f~

(1:&
~

~g;)

iJ.~r) (~ (~

i::::.' ~

;".:

A~~1~
1~

;'.:

--:-

e"R,

Fig. 103. Camera lucida

drawings of Penicillium
canescens SoPP. CSS
300.48. A , Detailed draw ings
of the penicilli . S, Mature
conidia. e, Habit sketches.

Penicillium canescens Sopp - Monographie, pp. 181 -182. 1912. See also Ch.
Thom, 'The Penicillia', pp. 347-348.1930; also Raper and Thorn , 'A Manual of the
Penicillia', pp. 316-319. 1949.

C olon ies up on CA (colour plate 103, A and a) spreadi ng broadly in some

strains, atta in ing a d iameter of 50 mm, and somewha t restrictedl y in othe rs,
attaining a dia meter of 35 - 4-0 mm in 14- days at room temperature (20-2 5C),

~1Il>

with surlacc more or less lana te consisting 01' a loose -tcxturcd network 01'
interwoven vcgetative hyphae arising from a close-tcxtured, compact basal felt ,
lending a deeply floccose appearance to the colon y surfacc; thin margin 2-1mm wide, surrounded by a fringe 01' submerged hyphae extending about 2 -3
mm beyond the aerial growth; colony surface convex; fairly abundantly
sporing throughout in pale gray shades (Sguy, Pi. XVI, 235); exudate
limited, pale amber, as minute droplets enmeshed in the mycelial felt; odour
lacking or indefinite; no coloured pigment diffusing into the surrounding agar;
colony reverse developing yellow shades (colour plate 103, a); penicilli variable
in sizc (fig. 103, A), borne either upon conidiophores arising directly from th e
substratum and varying in length up to 500 ,tm by 2.5-3 .5 ,tm in diameter, or
upon short branches from aerial hyphae, with walls ranging from
conspicuously warty to delicately roughened or nearly smooth , consisting 01'
variously arranged divergent structures, forming verticils 01' 2- 3 metulae
usually 10-25 ,tm by 2 .5-4.0 tui, bearing clusters 01' phialides 01' 4-12
elements in the verticil, mostly 6-10 ,tm by 2.5-3 .5 tut, bottle shaped,
terminating in a tapered, short narrowed neck; conidial chains tending to
adhere into loose columns at least in young cultures, but becoming more or less
divergent in age (fig. 103, C); conidia at first ovate to subglobose becoming
globose at maturity, with walls conspicuously roughened when observed under
the light microscope (fig. 103, B) and showing a definitely warty surface when
observed under the SEM (plate 18, top: left), usually 2.5-3.0 ,tm in diameter.
Colonies upon CYA (colour plate 103, B and b) spreading, attaining a
diameter 01' 50-60 mm in 14 days at room temperature (20-25C), consisting
01' a deeply velvety to lanate felt, deeply radiately and irregularly furrowed,
with concentric ridges, with centre depressed in a crateriform pattern; with
white even margin 5 mm wide surrounded by a narrow fringe 01' submerged
hyphae extending about 1 mm beyond the aerial growth ; abundantly sporing
throughout in brownish gray shades (Sguy, Pi. XXIX, 428); exudate limited,
as small droplets enmeshed in the mycelial felt; odour faint, 'mouldy'; reddish
brown pigment diffusing into the surrounding agar; colony reverse (colour
plate 103, b) developing deep brown, almost black shades ; penicilli essentially
as described aboye .
Colonies upon MEA (colour plate 103, C and c) growing restrictedly,
attaining a diameter 01' 25 mm in 14 days at room temperature (20-25C),
consisting 01' a loose-textured basal felt, lanate, somewhat funiculose, about 1
mm deep, not furrowed, with margin irregularly dissected in a denticulate
pattern surrounded by a narrow fringe 01' submerged hyphae extending about 1
mm beyond the aerial growth; heavily sporing throughout in mouse gray
shades (Sguy, Pi. XXXV, 523); exudate and odour lacking; reddish browncoloured pigment diffusing into the surrounding agar; colony reverse (colour
plate 103 , c) developing dark reddish brown colours; penicilli essentially as

describc d a bo ye but sho w iug a tc ndency lo p rod uce mo re co m puc t colu mna 01'

cunid iu.
'I'hc spccies d escr ip tio n is ce n tc rc d upo n strni n CIIS :~ 00 .4 8 - CM128 ,260 10,41 9 = NRRL9 10 = Q M 7550 .. I./F M :~ 866 , isola ted b y Miss E . Dale in
1';Ilf.{l an d , fro m soil, in 1912, cited b y T hom in 1930 a nd by Raper a nd T hom
i 11 t 949 as representing th e spcc ics.
l'enicillium canescens appears to be som ewha t tr ansitio nal betw een P.
j anthinellum a nd P. nigricans. It gro ws more res trictedly , at least up on M EA
I ha n the spe cies 01' the for rner seri es ; it prod uces deep brown rather tha n
pu rpl e-red colours in colony rev erse , suggestive 01' P. nigricans; a nd conid ia are
globose and rou gh as in the la tter series .
A'J'l: G

104 . Penicillium nalgiovense Lax a: fig. 104, colour plate 104, and SE M

plate 18 , (top: right ).

.:'

o O

00
BO O O
O 00
00
O 00

i.w.I..WW

10J.lm

0 0 0

fR.,

() Cj OO O

0 0

-r:
I..-----.--J

100J.lm

Fig. 104. Camera lucida

draw ings of Penicillium
nalgiovense Laxa, CBS
352.48. A , Detailed drawings
of the penicilli. B, Mature
conidia. e, Habit sketches.

Penicilli um na/giovense Laxa - Z. Bakt. Parasitenk . Inf. Hyg. 11 Abt. 86: 160-165.
1932.See also Raper and Thorn, 'A Manual of the Penicillia', pp. 319-322. 1949.

Colonies upon CA (colour plate 104, A a nd a) growing restrict cdly , a tta in ing 11
diameter of 30 mm in 14 days a t room ternpcraturc (20 - 25 C), co ns isting al' 11
fairl y closely woven network 01' vegetative hyphae lending a de eply lanat e
appearance to the colony 2- 3 mm deep, slightly radiately furrowcd, somewhat
zonate , at first white but gradually developing pale yellow- gree n shadcs wi th
the ripening of conidia; thin , eve n margin surrounded by a fringc 01'
submerged hyphae extending about 1-2 mm beyond th e a erial growth;
sparsely sporing over most of th e colony su rface , becoming yellow-green
(Sguy, PI. XXIII, 335) in fruiting areas and usually remaining white to ro sy
white in areas of purely vege ta tive growth ; exu da te abundantly produced as
large golden ye llow drops; od our la cking; yellow-coloured pigment diffusing
into th e surrounding a ga r ; co lon y rev erse in ye llow-ora n ge colou rs (colour
plate 104, a) ; penicilli strongly divaricate (fig. 104 , A) , variable in pattern ,
consisting o f occ asion al branch es, and metulae varying gr eatly in arrangc ments and in size; con id ial structures arising commonly a s short branches frorn
aerial hyphae , less frequently from th e substratum ; con id iophores ranging
frorn very short to up to 500 p.m in length by 2.8-3.0 p.m in diameter, with
walls gen era lly delicately roughened , more or less septate ; metulae mostly in
small clusters of 2- 3 elements in the verticil , usually 7-1 3 p.m by 2.8- 3.5 p.mj
phialides com p a ra tively small , bottle sh a ped , terminating abruptly, without
consp icuou s n eck , mo stly 6- 8 p.m by 3. 0 - 4 .0 tut , borne in clusters ol' 2-6
elements in th e ve rticil; conidia elliptica l, subglobose or globose, appearing
smooth walled when ob served under th e light mi croscope (fig. 104 , B) but ve ry
finely roughened when obs erved under th e SEM (plate 18, top: right), mostly
3. 5- 4. 5 p.m by 3. 0 - 3.5 tut: in diameter, sorne occasionally larger, up to 6.0 p.m
in long axis ; con id ial chains short, tending to adhere into narrow colu m ns (fig .
104 , C) .
Colonies upon CYA (colour plate 104 , B and b ) attaining a di ameter of 40
mm in 14 days at room temperature (20-25 C) , similar in texture to those
grown upon CA, but d eep er in the ce n tre and su lca te d by numerous radial
furrow s near th e margin, whi ch is su rro u n ded by a very narrow fringe of
su bmerged hyphae not exceed in g 1 mm bey ond th e aer ial growth; sporing very
poorly with colony surface remaining persistently white or turning to yellowish
tints; exud a te as very small drops em bedded in the mycelial felt, colou rle ss;
odo u r la cking; yellowish colou re d pigment diffusing into the su r rou nd in g agar;
colony reverse (colour plate 104, b) in yellow-ora n ge shades; penicilli
ess entially as described aboye .
Colonies upon MEA (colour plate 104 , C and c) growing very restrictedl y ,
co m pa ratively thin , attaining a diamet er of 20 mm in 14 days at room
temperature (20-25 C), consisting of a close-textured basal felt , deeply
vel vety to lanate , with fin ely denticulate m argin largely submerged exte nd in g
about 2 mm be yond the aerial growth, not furrowed ; colony surface remaining

persistently white in arca s 01' purcly vegetat ivo gro wth, sha ding to pale yellowgreen shad es in areas abundantly sporing, particularly in the colony centre
(S guy , PI. XXVI , 380); exudate and odo u r lacking; no coloured pigment
diffusing into th e surrounding agar; colony reverse in yellow-ora nge shades;
penicilli essentially as described ab oye but with conid ial chains lon ger a nd
columns somewha t tangled .
The spec ies description is centere d upon the typ e strain CBS 352.48 = C M I
39, 804 = ATCC 10,472 = NRRL 911 = IFa 8,112 = QM 7,600 = IJFM 7055 ,
isolated by O . Laxa from ElIischauer chee se (01' the C amembert type), in
Nalzovy, Southern Bohemia , in 1932.

105. Penicillium jensenii ZaIeski: fig. 105, colou r pIat e 105, and SEM
pIate 18, (centre : Ieft).

~':; :~ B' : ;2; : '

::.;:' ~

{~~ :~ ,:':i~

{ :;\

;fr'f r~ <' .,
L--...J

100 pm

Fig. 105. Camera lucida

drawings of Penicillium
jensenii Zaleski, CBS 216.28.
A , Detailed drawings of the
penicilli. B, Mature conidia.
e, Habit sketches.

Penicillium jensenii Zaleski - Bull. Acad . Polonaise Sci. Math . Nat . Sr. B, pp.
494-495. 1927. See also Ch. Thom, 'The Penicillia', pp . 346-347.1930; also Raper
and Thom, 'A Manual of the Penicillia', pp . 322-323. 1949.

Synonym
Penicillium chrzsszczii Zaleski -- Bull. Acad. Polonaise Sci. Math. Nat. Sr. B, pp.
464-466. 1927.
Colonies upon CA (colour plate 105, A and a) growing restrictedly , attaining a
diameter of25-30 mm in 14 days at room temperature (20-25 C), consisting
of close-textured basal felt about 500 ,tm deep , strongly folded and radiately
wrinkled, with marginal area abruptly raised and centre d epressed, with the
subcentral underlying area com m only pulling away the agar from the culture
dish and cra cking it , with colony surface appearing loos e , finely granular,
almost lanate, more or less zonate, with abruptly raised margin 1-2 mm wide;
light to medium sporing in pale yellow-green shades (Sguy , PI. XXV , 365) in
fruiting areas and u sually remaining white in areas of purely vegetative
growth; exudate abundantly produced, particularly in the subcentral and
cen tr al depression as deep orange, crystal clear, large drops; odour lacking;
orange-coloured pigment diffusing into th e surrounding agar ; colony reverse
(colour pi ate 105 , a) developing deep orange shades; penicilli typically
divaricate (fig. 105, A), often appearing ramigenous, in larger structures
commonly con sistin g ofwell-defined terminal clusters of2, 3, or more metulae ;
con id iopho res variable , ranging from smooth walled to con sp icu ou sly warty
with warts sparsely arranged on th e conidiophore wall , arising from the
substratum up to 500 ,tm or more in length by 2 .5- 3 .0 ust: in diameter , or
borne as lateral branches on trailing hyphae , u sually shorter, not ex ceeding
100 ,tm in length, with apices enlarged in a vesicle up to 5 .5 tui: in diameter;
metulae va ria b le, usually 10-18 ,tm by 2.0-2.5 tui, with a p ices infl ated in a
vesicle up to 4 .5 ,tm in diameter , always smooth walled ; phialides bottle
shaped, usually in clu sters of 3-10 elements in the verticil, mostly 8 -9 ,tm by
2.5-3 .0 tun , conspicuously tapering to a short narrowed neck; conidia globose
to subglobose , 2.0-2 .8 tut: in diameter, with walls conspicuously but delicately
roughened (fig. 105 , B, and SEM plate 18 , cen tre : left) ; conidial ch a in s
tending to adhere in well-defined loose, divergent columns (fig. 105, C) .
Colonies upon CYA (colour plate 105 , B and b) , spreading broadly,
attaining a diameter of 45 mm, consisting of a close-textured basal felt, deeply
velvety in appearance, with the surface strongly and irregularly wrinkled and
folded in a more or less radial pattern, with the central area folded in a
crateriform pattern; undulating margin surrounded by a fringe of submerged
hyphae extending less than 1 mm beyond the aerial growth; heavily sporing
throughout in blue-green shades (Sguy, PI. XXX, 442) ; exud a te and odour
lacking; amber-coloured pigment diffusing into the surrounding agar; colony
reverse (colour plate 105, b) in reddish orange to cinnamon shades; penicilli
essen tia lly as described aboye but more abundantly produced .
Colonies upon MEA (colour plate 105, C and c) growing very restrictedly,

~Ul

altaining a diameter 01' 20- 25 mm in 14- days at room tcmperaturc (20 - 25C),
co nsistin g 01' a loose-tcxtured basal fclt, more or less lanate, very slightly
Iurrowcd and raised in the centre; with undulating margin largely submerged,
:xtending more than 10 mm beyond the aerial growth; heavily sporing in dccp
hlue-green shades (Sguy, PI. XXX, 447) : exudate and odour lacking;
colou red pigment diffusing into the surrounding ag ar not produced ; colo ny
reverse (colour plate 105, e) in yellow-orange shades; penicilli essentialIy as
described aboye but more consistentIy simple (appearing monoverticillate and
lending to ramigenous), with conidial columns more strongly accentuated than
tlpon the aboye substrata.
The species description is centered u pon the type strain CBS 216.28 = CM J
:~ 9 , 768 = ATCC 10,456 = IFa 5,764 = NRRL 909 = QM 7,587 = IJFM 7076,
isolated by K. Zaleski from soil in Poland, in 1927.

106. Penicillium murcianum Ramrez el Martnez: fig. 106, colour plate

106, and SEM plate 18, (centre: right).

"

f( .

Fig. 106. Camera lucida

drawings of Penicillium
murcianum Ramrez et
Martnez, ATee 42,239. A,
Detailed drawings of the
penicilli. B, Mature conidia.
e, Habit sketches .

Penicillium murcisnum Ramlrez et Martlnez - Mycopathologia. 74. 35-49, 1981.

Colonies upon CA (colour plate 106, A ano a) growing restrictedly,

attaining a diameter of 20 mm in 14 days at room temperature (20-25C),
consisting of a tough , close-textured basal felt , strongly heaped, about 3 mm
deep , velvety in appearance , not furrowed , with a very thin and narrow
margin; colon y surface overlaid by tufts of fruiting aerial hyphae, lending a
coremiform appearance to it; heavily sporing throughout in pale blue-green
shades (Sguy , PI. XXIX , 425), especially in marginal zones but obscured in
central and subcentral areas by an overgrowth of fine closely woven aerial
hyphae ; exudate abundantly produced , as dark amber-coloured drops ; odour
lacking; reverse (colour plate 106 , a) orange with a gre en cast showing through
in marginal areas; amber-coloured pigment slowly diffusing into the
surrounding agar; penicilli typically asymmetrical divaricate, consisting of
fairly regular verticils of metulae arranged at the tip of the stipe , or branches
arising from the conid iophores at different levels bearing small verticils of
metulae (fig . 106 , A); conidial structures arising either from the substratum or
from trailing hyphae ; stipes of the conidiophores finely roughened, commonly
up to 400 .tm in length by 2.0-3 .0 .tm when borne directly from the
substratum; branches smooth walled, mostly 10-20 tui: by 2.5-3 .0 .tm;
metulae in clusters of 2- 4 or more elements in the verticil, usually 9 -18 .tm by
2.5- 3. 5 iun, usually inflated at the apex in a vesicle up to 4 .5 .tm in diameter,
strongly divergent, smooth wall ed ; phialides comparatively small, bottle
shaped, abruptly tapering to a very short, narrowed neck, in clusters of 6-12
elements in the verticil , mostly 4.0-7 .0 .tm by 2.5-3 .0 .tm; con idia
comparatively small, globose to subglobose , strongly echinulate (fig . 106, B,
and SEM plate 18, centre: right), usually 1.8-2 .0 .tm in diameter; conidial
cha ins tending to adhere in well defined divergent columns (fig. 106, C).
Colonies upon CYA (colour plate 106 , B and b), growing fairly rapidly,
attaining a diameter of 40 mm in 14 days at room temperature (20- 25C),
consisting of a close-textured basal felt, appearing deeply velvety, zonate,
radiately furrowed , slightly raised, with undulating margin surrounded by a
very narrow fringe of submerged hyphae extending less than 1 mm beyond the
aerial growth; heavily sporing throughout in zones of different shades of bluegreen (Sguy, PI. XXIX, 425, 428), depending upon the degree of ripcness of
conidial zones; odour lacking; exudate abundantly produced, as large drops
coloured in vinaceous shades; amber-coloured pigment diffusing into the
surrounding agar; colony reverse (colour plate 106, b) in reddish brown
shades; penicilli essentially as described aboye.
Colonies upon MEA (colour plate 106, C and e) attaining a diameter of
35-45 mm in 14 days at room temperature (20-25 C), consisting of a looser
textured basal felt , deeply velvety, slightly convex; with thin, finely denticulate

margin su rro unde d by a frin gc of sub me rgcd hy ph ae extcnd ing ove)' :~ mm

bcyond the aerial growth ; heavily spo ring eve nly throughout in mou se gra y
shades (S guy, PI. XXIX, 428 ); ex udate a nd od our lacking ; colony rever so
with dark brown shades sho win g throu gh (colour plat e 106 , e); no coloured
pigm ent diffusing into the surround ing aga r; penicilli esse ntially as des cr ibed
a boye.
The spec ies description is centere d upon the typ e stra in IJFM 7031 = ATC C
42 ,239 , isolat ed by th e au tho r (C. R .) from sandy soil, M adrid , Sp ain , March
1979. It is a t present kn own only from th e typ e strain.
Penicillium murcianum is a distin ct ive spec ies characte rized by its typ ically
di varica te penicilli , with phialides abru ptly tapering to a ve ry sho rt narrowed
neck , by its restricted gr owth up on C A and MEA , a nd by its co m pa ra tively
small, stro ng ly echi nulate globose con idia . The correct placement of the
species appears, in ou r opinion , to be mo st sa tisfac to rily assigna ble to th e P.
canescens series in the Asymmetrica- divaricat a sub-section .

107. Penicilliumturolense Martnez et Ramrez: fig. 107, colou r plate 107,

and SEM plat e 18, (bottom: left).

w..u..LJ.wJ

10 prn
@

@
Q B
} @

::o::(~

l'

100

jJm

c."R. .

Fig. 107. Camera lucida

drawings of Penicillium
turolense Martnez et
Ramrez, ATCC 42,242. A,
Detailed drawings of the
penicilli . B, Mature conidia.
C, Habit sketches.

Penicillium turolense Martlnez et Ramlrez - Mycopathologia: 74. 35-49, 1981.

Colonies upon CA (colour plate 107, A a/1(1 a) growing rather rcstrictedly,
attaining a diameter of 40 mm in 14 days at room temperature (20-25C),
consisting of a close-textu red basal felt , de epi y floccose, with margin raised and
central, and submarginal areas depressed : somewhat leathery in appearance,
persistently white in margin and with a creamish cast in the centre; even
margin surrounded by a narrow fringe of submerged hyphae extending about 1
mm beyond the aerial growth; colony surface overlaid by a closely woven
network of aerial hyphae influencing greatly the colony appearance; very
scarcely and sparingly sporing, not influencing the colony appearance; exudate
limited in amount as minute droplets enmeshed in the mycelial felt ; odour
lacking; very pale amber-coloured pigment diffusing into the surrounding
agar; colony reverse (colour plate 107, a) in yellowish cream shades; conidial
structures either arising from the substratum or from trailing vegetative
hyphae, typically asymmetric and divaricate, consisting of fairly regular
verticils of 2-4 diverging metulae , mostly 10-15 11m by 2.0-3.0 11m, with
apices enlarged very slightly in a vesicle; the stipes of the conidiophores are
comparatively long when borne directly from the substratum, mostly up to 300
11m by 2.5 11m, smooth walled; phialides in verticils of 3-8 elements per
cluster, bottle shaped, mostly 9.0-12 .0 11m by 2.5-4.0 11m, abruptly tapering
to a short narrowed neck (fig. 107 , A); conidia delicately spinulose, globose,
mostly 2.5 - 3. O 11m in diameter (fig. 107, B, and SEM plate 18, bottom: left);
conidial chains at first loosely parallel, becoming tangled in age (fig. 107, C).
Colonies u pon CYA (colour pi ate 107, B and b) spreading broadly, atlaining
a diameter of 45 mm in 14 days at room temperature (20-25 C), consisting of
a close-textured basal felt, overlaid by a closely woven network of fine aerial
hyphae lending a sublanate appearance to the colony surface , conspicuously
zonate, developing whitish shades in the centre and pale cinnamon colours in
the margin and in submarginal areas; radiately sulcated by somewhat shallow
furrows and irregularly folded in the centre ; very thin, even margin
surrounded by a very narrow fringe of submerged hyphae not exceeding 1 mm
beyond the aerial growth; very poorly sporing, with conidial structures arising
from the basal felt and from the network of interlacing hyphae and ropes of
hyphae ; odour strong; suggesting ammonia; exudate limited in amount, as
minute droplets enmeshed in the mycelial felt; reverse in orange shades (colour
plate 107, b); arnber-coloured pigment diffusing into the surrounding agar;
penicilli essentially as described aboye .
Colonies upon MEA (colour pi ate 107, C and c) growing very restrictedly,
attaining a diameter of 25 mm in 14 days at room temperature (20-25 C),
consisting of a close-textured basal felt, de epi y velvety to lanate, with a thin
white crenelled margin, surrounded by a fringe of submerged hyphae

extcnd ing about 2 mm or more beyond the a crial growrh; sporing thro ug ho u t
in gra y-grce n sha des (S gu y , PI. XXIX , 4:14); co lony S U r/i l CC pla nc, slightly
raised in the ce ntre ; ex udate lack ing or lirnitcd in amo un t, as minute d ro plets
en mes hcd in th e m ycelial fe1t; odour faint , ind cfinite ; rever se in bright yellowor ange sha de s (colour plat e 107 , c); golde n yellow -coloured pigm ent di ffusin g
into th e sur ro und ing agar; penicilli essentia lly as described aboye but with
conid iophores m ostly arising from th e subs tra tu m.
The species description is ce ntere d upon th e typ e strain IJ FM 7097 = ATCC
42 ,242 , and several other strains whi ch duplica te th e preceding in all
particul ars, all isolat ed by DI' A .T. M artnez from beech leal' litter in a beech
Iorest near N ancy, France, Septem be r, 1979.
Penicillium turolense is a d ist inc tive spec ies cha rac terized by its typi call y
div aricate peni cilli , with sho rt-po inted phiali des, by its spin ulose,
compara tively sm all and globose con id ia, and by its restricted gro wth upon C A
a nd M EA , a nd mor e rapid growth up on C YA. The correc t placement 01' thc
species , in our opin ion, appears to be m ost sa tisfacto r ily ass ignable to th e P.
canescens senes.

Penicillium nigricans ser ies

OUTS TAN DlNG F EAT UR ES

Colonies gro wing rather restrictedl y upon m ost subs tra te, cons isting usu ally
a close-textured ba sal felt with surface variable , consisting some times 0 1' a
network 0 1' tra ilin g hyphae 0 1' appearing m or e 0 1' less velve ty; con id ial areas
develop in g dull, 01'ten dark , gray to olive brown shades, almos t bl ack on so rne
su bstra ta; reverse typi cally develop in g deep orange -re d, very da rk brown , to
a lmos t black colou rs.
Conid iopho res vary ing gr eatl y in length and in origin, ar ising eithe r d irectl y
from th e substratum , 0 1' as short branches from ae rial hyphae.
Peni cilli typi cally strongly divaricat e a nd asym me tric, cons isting of
div ergent m etulae arisin g a t on e 01' more leveIs and bearin g clu sters of
ph ialides producin g conidial chains in loose colum n s 01' tangled ; sim ple
peni cilli (a ppearing mon overticillat e) are not infrequently encou ntere d.
Conid ia globose to su bglobose , stro ng ly ech in u late in sorn e species, smooth
a nearly so in othe rs.

0 1'

K ey to species

Page
l. Colonies not producing scierotia or masses of thick-walled cells. Penicilli
with the divaricate character well marked; growing rather restrictedly
upon most substrata; with metulae scattered on the conidiophore, or
usually aggregated into true verticils. Colonies developing typically dull
gray, dark brown, almost black shades (scarcely showing any green).
P. nigricans series .. .. 295

A. Conidiophoro walls eoarsely roughened, at least upon malt extraet

agar.
1. Phialides eonspieuously roughened .
P. radulatum Smith . ... 308
2. Phialides smooth.
a. Conidia eonspieuously eehinulate.
P. melinii Thom . ... 303
b. Conidia smooth or nearly so.
P. raciborskii Zaleski .. . . 306
B. Conidiophore walls smooth or nearly so upon all substrata.
1. Conidia smooth walled or nearly so.
P. yarmokense Baghdadi . ... 320
2. Conidia strongly eehinulate .
a. Conidia eomparatively very large, up to 7 .tm in diameter.
P. megasporum Orpurt et Fennell .... 315
b. Conidia large, up to 5 .tm in diameter.
P. albidum Sopp . ... 299
e. Conidia smaller, not exeeeding 3.5 .tm in diameter, rarely 4.0 .tm.
P. granatense Ramrez, Berenguer et Martnez .. .. 310
3. Conidia slightly roughened, provided with two roughly parallel thin
ridges.
P. inflatum Stolk et Malla .... 312
4. Conidia eonspieuously and finely spinulose.
a. Colonies growing very restrietedly upon Czapek's agar, attaining
a diameter of 15 mm ; margin abrupto
P. ovetense Ramrez et Martnez . ... 318
b. Colonies growing less restrietedly, attaining a diameter of 25-30
mm .
1'. Colonies heavily sporing in dark brown shades.
P. nigricans (Bainierl Thom .... 297
2'. Colonies slightly sporing, white or nearly so, or in pale bluegreen shades upon malt extraet agar.
P. kapuscinskii Zaleski . ... 301

M emb ers of th e series a re regularI y isolated as com pone n ts of the soil

m ycoflora a n d ha ve been recovered from sam ples a ll ov er th e worId. They a re
p articul arI y abundant in forest soils on or ga n ic constituents in the later stages
of d ecomposition. They a re occas ionally isolated from other su bs trata su bj ect
to a ir a nd so il co n tami natio n .
T he se r ies included originally (Raper a nd Thom , 194 9) the foll owing five
sp ecies: Penicillium nigricans (Bainier) Thom , P. kapuscinskii Zaleski, P. albidum
Sopp , P. melinii Thom , a nd P. raciborskii Zaleski, to which six additional forms

", ,1 ,

have been included in the cu rre nt study, namcly: Penicillium radulaium Sm ith,
P. granatense Ramrez, Berenguer et Martfnez , P. inflatum Stolk et Malla, P.
megasporum Orpurt et Fennell , P. ovetense Rarnrez et Martnez, and P. yarmokense
Baghdadi, all developing typi cal divaricate structures in sufficient numbers,
and dull, dark brown to almost black colours to necessitate pla cing the specics
here .

108 . Penicillium nigricans (Bainier) Thom : fig. 108, colour plate 108,
and SEM plate 18, (bottom: right).

0'.,' ~,,~ {)
C@C~~'"

f.6

(D O@

y~ y
L-........J

100pm

------

Fig. 108. Camera lucida

drawings of Penicillium
nigricans Bainier ex Thom ,
CBS 221.28. A, Detailed
drawings of the penicilli. B,
Mature conidia. C, Habit
sketches.

Penicillium nigricans (Bainier) Thom - 'The Penicillia' , pp. 351-353. 1930. See also
Raper and Thom, ' A Manual of the Penicillia', pp . 325-329. 1949.

Synonymies
Penicillium echinatum Dale - In Biourge's Monographie, La Cellule. 33: 278.
1923. discussed without name as C3 by E. Dale in Ann. Mycol. 12: 42. 1914; and
named by her in Ann . Mycol. 24: 137. 1926.

hy a narrow frin ge 01' su bmcrgcd hyph ae extc nd ing a bo u t 1 mm beyond the

ucriul growth ; hcavily spo ring throu gh out in dark brown shades; exudate as
lurgc crystal clear colourless drop s; odo ur ' m ouldy'; no coloure d pigment
dilfu sin g into th e surround ing agar; colony re ve rse (colour pla te 108 , b) in
uchre to yellow sha de s; penicilli esse ntially as descr ibed aboye, with con id ial
cha ins usuall y typi call y d ivergent , becoming tangled in a ge (fig. 108 , C ).
Colonies upon MEA (co lour plat e 108 , C a nd e) a tta in ing a diameter of 30
111 111 in 14 days a t room temperatu re (20-25C) , som ewhat looser textured, not
urrowed , plane , with a very thin , cre ne lla ted margin surrounded by a wid e
frin ge 01' su b me rge d hypha e exte nd ing about 5 mm beyond th e ae rial gr owth ;
hcavily spo ring through out in deep brown shades (Sg uy, PI. XXIX, 433); no
colou red pigment diffu sin g into the surro und in g agar; colony reverse (colou r
plat e 108 , e) in dark reddish brown shade s; penicilli borne on lon g
conid iophores, up to 500 p.m in len gth, arising p rim arly frorn the substratum,
slro ngly divaricate , with con id ial chains producing fairly definite colum ns up
lo 100 p.m long.
T he species description is ce ntere d upon th e typ e stra in C BS 221.28 = IJ FM
703 7, (T , P. j anczewskii Zaleski) , isolated by K. Zaleski in Poland frorn forest
soil under con ifers, also upon st ra in AT C C 18 ,380 = C BS 744 .70 = Q M 7,297 =
I.lF M 7,247 , (T. P. nigricans val'. sulfuratum Abe), isolat ed fromjapanese soil in
1952, and upon m any othe r strains recovered from beech fores t soil in the
Pr ovince of Navarra, Sp ain , by DI' A.T. M artnez in 1977.

109. Penicillium albidum Sopp : Fig. 109, colour pIate 109, and SEM

plate 19, (top : left) .

Penicil/ium albidum Sopp - Monographie, pp . 186-187. 1912. See also Ch. Thom,
'The Penicillia', pp. 350-351. 1930; also Raper and Thom, 'A Manual of the
Penicillia', pp. 329-330. 1949.

Colonie s on C A (colou r plate 109 , A and a) atta in ing a diameter of 40 mm in

[4 days at room temperature (20- 25C) , cons isting of a close -text u re d basal
lelt, ove rlaid by a closely woven netw ork of aerial hyphae lending a lanate
a ppeara nce to the colony surface, whi ch is persist entI y whi te rosy in colour,
with a pla ne marg in surrounded by a frin ge of su bme rged hyphae exte nd ing
a bo ut 2 mm beyond the aerial grow th, a nd th e ce ntre ab ru ptIy raised in an
umbon a te pattern m or e 0 1' less sulca ted by shallow radial furrow s; poorly
spo ring, with conid ial structu res very small a nd fra gm entary, cons isting
almos t ex clusively of ph ialides bo rne sepa rate1y 0 1' in stro ng ly divergent
clusters 0 1' phialides bo rne a pica lly upon branch es (a ppearing th en
mon overticilla te) 01' upon verticils 01' m etulae (fig . 109 , A) ; con id iophores
smooth walled, rangin g from very sho rt u p to 300 p.m in length by 2 .3- 3 .0 p,m

w.wwu

10

jJm

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0) ,_'i~..,1..,:,, @r7;}.

@ tS7 \:.~ ~1

'(
?0
~

re

. (-1

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B ffft'i
<% "C,;f

.;;
~ ~~ ~
"2.1 f '~
;--~. \.. \\

Y.:fI

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.~...,

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.

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~\1
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~ l (~' yf
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100 prn

Fig. 109. Camera lucida

drawings of Penicillium
albidum Sopp, CBS 254.29.
A, Detailed drawings of the
penicilli. B, Mature conidia.
C, Habit sketches .

in diamet er ; branches , 15-25 .tm by 2 .3-3 .0 tui ; metulae , in clusters of 2 - 3

elem ents in the verticil , mo stly 12-19 .tm by 2.5 -3 .0 iuu, with apices sligh tly
enla rge d in a vesicle up to 4 .0 .tm in di ameter ; phialides in clusters of 2-10
elements in the ve rticil, stron gly div ergent , bottle sha ped, abo ut 6 - 9 .tm by
2.5 - 4. 0 .tm, abr u ptly tapering to a sho rt narrowed ne ck ; co nid ia
conspicuo u sly roughened , with roughness arrange d in more or less parallel
bands (fig. 109 , B a nd SEM plat e 19, top: left) , globose , m ostly 3.5 - 5.0 tui: in
di amet er; conid ial chains loosely parall el , bec oming tangled in age ; odou r
pronounced, 'mouldy'; exudate lacking ; colony reverse (colour plate 109 , a) in
reddish cin namo n sha des (Sgu y , PI. XLVII , 694 , 695); no coloured pigment
diffusing int o the surro und ing ag ar.
C olon ies upon CYA (colour plat e 109 , B a nd b) spread ing broadl y , a ttaining
a diamet er of 55 mm in 14 days at room temperature (20-25 C ), consisting of
a close-textured basal felt , sim ila r in texture to th at des cribed aboye, but
sulcated by m any deep radial furrows, slightly and gra d ua lly raised towards
th e ce n tre, whi ch is depressed ; thin m argin coloure d in dark brown sha de s
owing to th e heavy spo r ing surface, surrou nde d by a frin ge of submerged
hypha e exte nd ing abo ut 3- 4 mm beyond th e a erial gr owth , velvety to lanate

.' U 1

in uppcarance ; colony su rface overlaid by l thin, close ly wovcn nctw ork 01'
ae rial hypha e in fl uenc in g gre atly th e colony appcarancc ; hcavily spo ring
ihroughour in dark br own shade s, particul arly cons pic uo us in marginal areas,
a nd somc what ob scured in ce ntra l and subcentral areas by the den se net work
01' ae rial hyph ae sha di ng to wh ite rosy sha des; ex udate ab undantly produced as
yc llow-ora nge d rops ; odo u r lacking; no coloured pigment diffusin g into th e
surround ing agar; colony reverse (colour plat e 109, b) in re ddis h cinna mon
shades; penicilli esse ntially a s de scribed abo ye bu t with th e d ivaricat e cha rac ter
more strongly acce ntuated than upon the abo ve su bs tratum .
Colonies upon MEA (colour pla te 109 , C and c) attain ing a d ia mete r 01' 50
mm in 14 d ays a t room temperature (20 -25C), consisti ng 01' a looser tex tured
basal felt, th in , velve ty, cons pic uo us ly zon at e , not fur rowed , pla ne, wi th a very
ih in m argin mostly submerged, ex tend ing abou t 5 mm beyond the ae rial
gro wt h; heavily sporing through ou t in deep olive bro wn shades (Sguy, PI.
XX IX, 426 , 427); exuda te a nd odour lacki ng; no coloured pigm ent di ffusing
i nt o the su rround ing agar; colony revers e (colo u r pla te 109 , c) wit h redd ish
hrow n shades show ing th rough ; penicilli essentially as described aboye but
with coni dio p hores arising prim a rily from the su bstratu m, b eing longer, up to
;OO p.,m in len gth .
T he spec ies description is ce ntered upon strain CBS 254 .29 = CM I 40, 219 =
ATCC 10,408 = NRRL 2,034 = QM 7,48 1 = IJFM 7036 , isola ted by D . J anke
Irorn soil in 1928 , a nd m any ot her strains isolated fro m beech forest soil in the
Provin ce 01' Navarra, Sp ain , by D I'. A.T. M artnez , in 1977 .

110. Penicillium kapuscinskii Zaleski: fig. 110, colour plat e 110 , and SEM

plate 19 (top: right ).

Penicillium kapuscinskiiZaleski - Bull. Acad . Polonaise Sci. Math. Nat. Sr. B, pp.
484-485.1927 . See also Ch. Thom, 'The Penicillia' , pp . 355-356.1930; also Raper
and Thom, ' A Manual of the Penicillia', p. 330. 1949.

T he following species description , tak en from T hom's Monograph ( 1930), is

inse rte d h ere since it adequately presents th e species in q uestion, whe n first
isolated by K. Za leski in 1927. T he ty pe strain CBS 218.28, maintaine d in
artifica l cult u re since its isolation has becom e pred ominantly sterile on mos t
substrata , a nd the penicilli pred ominantly sim ple (appea ring monove rticillate) , not showing a ny longer the typ ical cu ltural characteristics origi nally
describe d by the autho r, alt ho ugh still showing ce rta in characteristic det ail s 01'
rnorphology:
'Type strain growing fairly well about 20C, very slightly at 30C, or aboye;
colonies on Czapek's solution agar about 20C, restr ictedly growing 15-20 mm in
diameter in 12-14 days, appearing velvety but with very thin felt at base, buckled

S)~::~~ ~..
(ii

,b '(

~i"

t1I;

f}

I.--.J

100J,lm

tW

fR. .

Fig. 110. Camera /,

drawings of Penie"
kapuscinskii Zalesld
218.28. A, Detaile d
of the penicilli. B,
conidia . e, Habit s

and radiately wrinkled sometimes in quadrants, with eonidial area gray-gr

olive gray (Ridgway, PI. L1) at first, beeoming drab in age (brownish drab ,
PI. XLV) with myeelial hyphae, and marginal 3 mm white, showing s
hyphae reaehing beyond the submerged myeelium and re-entering the su
hyphae very delieate; reverse in pale orange shades; drops , small, colour
distributed over the eonidial area; eonidiophores variable in length, arising
substratum or from aerial hyphae; penicilli partly monoverticillate, rnostlv
showing one-braneh, a group of sterigmata (phialides) around the main a
first septum; branehes (or metulae?) 12-15 .tm long; sterigmata (phialidesl
by 2.0-2.5 tm with eonidial ehains divergent; eonidia 2.5-3.0 tm, witl1
roughening or spinulosity seen only under oil immersion .'

Our not es follow :

Colonies upon CA (co lour pl at e 110, A a nd a ) a ttain ing a diamet er O
in 14 d ays at ro om temperature (20 - 25 C) , co nsisting of a tougl
textured basal felt , a ppear ing typicall y ve lve ty to finel y lanate owing t
closely wov en network of fine white veget ative hyphae overla ying th

i;~;~.

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;.) : i

~r;;} \~~)

f2 r
1

l-,.-J

100 pm

'l Y

iR..

Fig. 110. Camera lucida

drawings of Penicillium
kap uscinskii Zaleski, CBS
218.28. A , Detailed draw ings
of the penicilli. S, Mature
conidia. e, Habit sketches .

and radiately wrinkled sometimes in quadrants, with conidial area gray-green, deep
olive gray (Ridgway, PI. L1) at fi rst , becom ing drab in age (brownish drab , Ridgway,
PI. XLV) with mycelial hyphae , and marginal 3 mm white, showing stolon-Iike
hyphae reaching beyond the submerged mycelium and re-entering t he substratum;
hyphae very delicate ; reverse in pale orange shades; drops, small, colourless, well
distributed over the conidial area; conidiophores variable in length, arising from the
subst ratum or from aerial hyphae ; penicilli partly monoverticillate, mostly variously
showing one -branch, a group of sterigmata (phialides) around the main axis at the
first septum; branches (or metu lae?) 12-15 ,tm long; sterigmata (phialides) 8-10 ,tm
by 2.0-2.5 ,tm with conidial chains divergent; conidia 2.5-3.0 ,tm, with delicate
roughening or spinulosity seen only under oil immers ion .'

Our notes follow:

Colonie s upon CA (co lour pla te 110 , A and a) attaining a diamete r of20 m m
in 14 days at room temperature (20-25 C), consisting of a tough , close textured basal felt , appearing typ ically velvety to finely la na te owing to a thi n,
close ly woven network of fine white vegetative hyphae overiaying the colony

su r facc

and

inllu cncin g grca t ly th e co lo n y a p pca ra ncc;

m a rgin

plane ,

su rro u ndc d b y a narrow frin ge 01' su b me rge d hyph ae ex te nd ing a bou t 1 mm

beyond the ae rial grow th ; co lon y surf acc a hru ptly raised in sub marg ina l arca
ami c1epressecl in a cra te ri for m pattern in th e ce n tre, sulca ted with few radi al
furrow s ; ve ry poorI y spo ring, with co lony su rface remaining per sistently white ;
exudate a nd odo u r lacking ; no colou re d pigme n t diffu sin g into th e
su rro und ing agar; colony reve rse (co lour pla te 110 , a) in ligh t cream shades
a nd stro ngly pulling th e agar a way fro m th e cult ure di sh ; pen icilli (fig. 110 , A)
with an ac tual majority a p pearing monoverticillat e, with co ni d iopho res
smooth walled a nd with numerou s septa, bearing few verticils of metulae ,
mostIy 6 -8 utt: by 2.8 -3. 0 tm ; phi alides in clus te rs of 2-6 eleme n ts in the
ve rtici l, usu all y 6- 10 ust: by 2.5 -3.0 tuu , ta pering to a sh ort narrowed neck ;
conidia globose de lica tely rou gh en ed (fig. 110 , B, and SEM plat e 19, top :
ri ght ), mostIy 2.5 -3 .0 tm in d iameter; conid ia l cha ins sh ort, loosely parallel
(fig. 110, C).
Colon ies upon C YA (co lo u r plat e 110 , B a nd b) attain ing a di amet er of 30
mm in 14 d ays a t roo m tempera ture (20 -25 C) , co ns isting 01' a close-textu re d
basal felt , a p pearing 'wet', translu cent , stro ng ly sulcated by m any radi al
fu rrows in m argin al and su bmarg inal areas, with cen tre strong ly raised and
irregularIy fold ed in a cra te rifor m pattern , pulling th e agar so stro n gly as to
p roduce cre vices in it and cracking th e colony su rface in the ce n tre; m argin
largely su bmerge d; al mo st ste rile ; ex uda te a nd odo ur lacking; yellow-coloured
pigme nt slowly diffusin g into the surro u nd ing agar; co lony reverse (co lour
pl at e 110 , b) uncoloured.
Colonies upon M EA (colour plat e 110 , C a nd c) a tta in ing a di amet er of2 0
m m in 14 d ays at room temperature (20-25C), cons isting of a typi call y
velve ty felt , co m paratively thin , with a wide f1at m argin surro u nde d by a wide
fringe of su b me rge d hyphae ex te nd ing a bo ut 5 mm 0 1' m ore beyond th e aerial
growth; cen tral a nd su bce n tral areas somewhat raised a nd radi at ely wrinkl ed
with cen tre d epressed ; m edium spo ring in ce n tral areas in pale blue-gray
shades (Sgu y . PI. XXXI , 455) ; exud ate a nd od ou r lackin g; no coloured
pigmen t d iffu sin g in to th e surrou nd ing agar; colony reverse (co lou r pla te 110 ,
c) in pal e ye llow shade s; penicilli esse n tia lly as d escribed a boye .
T he species description is ce n te red upon th e typ e strain C BS 2 18. 28 = C M I
40,56 7 = ATCC 10 ,4 57 = NR RL 2, 14 7 = IF a 77, 674 = Q M 7,588 = IJFM 7035 ,
isolat ed by K. Za leski in Poland , from forest soil under co nifers, in 1927 .
The species is ret ained to include occa sion al m embers of th e Penicillium
nigricans se ries whi ch produ ce globose, delicatel y rou ghened con id ia, and which
are ofte n less deepl y colou re d in th e obverse a nd rever se th an th e typical
re p rese n ta tives of P. nigricans itself.

111. Penicillium melinii Thorn: fig. 111, colour pIate 111, and SEM
plat e 19, (centre : Ieft).

':1

"

"

"

:'

"
""

"

"

"

'0 0

"

10 jirn

"

o'

",,
,,

(f)4iJ@

f&)';\~
'&'

Ylr
e

.F?,

'-----J

100)Jm

Fig. 111. Camera lucida

drawinqs of Penicillium
melinii Thom, CBS 218.30.
A, Detailed drawings of the
penicilli. B, Mature conidia.
C, Habit sketches.

Penicillium melinii Thom: In 'The Penicillia', p. 273. 1930; also Raper and Thorn,
'A Manual of the Penicillia', pp . 331-332 , 1949.

Colonies upon CA (colour plate 111 , A and a) growing restrictedly,

attaining a diameter of20-25 mm in 14 days at room temperature (20-25C),
consisting of a tough , close-textured basal felt, appearing typically velvety,
abruptly raised and strongly sulcated by many radial furrows near the
marginal areas and irregularly folded in th e centre and sometimes depressed in
a crateriform pattern; growing margin very thin , surrounded by a narrow
fringe of submerged hyphae extending less than 1 mm beyond the aerial
growth; heavily sporing throughout in olive brown shades; exudate lacking;
odour faint, ' mouldy '; no coloured pigment diffusing into the surrounding
agar; colony reverse (colour plate 111, a) uncoloured but with the olive brown
shades of the obver se showing through ; colony strongly pul1ing the agar away
from the culture dish ; penicil1i variable in pattern, commonly simple
(appearing monoverticil1ate) but frequently consisting of terminal groups of
strongly diverging branches or metulae (fig . 111 , A), irregularly arranged,

borne cithcr at thc apex 01' thc main axis 01' arising from lowcr nod es ;
conidiophores with walls strongly warty , sho rt , usu ally less than 100 Jtm in
Icn gth by 2.8-4.0 Jtm in diameter; metulae commonly 10-30 Jtm by 3.0 Jtm;
phialides comparatively large , in clusters al' 3-10 elements in the verticil,
mostly 9-15 Jtm by 3.0-5 .0 usu , conspicuously tapering to a long narrowcd
ncck, and tending to diverge; conidia globose, comparatively large, most1y
5- 6 Jtm in diameter, strongly echinulate , suggestive al' the chestnut fruit when
ob served under the light microscope (fig . 111, B) showing a strongly thorny
surface when observed under the SEM (plate 19, centre : left); con idial chains
111' to 100 tm in length, com monly divergent (fig . 111, C) .
Colonies upon CYA (colour plate 111, B and b) growing more rapidly than
a hoye, attaining a diameter 01' 40 mm in 14 days at room temperaturc
(20 - 25 C ) , similar in texture to thos e grown upon CA, but not abrupt1y
raised, and regularly sulcated by numerous radial furrows in subce ntral areas ,
with the centre irregularly folded in a more al' less cra te riform pattern, with a
(hin white margin, becoming light buff in age, surrounded by a narrow fringe
01' subm erged hyphae extending about 1 mm beyond the aerial growth; very
heavil y sporing throughout in dark olive brown shades, with the brown colour
pr edorninant , except in central areas whi ch remain persistently white;
cxuda te lacking; odour faint , 'mouldy'; no coloured pigment diffusing into the
surroun ding agar; colony reverse (colour plate 111 , b) apparently colourless
but with de ep brown shades showing through ; penicilli essentially as described
aboy e.
Colonies u pon MEA (colour pIate 111, C and c) att a ining a diameter 01' 50
mm in 14 days at room temperature (20-25C), consisting al' a looser textured
basal felt, appearing typically velv ety , not furrowed , plane , with centre
umbonate ; very thin margin largely submerged, exten ding about 5 mm al'
m ore beyond the aerial growth ; exuda te lacking; odour not pronounced ,
' mouldy '; no colou red pigment diffusing into the surrounding aga r ; colony
reverse (colour plate 111, e) in reddish brown shades with centre in deeper
brown colours; penicilli essentially as described above but with the stipes 01' the
conidiophores more coarsely roughened .
The spe cies descriptions is centered upon the type strain CBS 218.30 = CMI
40 ,216 = ATCC 10,469 = NRRL 2,041 = IFO 7,675 = QM 799 = IJFM 7053,
isolated by DI' Elias Melin from forest soil in 1929 in th e USA.
Penicillium melinii was assigned by Raper and Thom (1949) to th e
Asymmetrica-divaricata, sin ce an appreciable number 01' penicilli showed the
typi cal divaricate character 01' this group . Many , if not an actual majority , al'
th e penicilli , however, appear simple and a possible relationship to the
Monoverticillat a should not be overlooked . However, the globose, strongly
echinulate conidia and th e production al' deep brown colours with the ripening
a l' con idia are strikin gly suggestive al' th e Penicillium nigricans series , and

parti cula rly 01' P. nigricans itself lt dilfers I'I'O Ill P. nigricaus in its slro ng ly warty
co nid iop ho rcs .

112. Penicillium raciborskii Zaleski: fig. 11 2, colour plate 112 , and SEM

pIate 19, (centre: right).

4b"c~

r.l o

:
'
~l ~:~! ~ .
L - -jrrn
-J
100

"

Fig. 112. Camera lucida

drawings of Penicillium
raciborskii Zaleski, CBS
224.28. A, Detailed drawings
of th e penicill i. B, Mature
conidia. C, Habit sketches.
D, Penicilli on malt extract
agar .

Penici/lium raciborskii Zaleski - Bull. Acad. Polonaise Sci . Math. Nat. Sr. B, pp.
454-455.1927 . See also Thom , 'The Penicillia', pp. 318-319.1930 ; also Raper and
Thom, ' A Manual of the Penicillia', pp . 333-334. 1949.

Coloni es upon CA (co lour p lat e 112 , A and a) attaini ng a di ameter 01' 45 m m
in 14 days at room tempcrature (20 - 25 C) , consistin g 01' a clos e-textured basal
felt overlaid by a dense , clos ely wov en network 01' aerial hyphae lend ing a
la na te appearance to th e colon y surface , stron gly furrowed in an irregular
pattern , rem aining pe rsi stently white th rou ghou t , w ith th e exce p tion 01' the
inter-colony margins where con id ial structures a re a bu ndan tly produ ced in
vcry pale gray-grcen sh ades (Sguy, P I. XXX , 44 5) ; exu da te lim ited in

a mou nt , as minute dropl ets c n me shc d in thc my cclial felt ; od ou r lackin g or

slightly 'mouldy ': no colou rcd pigmcnt difusin g into the su rrou nd ing agar;
culony reverso (colour plate 112 , a) in dull pcach, somctimcs dcvcloping
vinaceous shades; pcnicilli usually strongly and typicall y divaricate (lig. 112,
A) . with con id iop hores slender, usually 1.8-2 .0 11m in diameter arising eithe r
lrom th e substratum 01' as branch es frorn aer ial hyphae , variable in len gth,
1'1 I'c1y exceeding 200 11m in length , with walls usually appearing smooth;
pcni cilli irregularly branched, with branches and metulae variable in size a nd
olt cn arising at th e same level ; branches usually 30- 100 11m in length by 211m
in diamet er ; m etulae in whorls 01' 2 -4 01' more elements in the verticil,
rom rnonly 17-25 11m b y 1.5-2. 5 11m, with apices conspicuou sly enlarged in a
vcsicle up to 4 .0 11m in di ameter; phialides com para tive ly small , bottle shaped,
usually cro wde d in clusters 01' 3- 10 01' more elements in the verticil , closely
pl ra llel, a bou t 7-9 11m by 2.3 - 3 .0 11m, tapering to a shor t narrowed neck ;
ro nid ia com p a ratively very small , thin walled, a bo u t 1.5-2.0 11m in di ameter,
with walls appearing smooth when observed under thelight mi croscope (fig.
11 2 , B) 01' delicately granular, when observed under the SEM plate 19 , ce n tre:
I'i g-ht).
Colon ies upon CYA (colour pl ate 112 , B and b) attaining a di amet er 01' 50
111m in 14 days a t room temperature (20 -25 C), sim ila r in texture to th e aboye
ilcscr ibe d colonies but with surfa ce regularly sulcated with many radial furrows
iu subcen tral a nd marginal areas, and with ce n tre irregularly fold ed , sligh tly
rai sed ; with thin , ev en margin su r rou nde d by a narrow fringe 01' submerged
h yph ae exte n d ing abou t 1 mm be yond the ae ri a l gr owth; heavily , sporing,
particul arly in ce n tral areas a nd alon g the intercolony margins in d eep gray~ I'ce n shades (S guy , PI. XXXI , 461) ; exu da te ab u n dan tly produced as cry sta l
clea r . colou rless drops along th e furrow s; od ou r lacking ; no colou red pigment
dilfu sing into the surrounding aga r ; colony reverse (col our plat e 112, b) in dull
peach to reddish cinnamon colou r s; peni cilli essen tially as descr ibed a boye.
Colon ies upon MEA (colour plate 112 , C a n d c) spreading broadly,
a ttain in g a di amet er 01'60- 70 mm in 14 days a t room temperature (20 - 25 C ),
cons istin g 01' a loo ser textured basal felt, co m p a ra tively thin, not furrowed ,
plane , with su rface m arked by a loose overgrowth 01' floccul ent hypha e; with
thin, eve n m argin su rrou n de d by a wid e fringe 01' submerged hyphae
cxte nd in g about 5 mm be yond th e a eri al growth ; heavily sp oring throughout in
dull gr ay-green sh ad es (Sguy , PI. XXIX , 434 ); ex udate a n d odour lacking ;
110 colou red pigment diffusing into the su r rou nd ing a ga r; colony reverse
(colour plat e 112 , e) in reddish brown in ce n tral a re as to yellowish brown
sha des in m argin ; penicilli usually essen tia lly a s d escribed ab oye but oft en
larger and with conidiophores conspicuously gran u la r (fig. 112 , D) , a n d with
co n id ial cha ins forming 2, 3, 01' m ore fairl y d efinite colu m ns up to 100 11m in
len gth (fig. 112, C ) .

I' he spe cies dcscript ion is ee nte re d upon the typ e strain c ns 224. 28 - e M I
40 ,56B - .vrc o 1O ,4B8 = NRR L 2,150 - I Fa 7,67 6 - IJ FM 7052, isolat ed by K.
Zaleski from forest soil under eonifc rs, in Pola nd , in 1927. 1t is kn own only
from th e typ e strain.

113 . Penicillium radulatum Smith: fig. 113 , colou r plate 113, and SEM
plat e 19 , (bottom : left) .

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Fig. 113. Camera lucida

drawings of Penicil/ium
radulatum Smith , CSS
280.58. A, Detailed drawings
of the penicilli. S, Mature
con idia. e, Habit sketches.

Penici/lium radulatum Smith - Trans . Brit. Myeol. Soe. 40: 484-485. 1957. See also
M.M. Kulik, 'A Compilation of Deseriptions of New Penici/lium Speeies' . Agrie .
Handbook No. 351. Agrie. Res. Ser . U.S. Dep . Agrie . pp. 68-69. 1968.
Smith's original diagnosis follows, sinee this adequately presents the species in
question when first isolated :
' Colonies on Czapek agar spreading broadly, circular with regular margins, dark
bluish green with narrow paler edge during the rapidly growing period, beeoming
very dark smoky blue -green , smooth , velvety; respiration drops (exudate) minute,

yollow; reverse somewhat greenish yellow, with the colour diffusing throughout the
r; on malt agar or potato agar spreading fairly rapidly, 23-30 mm in 9 days at
/loe , dark grayish blue -green with paler bluish margin, velvety; drops minute,
yollow; reverse yellow to orange, with the colour diffusing slightly; penicilli
mnrkedly divaricate with the branches loosely divergent, usually with a terminal
luster of metulae and phialides mixed, and with a series of branches from lower
nodes, bearing metulae and phialides or phialides alone; conidiophores arising from
tho substrate, very rough, fairly long and 2.2-2.5 .tm in diameter; rami and metulae
ou on indistinguishable and very variable in length, rough; phialides rough, fairly
ihruptlv pointed, 8-9 by 2-2.4 .tm; conidia globose, spinulose, mostly 2.8-3.2 .tm
111 diameter, but occasionally up to 4.5 .tm, in chains which are usually divergent at
Ilrst, becoming tangled, but are sometimes more or less parallel or even in loosely
olumnar rnasses.'

O u r notes follow :
C olonies upon CA (colour plat e 113, A a nd a) growing rather restrictedl y ,
Illaining a diameter of 35 mm in 14 days at room temperature (20-25 C) ,
I'I lIIs isting of a close-textu red basal felt , typi cally velvety, not furrowed , plane ,
with thin undulating m argin su rrou nded by a fringe of submerged hyphae
xtc n d in g ab out 1 mm be yond th e aerial gr owth ; heavily sporing throughout in
d ilfcrent shades of blue-gr een, depending on th e degree of rip eness of conid ia ,
IlCing darker in m arginal area s (Sgu y , PI. XXX, 447) ; exudat e abundantly
produced , particul arly in th e colony cen tre, as deep or ange-coloured dropl ets;
odo ur lacking; deep yellow-oran ge-c oloure d pi gm ent diffusin g into th e
surr ounding agar ; colony reverse (colour pIat e 113 , a ) in or ange shades;
conidial structu res essentia lly as des cribed aboye (Smith 's original diagnosis)
( Iig. 113, A) , with conid iophores cons picuo us ly warty and phialides definitely
rou gh; con id ia are strongly rou ghened, appearing spinulose wh en obs erved
u nd er th e light microscop e (fig. 113, B) but th ey show stro ng echin ula tion
whe n ob serv ed under th e SEM (pla te 19, bottom : left) , mostly 3. 0 - 4 .0 /lm in
di ameter: conidial chains are loosely div ergent (fig . 113, C ).
C olonies upon CYA (colour plate 113, B a nd b) growing m or e rapidly th an
I he preceding, a ttaining a diamet er of 50 mm in 14 days at room temperature
(2- 25 C ), con sisting of a close -tex ture d basal felt , velvet y, with m argin
plane , not furrowed , but with cen tra l area deeply radiately furrowed and
slightly rais ed , cons picuo us ly zonate; medium spo r ing in pale drab sha de s
(S gu y , PI. XL, 590); exudate abu nda ntly produced as deep reddish or angecoloured large drops ; odour lacking; reddish brown-coloured pigm ent d iffusin g
i nto th e su rro und ing a ga r ; colony revers e (colou r plate 113 , b) in deep redhr own shades; penicilli esse ntially as described abo ye .
C olonies u pon M EA (colour plat e 113, C a nd c) spreading broadly,
a ttain ing a di ameter of 50 mm in 14 days a t room temperature (20-25 C),

cons isting 0 1' a looser tcxtu red busal Iclt , vclv ct y , plnn c , more 0 1' less zo na te in
m argin al a rca, not Iurrow ed , wi th thin m a rg in surrounclcd by l wid c frin ge 0 1'
su b merged hyphae extendi ng 5 m m 0 1' m ore bcyond the aer ial gro wt h; ve ry
h eavily spo r ing throu gh ou t in ve ry dark gr cen , almost b lac k shadcs (Sgu y , PI.
XXX , 44 6); exu date a nd odo ur la ckin g 0 1' faint , a ro matic; no colou rcd
pi gment diffusing into th e surro un d in g agar; colony reverse (colour platc 113 ,
c) in yellowish shades with a green ca st show ing throu gh ; penicilli essen tially as
described aboye .
The species description is ce ntere d upon the ty pe strain CBS 250 .58 = C M I
71 ,624 = ATCC 18 ,383 = Q M 7, 526 = IJFM 7039 , isolat ed by M r G .W .Sewe ll,
01' th e R oyal H ollow ay College , from Calluna hea thl and soil from Puttenharn
Com mo n , Su r rey, U K, M arch , 1954 . Several strains 01' Penicillium radulatum
we re recove red by G. W .Sewell from hea thl and soils in N ewb or ough ,
A n glesey, in Godlingsto n H eath a n d Pl at eau H eath , D orset , in K ell in g H eath ,
Norfolk, a nd in Rhossilli D owns, Gl amorganshi re , Wales.
Sp ecies 0 1' Penicillium in wh ich b oth co n id iophores an d phialides are
d efinitely rou gh a re few in number; a mo ng them : P. argillaceum St olk , Evans el
Nil sson , P. pallidum Smith, P. putterilli T hom , a n d P. namy slowskii Zaleski have
smooth wall ed con id ia , but no other spe cies is kn own so far , except P. radulatum
Smith , h a ving all pa rt s of the penicillus , in c1uding th e phialides, as well as th e
co n id ia, co nsp icuo us ly as pe ru la te .

114. Penicillium granatense R amrez , M artnez et Ber en gu er: fig. 114,

colou r pl at e 114 , and SEM plat e 19 , (bo ttom : right).
Penicillium grana tense Ramrez, Martnez et Berenguer - Mycopathologia. 72.
27-34 . 1980.
Colon ies upon C A (colour plat e 114 , A a n d a) gro w ing ra th er res tr icte dly ,
a tta in ing a di ameter 01' 30 mm in 14 da ys a t room temperature (20 -25 C ),
consisting 01' a tough , close -tex tured b asal felt , a ppear ing deeply velvety to
su bla nate , zo nate, with undulating m argin en tire; colony su rface d eeply
radiat ely furrowed in m arginal a reas a nd irregularly folded a n d rais ed towa rds
the ce ntre in a crate r ifor m pat tern ; hea vily spor in g in zo nes in blue-green to
blue-gra y shad es (S guy , PI. X XX , 424, a n d PI. XL , 449, respectively);
ex u date crystal c1ear , abundantly produ ced a s small colou rless d ropl et s; od ou r
' mo uldy' ; co lon y rev er se (co lou r plat e 114 , a) in re dd ish cin n a m on sh ades
(S gu y , PI. XLVII, 698) ; reddish brown-coloured pigment diffusin g into the
su rro u nd ing agar ; penicilli ty pically strongly di varica te asym metr ical,
irregularly branch ed , a ppear ing sometimes sim ple (fig . 114 , A ), with
con id iophores usu all y arising as sho rt branch es from tr ailing h yphae , srn ooth
walled, co m monly not exceed ing 100 .tm in length by 2.0- 2.5 .tm in di amet er,

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drawings of Penicillium
grana tense Ramrez,
Martinez et Berenguer,
A TCC 42,233. A , Detailed
drawings of the pen icilli. B,
Mature conid ia. C, Habit
sketches .

w it h ap ices somewhat enlarged in a ves icle up to 3. 0 /lm in diamet er; m etulae

I'('w in th e ve r ticil, gen erally tw o eleme nts, stro ng ly di vergent , mostl y 10-1 5
{I n in len gth , so me time s up to 25 tui: long, by 2.0 -3 .0 /lm in diameter , usu all y
nI' un equal len gth ; phialides bottl e shaped, in clusters of 3 - 8 ele me n ts in th e
vert icil, di ver gen t , m ostly 7. 0 -9 .0 tuti by 2.0 - 3 .0 tun , tapering to a very shor t
narrowed neck ; conidia stro ng ly echin ula te (fig. 114 , B a nd SEM plat e 19 ,
hot tom : ri gh t) , glob ose , usuall y 3 -4 /lm in diamet er ; coni d ia l cha ins
p rodu cin g loose colu m ns (fig . 114 , C).
Co lon ies upon C YA (co lour pla te 114 , B a nd b) sp read ing m ore widely th an
un the preced ing su bst ra tu m, attaining a diamet er of 50 mm in 14 days at
room temper ature (20- 25 C ), sim ila r in texture a nd colony a ppeara nce to
I hose gro wn upon C A, but m ore deepl y radiately furrow ed , with ce n tre
irreg ula rly fold ed a nd raised , with white eve n margin su rrou nded by a narrow
fringe of subm erged hyphae , ex te nd ing less th an 1 mm beyond gro wth; co lony
su rface cons p icuou sly zanate , owing to th e va r ious degr ees of ripen ess of
co nid ial zo nes ; heavily spo ri ng throug ho u t in blue-gray sh ades (Sguy, PI.
X X X , 449) a t m aturity, wh ich sta rt s in m argin al zo nes towa rds th e cen tr e ;
eolony su rface ove rla id by a th in closely inter woven network of fine ve ge ta tive

whit e hyph ae partiall y obsc uring th e co nid ia l struc tu res, es pcc ia lly in the
ce n tre; cx udate a bun d a ntly produ ced , as uncoloured, crys tal c1ear drops;
odo ur fa int , indefinite, 0 1' la cking; very pale rcddish br own pigment diffusing
into th e surrou nd ing a ga r; co lony revers e (colour plate 114 , b) in pale rcddish
cin namon shade s; penicil1i esse ntially as described aboye.
C olonies upon M EA (co lour plat e 114 , C and c) gro wing restrictedly,
attain ing a di ameter of 35 mm in 14 days at room temperature (20 - 25 C ),
co ns isting of a looser textured bas al felt , not furrow ed , plane , typically velvety,
with white , finel y cre ne lla ted m argin with a frin ge of submerged h yphac
exte nd ing a bo u t 2 mm beyond th e ae rial gro wth ; very heavily spo ring
th rou gh out in very dark blue-brown, almost black shade s (S guy , PI. XXXIV,
506) ; no colou re d pi gm ent diffusing into th e surrounding agar; colon y reverse
(colour plate 114 , c) developing reddish brown shades; exudate la cking; odou r
aromatic ; penicilli esse n tially as described a boye .
The species description is ce nte re d upon th e typ e stra in IJFM 5965 = ATCC
42,233 , isolated b y th e author from th e a ir of Madrid , Spain , Apri11979 . It is
ac tually kn own only from th e typ e strain .
Penicillium granatense is a distinctive species c1e arly characte rized b y its
strongly di varicate pen icilli , th e stro ngly ech in ula te conid ia, and by the very
d ark, almost black colou rs develop ed b y th e colon ies gro w n on MEA,
suggestiv e of th e P. nigricans se ries, to which th e species appears to be most
sa tisfacto rily a ssignable.

115. Penicillium inflatum Stolk et Malla: fig. 115, colour plate 115, and

SEM plate 20, (top: left).

Penicillium inflatum Stolk et Malla - Persoonia. 6: 197-200. 1971.
Stolk and Malla's original diagnosis follows, since this adequately presents the
species in question when first isolated:
'Colonies on Czapek agar growing very restrictedly, attaining a diameter of about
10 mm in two weeks at 25C ; central areas generally raised, mostly somewhat
wrinkled and buckled, consisting of a tough, c1ose-textured felt of hyphae, lightly
sporing. Surface of central areas almost white, the very narrow plane marginal
areas showing pale olivaceous to buff shades near deep olive-buff (Ridgway, PI. XL;
Rayner, 1970,21" 'b). Exudate lacking or in Iimited amount and then collecting in
very small pale orange droplets. Margin abrupto Reverse of the colonies pinkish to
orange-brown from Iight pinkish to cinnamon (Ridgway, PI. XXIX; Rayner 15" d to
15"), sometimes showing a small brown zone .
Colonies on malt agar growing slightly more rapidly than those on Czapek agar,
attaining a diameter of 25 mm in two weeks at 25C; plane, slightly raised and
furrowed in the centre of the colonies, velvety at the margin, becoming looser-

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drawings of Penicillium
inflatum Stolk et Malla, CSS
682.70. A, Detailed drawings
of the penicilli. B, Mature
conidia. e, Habit sketches.

textured in the submarginal and central areas, consisting of a thin network of

trailing hyphae; slightly zonate in age; sporulating well. Surface showing grayish to
olivaceous shades ranging from light grayish olive to grayish olive (Ridgway, PI.
XLVI; Rayner, 21" b to 21"). Reverse of the colonies yellowish brown.
Vegetative hyphae hyaline, branched, 1.5-4 .tm in diameter, the submerged
hyphae often showing inflations up to 8 .tm in diameter. On malt agar conidiophores
arising at the margin from submerged hyphae, terminating in penicilli, ascending at
first, but increasing length becoming procumbent and developing many short,
seprate side-branches with penicilli; in submarginal and central areas
conidiophores arising as short branches from aerial hyphae .
Conidiophores hyaline, smooth- and thin-walled, varying greatly in length,
ranging from very short when arising from aerial hyphae to 500 .tm or even longer,
by 1.5-3 .tm in diameter; enlarged at the apex to form a vesicula-like structure
3.5-6 .tm in diameter. Branches irregularly arranged along the conidiophores,
measuring 5-30 by 1.5-2.5 .tm, swollen at the apex to 3-4 .tm in diameter. Penicilli
biverticillate-divaricate, consisting of 2-10 mostly strongly diverging metulae
bearing phialides; when many metulae are present they may be somewhat radiately

arranged, glving the peni cillus and Asporgillus-lIke appearance; also reduc ed forms
consisting of only two metulae and monoverticillate penicilli occur; all elements of
the penicillus are hyaline and smooth-walled . Metulae developing successively on
the vesicula-like apex of the conidiophore, occasionally occurring also on its
subterminal portion; c1ub-shaped, rarely branched, 5-10 ,tm long, narrow at the
base, 1.5-2.2 ,tm in diameter, broadening gradually to the swollen apex, 3-6 ,tm
diameter. Phialides developing successively on the swollen apex, occasionally also
on the subterminal portion of the metula , occurring in small c1usters of 3-8, slightly
diverging, 5.5-7.5 ,tm by 2-3 ,tm, less wide at the base, narrowing at the top
abruptly to a small, but definite conidium-bearing tip (neck) (0.5-1 ,tm long).
Conidia brownish, globose to subglobose, slightly roughened, provided with mostly
two roughly parallel, very thin bands, 1.7-2.5 ,tm in diameter. Conidial chains
diverging, somewhat tangled, short, up to 60 ,tm long.
Minimum temperature 5C , optimum temperature 20-25C, maximum
temperature 30C .
Because of the swollen apices of the conidiophores and its many strongly
diverging metulae, which in large heads may be radiate, the habit of this new
species suggests a species of Aspergillus. It differs, however, from this genus in
some important characters. No defin ite foot-cells are present, the metutae do not
develop simultaneously on the apex of the conidiophore as is known to be the case
in true Aspergillus, but they develop successively. Moreover the conidiophore wall
is thin, whereas most Aspergillus have thick walls. The species shows much closer
affinities to the genus Penicillum, especially to the P. nigricans series. Its cultures
are grayish to olivaceous-buff like in this series, the conidia are brownish coloured,
while the structure of the colony agrees completely with P. nigricans (Bainier)
Thom.'

The origina l diagn osis presented aboye esse n tially con forms with our own
ob servations of th e typ e strain in con nection with th e present study. Our
additional cult ural observatio ns follow:
C olon ies upon CA (co lour piate 115, A and a) growing ve ry restrictedl y,
attaining a diameter of 10 mm in 14 days a t roo m temperature (20 -25C),
co nsisting of a tough , cio se-t extured ba sal felt , a ppearing velv et y , ove rla id by a
ciosely woven network of white vegetative h yphae influencing greatly th e colony
appearance , colony surface strongly convex, abo ut 3 mm deep ; ve ry slightly
spo ring, remaining persistently whi te; ex udate lacking; odo ur non e; no
colou red pigment diffu sin g into th e surroundin g a ga r; colony reverse (colou r
plat e 115, a ) in pinkish cin nam on shades .
Colonies upon CYA (co lour plate 115, B and b) growing more rapidly th an
upon C A, att ai n ing a di amet er of 25 mm in 14 days a t roo m temper ature
(20 - 25C) , cons isting of a close-texture d basal felt , ap pearing deepl y velvety
to lanate , stro ng ly rad iately wrinkled and grad ually rai sed tow ards the

J I ",}

ub ccntral arca , thcn a bru pt ly raised in th e ce ntre ami dcprcsscd in a

r ru te ri fo rm pattern: colony su r facc entire ly ove rla id by a denscly woven

nct work ol' white vegetati vo hyphae, inlluen cin g thc colony appe a ra nce a nd
OI.lsl:lII'ing com pletcly th e otherwise medium spor ing areas, whi ch remain
pcrsis ic ntly sno w white in colou r; exuda re abundantly produced as minute ,
crystul clear, colourless droplcts enmes he d in th e m yceli al felt ; odo ur lacking ;
pille arnber-colou re d pigment d iffus ing into th e surro und ing agar ; colony
n -verse in ye llow-cinnamo n shades (colour plat e 115 , b).
C olon ies upon MEA (colour plate 115, C and e) attaini ng a diameter 0 1' 30
111 111 in 14 days a t room temperature (20-25 C) , cons isting 0 1' looser textured
hnsal felt , not furrowed , thin, plane , with cen tre umbonate , cons picuo usly
zu narc : very thin m argin largeIy su bme rge d, ex te nd ing abo u t 10 mm beyond
11)(: ac rial growth; spo ring abundantly in pale brown to buff shades (Sguy,
X I,VII, 705); exuda te and odour lacking; no coloured pigment di ffusin g int o
1111: surro u nd ing agar; colony revers e (colour pia te 115, e) in ye llow ish b rown
ru lours ; penicilli esse ntially as described in th e origi na l d iagn osis (fig. 115 , A
und C); conid ia (fig. 115 , B) essentially a s described aboye whe n obse rve d
uu cle r th e light m icros cop e , but showing well-d efin ed spirally ridges wh en
oh scrved under th e SEM (pla te 20 , top : left) .
The species description is centered upon th e typ e stra in C BS 682 .70 = IJFM
7()(j2, isolat ed with four addi tional strains from the root surface 0 1' Picea excelsa
Link , in th e L venholm di strict , ]utland , by D .S . M alla , Sta tens For stli ge
I'orsgsva esen , Springforbi , D enmark , in 1970 , a nd in the H rsh olm di strict
in the northern part 01' Sjaelland , D enmark , in 1969 . An add itional stra in wa s
ixolated by R.A . Sa mso n, C .B. S. , Ba arn , The N etherlands, from forest soil
lindel' Quercus robur L. , in Spaande rswo ud nea r Hilversum , The N etherl ands ,
in O ctober 1970 .

11 6. Penicillium megasporum Orpurt el Fennell: fig. 116, colour plate 116,

and SEM plat e 20, (top: right).
Penicillium megasporum Orpurt et Fennell - Myeologia 47: 233-237. 1955. See also
M.M . Kulik, 'A Compilation of Deseriptions of New Penici//ium Speeies'. Agrie.
Handbook No. 351. Agrie . Res. Ser. U.S. Dep. Agrie. pp . 54-55. 1968.
Orpurt and Fennell's original diagnosis follows, sinee this adequately presents the
species in question when first isolated:
'Colonies upon Czapek's agar growing fairly rapidly, attaining a diameter of
40-45 mm in two weeks at room temperature; eonsisting of a thin, c1ose-textured
mycelium, forming an almost brittle felt; radiately furrowed, central area often
raised and bearing Iimited eonidial struetures, becoming olive-buff to deep olive
(Ridgway, PI. XLV); surrounded by a zone 2-3 mm wide, beeoming dull greenish

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Fig. 116. Camera lucida

drawings of Penicillium
megasporum Orpurt et
Fennell, CSS 256.55. A,
Detailed drawings of the
penicilli. S, Mature conidia.

blaek (Ridgway, PI. XLVII) due to the presenee of abundant eonidial struetures,
shading into a white submarginal area; exudate laeking; odour not distinetive;
reverse olive-oehre to yellow-olive at 10 days, near Prout's brown (Ridgway, PI. XV)
in age, with the pigment diffusing into the agar; eonidiophores arising from the substrate or the aerial felt, septate, smooth, hyaline, 3.0-4.5 .m in diameter and up to
300 .m in height; penicilli biverticillate, asymmetrie, divarieate, variable in pattern
but usually eonsisting of a verticil of 2-6 divarieate metulae; mostly 4.0 by 10-12
.m; phialides 3-5 per metula, divergent, bottle shaped, 5.0-7.0 by 3.0-4.0 .m;
eonidia in short divergent ehains, spherieal, averaging 6.7 .m in diameter (5.0-7.0
.m), thiek walled, dark olive-brown at maturity, eoarsely eehinulate, with dark eoloured spines 1.0-1.5 .m in length; prominent disjunetors 1.0-2.0 .m long by
1.0-1.5 .m wide.
Colonies on malt extraet agar growing more slowly, attaining a diameter of 35-40
mm in 10 days, flat, unfurrowed, vegetative myeelium submerged, not forming a
brittle felt, heavily sporulating, abundantly velvety but showing a thin surfaee network of aerial hyphae bearing seattered conidial structures, with thin marginal area
white, shading rapidly through shades of American green to dull blackish green
(Ridgway, PI. XU, finally becoming olivaceous black (Ridgway, PI. XLVII); no
exudate; reverse uneoloured or nearly so; penicilli as deseribed above but oeeasio-

nolly showing much more complex structures rebranched as many as five or six
times below the level of the phialides, consistently producing the coarsely
hinulate, spherical conidia characteristic of the species.'

' I'he original diagnosis presented aboye essentially conforms with our own
nhscrva tion of the type strain in connection with the current study . Our own,
ud d ition a l, cultural observations follow :
C olonies upon CA (colour plate 116, A and a) attaining a diameter of25 mm
iu 14 days at room temperature (20- 25C), consisting of thin, close-textured
husal felt, not furrowed, flat , with thin margin surrounded by a narrow fringe
111' submerged hyphae extending about 1 mm beyond the aerial growth, colony
xu rl ace persistently white in marginal areas and showing pale yellow shades in
submarginal and central areas ; almost sterile , with colony surface overlaid by a
I hin, closely woven network of vegetative hyphae ; exudate and odour lacking;
ura ngc-colou red pigment diffusing into the surrounding agar; colony reverse
i 11 pal e yellowish shades (colour plate 116, a) .
Colon ies upon CYA (colour plate 116, B and b) growing more rapidly,
nu a ining a diameter of 40 mm at room temperature (20-25C) , similar in
tcxture to the preceding, but with undulating white margin slightly furrowed,
surr ounded by a fringe of submerged hyphae extending about 1 mm beyond
1he aerial growth; colony surfac e very poorly sporing, usually with conidial
a rcas obscured by a dense network of aerial vegetative hyphae lending a pale
llrange colour to the colony , which is almost plane and not furrowed in central
a nd submarginal areas; orange coloured pigment diffusing into the
surrounding agar; colony reverse sim ilarly coloured (colour plate 116, b) .
Colonies upon MEA (colour plate 116, C and c) attaining a diameter of 25
111m in 14 days at room temperature (20-25 C) , appearing 'wet' , white,
slightly raised in the centre , not furrowed ; slowly shading towards deep olive
hrown colours, becoming almost black with the ripening of conidia after 21
days (colour plate 116 , C); heavily sporing throughout ; colony surface overlaid
by a thin network of tloccose vegetative hyphae and ropes of hyphae
intluencing the colony appea rance during the first two weeks, then being
obscured by the ripening of conidia ; colony reverse in pale yellow shades
(colour plate 116 , c).
Penicilli essentially as described in the original diagnosis (fig . 116, A), and
conid ia attaining a diameter of 7 tui, conspicuously echinulate (fig . 116, B and
SE M plate 20, top : right).
The species description is centered upon the type strain CBS 256 .55 = ATCC
12,322 = NRRL 2,232 = QM 6,879 = WB 2,232 = IJFM 7046, isolated by] . H .
Warcup from heathland soil in Suffolk, UK, in 1947. Similar strains have been
isolated by P .A. Orpurt and D .!. Fennell from grassland soil collected at a
depth of one inch at five stations in southern Wisconsin , USA . between May

195 2 and M ay 19:>3. 0 1' th e nin c strai ns isolat ed Ih >111 Wis consin soils , cight
duplicated th e pattern presented by the typc . Howevcr, une isolat e , d iffered
fro m th ese stra ins in devel oping more slowly-grow ing, deep er , almos t lan at e
colo nies; in produ cing m ore variable and o fte n fractional or m al formed
penicil!i; and , in bearing ' wa r ty' rather th an stro ng ly ec hin ula te co nid ia .
Stra in C B S 529 .69 = VK M F-l ,824 wa s isolated by G .C. Blatt from forest soil
under Thuja occidentalis, in Canada , 1969 . Strain C BS 121. 65 was isolat ed by J.
Nicot from beach sa nd , in th e Leb anon , in 1965 .
Penicillium megasporum appears most satisfact orily relat ed to sp ecies of th e P.
nigricans se ries, espec ia lly to P. melinii, altho ugh it differs m arkedly from th e
latter sp ecies in cer ta in striking fea tures , namely : The globo se, large, rough
co nid ia of P. megasporum exc eed ing 6.5 Ilm in di amet er, wh ereas th ose of P.
melinii m easure 3.0-3 .5 uui ; a nd th e co n id iopho res of P. megasporum are
smooth upon al! substrata while th ose o f P. melinii a re cons picuo us ly warty .
T he sp ecies m ay be rega rd ed as a soil org a nism world-wide in d istribution .

117 . Penicillium ovetense Ramrez et M artnez: Fig. 117 , colou r plat e 117,
and SEM plate 20 , (centre: left).

~~~Jr
1:-- t
100l'm

f.R-

Fig. 117. Camera lucida

drawings of Penicillium
ovetense Ramrez et
Martnez, ATCC 42,237. A,
Detailed drawings of the
penicill i. B, Mature conidia.
C, Habit sketches.

nicillium ovetense Ramlrez et Martln ez - My copath ologia. 74, 35-49, 1981,

:olon ics up on C A (co lou r plat o 117, A a nd a) gro wing very rcstrictcdly ,
uruinin g a di amet cr 01' 15 mm in 14 days a l roo m tc rnpc ra tu re (20 -25 C),

l'onSisling o f a tou gh , close-tcxtu rcd ba sal Iclt , a ppcaring vclv et y , with a

nurruw u ndulatin g marg in su r ro u nde d by a n arro w fr inge a l' su b merged
h yphae exte nd in g a bo u t 1 mm beyon d the ae r ia l grow th, the n a b ru p tly raised
11) a plat eau somewhat buckled a nd fu rro wed, covered by large drops 01'
cx u rlatc be a ut ifuJly coloured in top az ye llow shades , becoming d eep vinaceous
1'1 '11 in age; co lo ny surface ove rlaid by a close ly woven ne twork 01' whi te to r osy
white vege ta tive hyphae in flue nc ing the colony a p pearance; hea vily sporing in
Illarg-i nal a reas in bl ue- green shades (Sguy, PI. X X X I, 46 1); no odo u r; pale
u rubcr -colou red pigment slow ly di ffu sin g into th e su rrou nd ing agar; colony
n -verse (co lou r plat e 117 , a) in pal e red dish cin namo n shades; pe n iciJli
ryp icall y asym metr ica l a nd st rongly divaricate, co ns ist ing al' ve rticils 01'
ruerul ae 01' b ranch es arising a t different levels on th e co n idiopho res (fig. 117 ,
A); con id iophores smooth walled, a ris ing p r imar ily from trailing hyphae ,
xho rt , not ex cee d ing 100 .tm in length by 2 .0-3.0 .tm in di amet er , w ith a p ices
"l lIarged in a ves icle up to 3.5 tui: in di amet e r ; branch es up to 25 .tm in length
b y 2.0 .tm in di ameter; m e tul ae in c1us ters 01' 2- 4 eleme n ts in th e ve rticil,
s lro ngly di ve rgent, mostly 20 -35 .tm in length b y 1.5 - 2. 0 .tm in di amet er ,
w it h a p ices co ns pic uo us ly en larged in a ves icle up to 5 .tm in di amet er ;
p hialides in cro wde d c1usters al' 12 a l' m ore elements in the vertic il, b ottle
sha ped , taperin g to a sho r t, narrow ed neck , a bou t 9-11 .tm by 3 .0-4 .0 tuu ;
ro nid ia globose, cons picuo us ly sp inu lose (fig . 117, B a nd SE M pla te 20,
r cnt re : left), a bo u t 2 .8 -3 .5 .tm in di amet er ; co n id ia l cha ins form in g well d cfined dive rge nt colu m ns (fig . 117 , C) .
Colon ies upon C YA (co lou r plate 117 , B a n d b), attain ing a d iamet er a l' 25
l o 30 mm in 14 da ys a t room temperature (20-2 5 C) , fine ly vel vet y , co ns isting
01' a c1ose-tex tured b a sal felt , w ith undulat in g m a r gin radiat ely fu rrowed ;
subce ntral a nd cen tral areas rai sed ab rup tly a nd depressed in a crate r ifor m
pattern ; abundantly spo r ing in pal e blue- gray shades (Sguy, PI. XXX , 450) ;
colony su rface covered in cen tral areas b y small tufts 01' white vegeta tive
hyphae ; exudate ab u ndan tly p roduced as small d roplet s beautifull y co loured in
deep gren ate red shade s; odo u r faint , indefinite ; a rnbe r-c olou red pi gment
diffu sin g into th e surro u nd ing agar; becomin g vin aceous in age ; colo ny reverse
(colour pl at e 117 , b ) developing ve ry dark blue-black shades (Sguy , PI.
XXX IV, 506) ; penicll i esse n tia Jly as described a boye .
Colon ies upon M EA (co lour pl a te 117 , C and c) , gr ow ing very restrictedly ,
a tta in in g a di amet er 01' 15- 20 mm in 14 day s at room temperature (20- 25 C ),
consistin g 01' a looser textured bas al felt , ty pically velvety, somewhat zanate,
with a white , fin ely crene lle d margin surrounded b y a fr inge 01' su b merge d

hyphae extend ing abou t 2 mm beyond the aerial gro wt h; colony surf ace fla t,
not furrowed ; heavil y sporing throu gh out in dcep olivc-gray shades (S guy, PI.
XXIX , 433 ); exuda tc and odour lacking; no colou red pigm en t diffusin g into
the su rro und ing ag ar; colony reverse (colour plate 117, e) brown with a gr een
cas t show ing throu gh . Penicilli essentially as described aboye .
The spec ies description is ce ntere d upon th e type stra in IJFM 7029 = AT CC
42,237 , isolat ed frorn sandy soil in M adrid , Spain , in 1979 , by th e author. It is
at present known only from the typ e strain .
Penicillium ovetense is a di stinctive species characterized by its typ ical
divaricate penicilli, although sim ple forms are not unfrequ ently enco untered
(appearing monoverticillate), by its globose, spin ulose conid ia , by its restricted
gro wth on most su bstra ta, a nd by th e ve ry dark olive-gray shade s a l' its colon ies
u pon MEA , su ggestiv e a l' th e species inclu de d in the P. nigricans seri es to which
it appea rs mo st satisfactor ily ass ignable .

118 . Penicillium yarmokense Baghdadi : fig. 118, colour plat e 118, and

SEM piate 20, (centre: right).

y:~

1.....----1

100)Jm

Fig. 118. Camera lucida

drawings of Penicillium
yarmokense 8aghdadi, C8S
410.69. A, Detailed drawings
of the penicilli. 8, Mature
conidia. C, Habit sketches .

nlcillium vermokense Baghdadi - Nov. Syst. Nlz. Rast. pp. 99-101. 1968.
~ ol o n i e s

upon CA (colour plate 118, A and a) growing restrictedIy,

uiuining a diameter of 25 mm in 14 days at room temperature (20-25C),

co nsisting of a tough, close-textured feit, vclvety in appearance, strongIy
rud ia tcly furrowed, with centre raised and irreguIarly wrinkIed in a crateriform
patlern; with finely crenelled margin surrounded by a fringe of submerged
hyphae extending about 2 mm beyond the aerial growth; heavily sporing
I hroughout in gray-green shades (Sguy, PI. XXIX, 428); exudate abundantly
produced, as crystal clear colourless drops; odour lacking; no coloured pigment
r1ilTusing into the surrounding agar; colony reverse (coIour pIate 118, a) in
ycllowish ochre shades; penicilli typically divaricate asymmetrical (fig . 118,
A}, with conidiophores smooth walled, arising either from the substratum or
lrom aerial hyphae , 70-500 m in length by 2.5-3.0 m in diameter; penicilli
c:onsisting of branches arranged at different levels of the conidiophores
I erminating in a verticil of metulae of unequallength, or of verticils of metulae
nlo nc at the tip of the conidiophores, bearing clusters of phialides; branches,
usually 25-30 m by 2.0-2.5 m; metulae in clusters of 2-3 elements in the
vc rticil, mostly 8-20 m by 2.3-3 .5 m, slightly enlarged at the apex in a
vcsicle up to 4 .0 m in diameter; phialides bottle shaped, in clusters of 2-12 or
more elements in the verticil, 9-11 m by 3.0-3.5 m, conspicuously tapering
lo a narrow, short neck; conidia globose to subglobose, smooth walled when
observed under the light microscope (fig . 118, B) but very delicately granular
when observed under the SEM (plate 20, centre: right), mostly 3.0-3 .5 utt: in
rliameter; conidial chains forming loosely parallel columns (fig. 118, C) .
Colonies upon CYA (colour plate 118, B and b) spreading broadly, attaining
a diameter of 45 mm in 14 days at room temperature, similar in texture to
I hose grown upon CA, but more regularly furrowed in a radial pattern,
I ypically velvety, conspicuously zanate owing to the different degree of
ripeness of zones, gradually raised towards the centre; uneven margin
surrounded by a narrow fringe of submerged hyphae extending about 1 mm
beyond the aerial growth; heavily sporing throughout in different shades of
gray-green, depending upon the degree of maturity of conidia (Sguy, PI.
XXIX, 427,428,429); exudate very abundantly produced as large, light yellow
drops; odour lacking; pale amber-coloured pigment diffusing into the
surrounding agar; colony reverse (colour plate 118, b) in reddish to ochre
shades; penicilli essentially as described aboye.
Colonies upon MEA (colour plate 118, C and c) attaining a diameter of 40
mm in 14 days at room temperature (20- 25C), consisting of a looser textured
basal feit, thin, flat , typically delicately velvety, with thin margin irregularly
dissected, surrounded by a fringe of submerged hyphae extending about 3 mm
beyond the aerial growth; somewhat zanate; heavily sporing throughout in

gray-green to arternisia green sha de s (Sgu y, P\. XX 1X , 429 , 4:H) ; cxudate

a nd odour lacking; no co loured pigment d iffusin g int o the surrounding agar;
colony reverse in yellowish shades (colour plate 118, e); pcnicilli essent ially as
des cribed aboye but with conid iophores mostly arising from the substratum .
The species des cription is centered upon the type strain CBS 410 .69 = VKM
F-l ,076 = CMI 140,346 = IJFM 7044 , isolated by V . Ch . Baghdadi from
Syrain desert soil , Es-Suveida, Syria, O ctober, 1964 .
Penicilliumy armokense is a distinctive species which clearly appears to be mo st
satisfactorily assignable to th e Asymmetrica-divaricata sub-sect ion , particula rly to the P. nigricans series, on account of its cultural characteristics.

Penicillium series atro-sanguineum ser. nov .

Coloniae in agaro Czapekii restrietae ereseunt; penieilli asymmetrici divarieati;
eonidia parva; eum pigmento rubro sanguineo diffluente in medio.
Typus: Penicillium atro-sanguineum Dong.

Penicillium atro-sanguineum series

The newly erected Penicillium atro-sanguineum series is ad m itt edly artificial in
con cept and has been designed to cover two forms whi ch cannot otherw ise be
satisfactorily placed in a ny of Raper and Thom 's (1949) recognized series ,
namely: Penicillium atro-sanguineum Dong, a nd P. griseo-purpureum Smith . A close
relationship between these species is not presumed , and they are considered
together primarily as a matter of convenience . They do, how ever, show certa in
characteristics in common, particularly th e strongly divaricate cha racte r of th e
penicilli, usually with con id ial chains showing a tenden cy to produce loosely
divergent colu m ns; the com pa ratively sm all , globose to subglobose , smoothwall ed con idia ; and espec ially the production of a deeply red pigment diffusing
into the surrounding agar on certain substrat a .

119. Penicillium atro-sanguinem-Dong: fig. 119, colour pIate 119, and

SEM plate 20, (bottom: Ieft).
Penicillium atro-sanguineum Dong - Ceska Mykologie. 27: 174-176. 1973.
Dong's original diagnosis follows:
'Colonies upon Czapek-Dox agar, attaining a diameter of 30 to 35 mm in 12 days
at 22 to 24C, floeeose, gray-green (Sguy, PI. XVI , approx. 310), eolony reverse
bright yellow-green (Sguy, PI. XVII , approx. apple green, 268); exudate laeking;
white margin, 2 to 3 mm w ide.
Colonies upon malt extraet agar (2%), attaining a diameter of 60 mm in 12 days at

0 0

08 00

0 0
O

co

00

o o 0 00
o 00 O o
00 o
o o

R.

Fig. 119. Camera lucida

drawings of Penicillium etrosanguineum Dong, CBS
380.75. A, Detailed drawings
of the penicilli. B, Large
mature conidia. C, Small
mature conidia. D, Habit
sketches.

22 to 24C , tanate , grayish-green , with radial zones, spotted in violet red shades ;
colony reverse and agar similarly coloured in deep red shades approximating ox
blood (Sguy, PI. X, 141); exudate Iimited in amount, colourless or lacking .
Conidiophores arising from aerial hyphae, 25 to 150 Ilm in length, smooth-walled.
Penicilli irregular in pattern, mostly divaricate, consisting of c1usters of 2 to 4
metulae, although simple penicilli (appearing monoverticillatel and irregu larly
branched are not unfrequently encountered. Branches, 8 to 15 1lm by 2.0 to 2.5Ilm;
metulae, 6 to 12 1lm by 2.0 to 2.5Ilm; phialides bottle-shaped, 5.0 to 7.5 1lm by 1.7
to 2.2 Ilm , in c1usters of 5 to 8 elements in the verticil , sometimes single on the t ip of
the metula. Conidia globose, varying in dimens ions , 1.5 to 1.8 1lm or 3.0 to 3.5 1lm in
diameter, smooth-walled , in short diverging chains.' (Translated from the French by
the author) .

Our no tes follow :

Colonies u pon CA (co lour plate 119 , A and a) attaining a diameter of 45 mm
in 14 days at room tempe rature (20 -25 C), deeply velvety to lanate ,
cons isting of a close-textured basal felt, overlaid by a looser textu red network

01' floccosc aerial hyphae in lucn cin g gre a tly (he colo ny a ppcara ncc , not
furrowed, slightly and gradually rai sed towards the centre ; thin evcn margin
surrounded by a submerged fringe extending about 1 mm bcyond the aerial
growth; heavily sporing throughout in dark blue-grcen shades (Sguy, PI.
XXX, 447) in sectors not ov erlaid by the nctwork 01' floccose hyphae, 01'
showing arte misia gre en shades in zones covered by th e m ention ed network 01'
vegetative h yphae; odour lacking; exudate abundantIy produced as minute,
crystal clear droplets enm eshe d in the my celial felt; bright yellow-orange
coloured pigment diffusing into the surrounding agar; colony reverse (colour
plate 119, a) developing bright yellow colour; penicilli (fig. 119 , A) typi cally
div aricate and asymmetrical, with con id iophore s smooth walled, arising from
a erial hyphae, up to 200 Jlm in length by 2.8-3 .0 Jlm in diameter, consisting 01'
branches variously arranged along the conid iophore, and 01' clusters 01' metulae
01' 2-4 elements in th e verticil, bearing cro wded whorls 01' phialides; branches
usu ally ranging from 10 to 30 Jlm in length by 2.5-3.0 tut: in diameter;
mctulae , 8-15 Jlm by 2.5-2 .8 uiu, with a pices slightIy enla rged in a vesicle up
to 3.5 Jlm in diameter; phialides bottIe shaped , in clusters 01' 3-6 elements in
th e verticil , div ergent , mostIy 7-8 Jlm by 3.0 ust , conspicuously tapering to a
short , narrowed ne ck ; conidia glob ose, smooth wall ed wh en obs erved under
the light microscop e (fig . 119 , B and C), but finel y rough ened when obs erved
under th e SEM (plate 20, bottom: left) , ranging from very small, 1.5-2 .0 Jlm
diameter (fig. 119, C) to larger, about 3 .0 Jlm in diameter (fig . 119 , B) ;
conidial cha ins producing short div ergent colu m ns (fig . 119, D).
C olonies upon CYA (colour plate 119 , B and b), attaining a diameter 01' 45
mm in 14 days at room temperature (20-25 C) , cons isting 01' a close-textured
basal felt overlaid by a looser-textured network 01' floccose hyphae lending a
lana te appearance to th e otherwise deeply velvety colony surface, whi ch is
deeply radiately furrow ed with th e centre irregularly foldcd and depressed ;
thin white m argin surrounded by a fringe 01' su bm erged hyphae extending
about 1 mm beyond th e aerial growth; conspicuously zanate owing to the
variou s degrees 01' ripeness 01' con id ial zones ; heavily sporing through out in the
same shades a s th e preceding, with conidial areas more 01' less obscured by the
deep floccose ov er growth 01' aerial hyphae ; exudate abundantIy produced , a s
minute yellow droplets enmeshed in th e mycelial felt ; odour lacking; am bercolou re d pi gment diffu sing into the surround in g agar; colony rcvers e (colour
plat e 119 , b) developing reddish cinnamon shades ; penicilli essentially as
des cribed aboye.
Colonies upon MEA (colour plate 119 , C and e) attaining a diameter 01' 35
mm in 14 days a t room temperature (20 -25 C) , consisting ofa looser tcxtured
basal feIt, deeply velvety , overlaid by a loo sely wov en network 01' floccose aerial
hyphae lending a lanat e appearance to th e colony surface , not furrowed ,
slightIy rais ed tow ards th e centre , which is m arked by a loose overgrowth 01'

,J';;

loccul cn t hyphae; heavily spo r ing th rou gh out in oliv e-gr een shade s (S guy,
PI. XXIX, 422), partially obsc u rc d by the Iloccose ove rgrowt h 01' aer ial hyphae
whi ch dev elop blue- gr een sh ades (Sguy, PI. XXIX, 425) ; odo u r and exudare
lacking; deep gr enat e , blo od red, to deepl y vinaccou s, beautifully coloured,
pigment diffusing into th e su r ro u nd in g agar; co lony reverse (co lour plate 119,
e) dc veloping deep purple red to blood red colou rs; penicilli essen tially as
d escribed above.
The sp ecies description is ce n te re d upon th e type st rain C ES 38 0 .75 = C M I
197,488 = IJFM 7058 , isolat ed b y DI' Bui xu a n D ong , Faculty 01' Pharmacy,
U niv ersity 0 1' H anoi, Vi etnam , from see ds 0 1' Trit icum vulgare L. in a cereal
preservin g pit , Prague , Czechoslovakia, April , 1971.
Penicillium atro-sanguineum is a ve ry distinctive sp ecies characte rized
particul arly by th e deep red pi gment develop ed upon MEA a nd diffusing into
t he su rrou nd ing a ga r , by its typicall y divaricate penicill i produ cin g co nid ia 0 1'
two di ffer ent sizes , eithe r in th e sa me penicillus 0 1' in different st r uc tures,
whi ch is one 01' th e most striking features 0 1' thi s organism o
Ir ap pe a rs to be m ost sa tisfac to r ily assi gn able to th e Asymmetri ca-divaricata sub-section , bu t it ca nno t be placed in a ny 0 1' R ap er a nd
Thom ' s (1949) recognized series , hen ce its inclusion here .

120. Penicillium griseo-purpureum Smith: Fig. 120, Colour plat e 120, and
SEM plate 20, (bottom right).
Penicil/ium griseo-purpureum Smith - Trans . Brit. Mycol. SOCo 48: 274-275. 1965.
See also M.M. Kulik, 'A Compilation of Descriptions of New Penicil/ium Species' .
Agric. Handbook No. 351. Agric. Res. Ser. U.S . Dep. Agric. p. 43. 1968.

Smith's original diagnosis follows:

'Colonies on Czapek agar growing somewhat slowly, 16 to 17 mm in diameter in 7
days at 24C, bluish gray green at first, rapidly becoming almost pure gray, with no
white margin, thick velvety, with reverse at first dull purple, not diffusing, becoming
very dark brownish purple; colonies on malt agar growing slightly more rapidly, 20
mm in diameter in 7 days, slightly funiculose, coloured as on Czapek agar, with
reverse deep dull purple, diffusing somewhat; penicilli consisting of whorls of
rnetulae, somewhat divergent, and often unequal in length in a single verticil,
occasionally with a secondary branch from the next lower node of the
conidiophore, this bearing phialides only or with a small cluster of metulae each
bearing phialides; con idiophores arising mostly from the substratum, usually about
500 p'm long and 2.4 to 3.2 p'm in diameter, slightly rough; metulae 8 to 18 p'm by 2.5
to 3.5 p'm; phialides with fairly long-pointed tips, 10 p'm by 2.0 to 2.2 p'm; conidia
globose to subglobose, sligltly rough, 2.2 to 2.6 p'm in diameter, in chains, at first
divergent or in iII-defined columns, becoming more or less tangled.'

:1 ~ 1>

O~o

0
0 0 O
00 B O

O O 0000
O O

'."
'1''''.i("

'.

""<'1

. J/
'

LL=lwoJ

O 10).lm

c ;'
.'
r'')) ji:.

L-----I

100,JJm

l'

fR. ,

Fig. 120. Camera lucida

drawings of Penicil/ium
grieso-purpureum Smith,
CSS 406.55. A, Detailed
draw ings of the penicilli. S,
Mature conidia. e, Habit
sketches .

Our not es follow:

C olonies upon CA (colour plat e 120 , A and a) gro wing restricted ly,
a tta ining a di ameter of 25 mm in 14 days at room tem perature (20 -25 C),
cons isting of a tou gh , close -textured basal fe1t, deepl y velvety in appearance,
ove rla id by a thi n, close-t extured network of fine aerial white hyphae
influe nc ing greatly the colony appearance, lending it a su bla nate aspect; thin
flat m argin surrounded by a na rrow fringe of su bmerge d hyphae extend ing
abo ut 1 mm beyond th e ae rial growth; submargi nal area abruptly raised and
radiately furrowed, w ith ce nt ra l area de pressed in a crateriform pa ttern ;
heavily spor ing in pale blue- green shades (Sguy, PI. XXXII I, 484)
particul arly conspicuo us whe n conid ial areas are not obs cured by th e
overgrowth of fine wh ite vegetative h yph ae , as in m arginal areas , othe r areas
rem aining pe rs istently wh ite ; exuda te a nd odour lackin g ; pale yellow ish purple
pigm en t di ffusin g slowly into the surro unding agar; colony reverse (colour
plat e 120 , a) at first in cinnamo n shades, becom ing soo n ve ry da rk violet
purple (Sguy , PI. XLV, 666) in colour , sta rt ing from th e margin towards the
centre; penicilli esse nt ially as described in Smith's origina l diagnos is (fig. 120 ,

1\), with co n id io p ho res sm oo th wall cd 01' ncarl y SO, sc p ta tc, a r ising' di rectl y
Iro m the su bs tru tu m, c nla rgc d a l th e a pex in a vesicle u p lo :> un in di um et cr ,
usu all y up l o 1:>0 un in lcn gth by 2.0 - 2 .:> J.l1II in diamct er , bcaring who rls 01'
mctuluc a r mare a l' lcss unequal len gth , 10- 2:> J.l1II by 2.0 -3.0 iutv , enla rge d a t
th e a pex in a vcs icle u p to 4.5 J.lm in dia m ct cr ; phialides in clu sters 01' 3 -8
clc mc n ts in th e ve r ticil, bottl e shaped , m ostly 10 iut: by 2.5 -3 .0 uin ,
conspicuously tapering to a shor t narrow ed neck ; co n id ia com para tively small,
xmooth wall ed 01' nearl y so wh en obse rve d under th e ligh t m icr oscop e (fig. 120 ,
B) but delica tely gra n ula r wh en obse rve d under th e SEM (pla te 20, bottom :
rigbt) , mostl y 2 .0 -2 .5 J.lm in lon g ax is; co nid ia l chains sho rt, in loose colu m ns
( Iig. 120 , C) .
Colonies u po n C YA (co lour pl at e 120 , B a nd b) gr owing more rapidly than
ahove, a ttain ing a di am et er 01' 35 mm in 14 d ays at roo m temperature,
co nsisting 01' a close -textu re d basal felt , typicall y a nd fin ely velvety,
cons picuo us ly za nate owi ng to the various degr ees 01' ripe ness 01' co n id ial
structures; m argin Ila t, not furrowed , su rrou nde d by a narrow fringe 01'
subme rge d h yphae ex te nd ing a bo u t 1 m m beyond th e ae r ia l growth;
su bm arg inal a rea a nd cen tre d eepl y radia tely fu r ro wed with ce n tre irregularly
olde d in a crater ifor m pattern a nd sho wing crev ices; heavil y spo ring
rhroughout d evel oping brownish gree n sh ades , becoming blue- gr een with the
ripe ning 01' co n idia (Sguy, PI. XXXI , 453) , sta r ting in m arginal a reas;
cxudate a b u ndan tly produced as large drops developing deep vinaceous red
colou rs; odo u r lacking; deep red -colou red pi gm en t diffusin g into the
su r rou nd ing agar; colony reverse (colour plate 120 , b) in the same d ark violet
pu rple show n on C A; th e agar being stron gly pulled away fro m the Petri d ish ;
penicilli esse n tia lly as described .
C olonies upon M EA (colour plate 120 , C a nd e), sp read ing broadly ,
a ttain ing a diameter 01' 50 mm in 14 da ys at ro om temperature (20 -25 C),
cons isting 01' a looser textured basal felt , typi call y velvet y , flat , not furrowed ,
with thin m argin su rrou nde d by a wide frin ge 01' su bme rge d hypha e exte nd ing
abou t 5 mm beyond th e ae r ia l growth; heavily sporing throu gh out in graygree n shade s (Sguy, PI. XXI , 309); ex udate a nd odour la cking; no coloured
pigm ent di ffusin g into th e surro u ndi ng agar; colony revers e (colour plat e 120 ,
c) developing reddish brown shades sta rti ng at th e ce ntre a nd exte nd ing
gra dually towards the m argin ; penicilli esse n tia lly as described a boye .
The spec ies description is ce nte re d upon th e typ e strain C BS 406 .65 = C M !
96 ,1 57 = A TCC 22 ,353 = !J FM 7042, isolat ed by B.A . Indraratne from forest
soil under Pinus sp ., Freshfield Forest , Lancaste r, UK , 1962.
Penicillium griseo-p urp ureum clearly belon gs to th e Asym me trica - d ivarica ta
su b-section, a nd is distinguish ed by th e deep violet -purple colou rs sho w n by
th e colony reverse upon CA a nd CYA , by th e deep red-coloured pi gm ent
di ffusin g into th e surrou nd ing agar upon C YA , a nd by its co m parative ly sm all

:1:U

conid ia . Howcver , it cannot be sutis fuctor ily assigned lo any 01' the Raper and
T hom's ( 1949) recogn ized se ries, hen ce its in clu sion here. lt is actually know n
only from the type st ra in .

Penicillium series brasilianum ser. nov .

Coloniae celeriter crescunt, floccosae; penicilli asymmetrici divaricati; conidia
fortiter elliptica , plus minus apiculata , fortiter asperula .
Typus: Penicillium !2rasilianum Batista.

Penicillium brasilianum series

OUTSTANDING FEATU RES

T he Penicillium brasilianum series is another newly erected group 01' Peni cillia ,
admitted artificially in concept, wh ich has bee n designed to cover five for ms
wh ich cannot otherwise be satisfactorily placed in any 01' R a per and T hom' s
(1949) recognized series, namely : Penicillium brasilianum Ba tista , P. kabunicum
Raghdadi, P. moldavicum Milko et Belja kova, P. onobense R am rez et Martnez,
and P. castellonense R am rez et Martnez. A close re lationship be tween these
species is, however , not necessarily presumed, and they are considered
together primarily as a matter 01' conven ience. T hey do, however, show certain
characteristics in common, particularly the divaricate character 01' th e penicilli,
its broadly spreading co lonies upon most substrata, the deeply floccose felt 01'
the colony surface, and its us ually strongly ellip itical, more or less apiculate
and rough conidia.

Key to species
Page
l. Colonies not producing sclerotia or masses of thick-walled cells.
Colonies showing some shades of green, gray , gray green, or blue-green
with the ripening of conid ia. Penicilli with the divaricate character
evident. Colon ies spreading broadly upon all substra ta, appearing more
or less floccose.
P. brasilianum series
A . Conidia globose or subglobose .
1. Conidia conspicuously spinulose.
P. castellonense Ramrez et Ma rtnez
2. Conidia smooth-walled or nearly so, pyriform , w ith one apiculate
end .
P. kabunicum Baghdad i
B. Conidia Iimoniform , conspicuously spinulose.
P. moldavicum Milko et Beljakova

. ... 328

... . 338

. . .. 331
. .. . 334

C. Conidia elllptical, conspicuously roughened, with roughness

arranged in a spiral pattern.
1. Colonies upon malt extract agar heavily sporing in deep gray-green
shades; colony reverse developing green-black colours.
P. brasilianum Batista . .. , 329
2. Colonies upon malt extract agar sporing in pale gray shades; colon y
reverse developing deep orange colours.
P. onobense Ramrez et Martnez . ... 336

121. Penicillium brasilianum Batista: fig. 121, colour plate 121 , and SEM
plate 21, (top: left).

.'

;:.

o@ BW'@
JJ

~~ : 8~
Lu.uJ.J.wJ rl'<l

10pm

ti"

i l rt~at.

Fig. 121, Camera lucida

drawings of Penicillium
brasilianum Batista , CBS
253.55. A, Detailed drawings
of the penicilli. B, Mature
conidia . e, Habit sketches .

Penicillium brasilianum Batista - Ann. Soco Biol. Pernambuco. 15: 160. 1957. See
also M.M. Kulik, 'A Compilation of Descriptions of New Penicillium Species'.
Agric. Handbook No. 351. Agric . Res. Ser. U.S . Dep. Agric. p. 21. 1968.
Batista's original diagnosis follows:
'Colonies upon neutral Czapek's agar attaining a diameter of 35-40 mm in 10

days at room temperature, consisting of a thin f elt, gray-green, velvety, superfic ially
f loccose, wrinkled , plane, sometimes umbonate, with radial furrows, w ith even
margin, 2-10 mm wide; at first, after 3 days old , they are white; azonate; odour and
exudate lacking ; colony reverse yellow, appearing orange-yellow in some stra ins,
soon becoming dark brown .
Vegetative submerged hyphae hyaline. Conid iophores arising f rom submerged
hyphae, erect, slightly rough walled , with walls th in and yellow-coloured , septate,
spread ing abundantly all ove r the colony and up to 480 .tm in lengt h by 2-2.8 .tm in
diamete r.
Penicilli borne singly on the conidiophore or on strongly divar icate branches.
Branches 8-20 by 2-3 .tm . Metulae truncate, incurved, in c1usters of 2-6 elements,
atta ining 7 .5-12 .tm by 2.5-3.0 .tm .
Some st rains do not show branches and the verticils of metulae may appa rently
be looking as monoverticillate structu res. Phialides 5-8 per metula , aproximately of
equal length , appearing fusiform tapering to a narrowed neck, 5.5-10 .tm by 2.5
.tm.

Conidia form ing short or long chains, not exceed ing, however, 200 .tm in length;
they are elliptical to subglobose, yellow coloured , ech inulate, thin walled , 3.6 .tm by

4.5-5.0

.tm .

Cleistothecia and sclerotia lacking.'

(Translated from the Portuguese by the autho r).

Our not es follow :

Colonies u pon CA (e olour p lat e 121, A and a ) spreadin g broadly, atta ining a
di amet er of 55 mm in 14 da ys a t ro om temperature (20-2 5C) , eons isting of a
close -te x tu re d basal felt deepl y velvety , ove rlai d by a clos ely woven n etwork of
floeeo se white aerial hyphae influen ein g gr eatly th e eolony a ppe a ranee, lending
it a lanate appearan ee , irregu larly wrinkled with shallow radial furrows,
almost p lan e ; heavily sp oring throughout in dark oliv e green shades (S gu y,
PI. XXIX , 431 ), w ith fruitin g areas alm os t eom ple tely ob seured b y th e dense
n etwork of white floeeo se h yphae ; very thin , eve n m argin su r rou nded by a
wide fringe of submerged hyphae ex te nd ing ab out 10 mm beyond the ae r ial
growth ; exu d a te produeed a s sm all ye llow drops; odou r none ; no eo lou re d
pigment diffusing into th e surrounding a ga r; eol ony re ve rse (eolour p lat e 121 ,
a) at first in ein namon shades, soon beeoming ora nge in the m argin ; penieilli
typi eall y asym me tr ie di varieat e (fig. 121 , A), eo nsistin g of ve rtieils of m etulae
more or less of eq ual len gth in th e wh orl ; eo n id iopho res a rising primarily from
th e su bstratu m, ereet, delieat ely rou ghened , altho ugh srn ooth-walled
eon id iophore s a re not infrequently seen , mostl y 100- 200 uit: in le n gth by
2.0-3. 0 .tm in di amet er; m etulae more or less eylind r ieal a nd ineurved , 10-15
Lm by 2.5 -3 .0 Lm with a p iees ver y slightly enlarged ; phialides bottle sh a pe d ,
in clusters of 3- 8 eleme n ts in th e ver tieil, mo stl y 11-1 5 Lm by 2.5-3 .5 iut ,

rupcriu g lo a co nspicuously narrow cd short neck : co nid ia co ns picuo us ly ro ug h,

wirh roughncss a r ra nge d in a spiral pa ltcrn (fig. 121, B a nd SEM plat e 2 1, top :
lclt) , stro ng ly ellip tical, a bo u t 3.2 -4. 0 J,t1l1 by 2 .2 - :1.0 tun in di ameter; eo n id ial
chuins a rr angcd in looscly di ve rgen t columns (lig . 121 , C); sing le pcnieilli
(a pp cnr] ng rnonovertieillatc) a re also obse rved .
C olonies upon C YA (co lou r plate 121, B a nd b) attaining a d iamet er of55 mm
in 11 d ays a t room temperature (20 -25 C) , sim ila r in texture to the pr eced ing
hlll decply radiately fu rrowed in su b margi nal a reas and deepl y sulcated by
l'llllce ntrie furrows in ce n tral areas , sligh tly raised ; thin , eve n m argin
su rr ou nded by a frin ge of sub me rge d hyphae ex te nd ing a bo u t 2 mm beyond
I he acrial growth ; exudate a b u ndan tly produced , as pal e yellow drop s; odo ur
IIll11e ; no eolou re d pigmen t di ffusin g into th e su r ro u nd ing agar; co lon y reverse
(el,lour pla te 121 , b) a t first in cinnam on tints shad ing to sulph ur yellow
l'lJllJu rs in a ge ; pe nicilli essen tially as described a boye, with heavily spo ring
nrcas devcloping sim ilar olive green shades a nd m ostly com pletely obsc ured b y
t lu: den se net work of white floccose ae ria l hyphae .
Co lonies upon M EA (co lour pl at e 121 , C a nd c) attain ing a di amet er of60
111111 in 14 d ays a t room temperature (20- 25 C), co ns isting of a looser textured
hasal felt , a ppear ing typicall y deep velve ty, a lmost flat , wit h few radial shallow
lu rr ows, with ve ry thin eve n m argin la rgely su b merged, exte nd ing a bo u t 5 mm
hcyond the ae r ial gro wth; heavily spo ring th roughout in blue-green shades
(Sg uy, PI. XXXI , 461) ; ex udate lacking; od our fain t, a ro matic; no coloured
pigmen t d iffus ing into the su r ro u nd ing agar; colon y reverse (colou r pla te 121 ,
e) developing ye llowis h shades in margin al and su b marg inal areas and ve ry
c1 ark gre en, almost black co lou rs in ce ntral areas; penicilli esse n tia lly as
dcscribed a bo ye.
Penicillium brasilianum description is centere d upon th e typ e st ra in CBS 253.55
= ATCC 12, 072 = Q M 6 ,947 = IJ FM 7041 , isola ted by Prof. A.C . Batist a ,
In sti tute of M ycology , U n ivers ity of R ecife , Pernambuco , Brazil , as a cultu re
co nta m ina nt, 1954. It is actually kn own only from th e typ e strain a nd from
ano the r strain isolat ed from soil in th e Province of Navar ra, Spain, b y D I' A .T.
M artnez as IJ FM 7080 , in 19 78 .
Penicilli um brasilianum is a distincti ve spec ies characte rized by its typi call y
divarica te penicilli, its delica tely ro ug he ne d co nidiopho res, and th e stro ng ly
clliptical rough co n id ia , with the rou ghness ar ra nge d in spira l band s. It
a p pears to be m ost satisfacto rily ass ignable to th e Asymmetrica-divaricat a
su b-sect ion, but , however , it ca n not be place d in a ny of R a per a nd T ho m's
( 1949) recognized series, hence its inclusion here .

122 . Penicillium kabunicum Baghdadi : fig. 122, colour pIat e 122 , and
SE M plate 21, (top : right).

10

'O

jirn

OO

OO O

110

6'

o"J~~ t 1
'----J

100pm

.f{.
. . -~.'' ~;\l{ ~_. :1

~G:~

Fig. 122. Camera lucida

drawings of Penicillium
kabunicum Saghdad i, CSS
409.69. A , Detailed drawings
of the penicilli. S, Mature
conidia . e, Habit sketches.

Penicillium kabunicum Baghdadi - Nov. Syst . Niz. Rast. pp . 99-99. 1968.

C olonies on CA (colour plate 122 , A and a) spread ing broadly, a ttaining a
diameter of 60-65 mm in 14 days (20- 25C ), cons isting of a close -textu re d
basal felt , velvet y in appeara nce, overl aid by a deep closely woven net work of
floccose ae r ial hyphae influen cin g gr eatl y th e colony appearance , lending to it
a lanat e a spect; with undulating m argin surrounded by a fringe of submerged
h yphae ex te nd ing abo u t 1 mm beyond the ae r ial gro wth ; colony surface almost
plane with ce nt re umbon a te , ow ing to the deep floccose net work ofw h ite aeri al
h yphae; spa ringly spo ring throu ghout , with conid ial areas va r ious ly coloure d,
dep ending on the degree of r ipe ness of con id ia, ran gin g fro m yellow in
margin al areas, pink in su bce n tra l areas , and pale gray ish buff to m ou se gray
(Sguy, PI. XXXIII , 493) in th e ce ntre, m or e or less obsc ure d by th e floccose
ove rlay ing netw ork of ae rial hyphae ; odo u r faint , ind efinite ; exudate
abu ndantly produced as minute crystal clear droplets enme she d in th e m ycel ial
felt ; no coloure d pigm ent di ffusing into th e surro und ing a ga r; colony reverse
(colour plat e 122, a) a t first developing pale reddish brown colours turning to

.Ieep brown s hades in nge : pcni cilli (Iig . 122, A) typically div nri cat c , co ns isrin g'
01' ve rti cil s 01' m etul ac 01' tairly eq ual lc n g th in th c vcrticil 01' b ranch cs al
rli llcrcn t nodcs al' th c co nidiophorc, bearin g cro wde d clus ters 01' phi alid cs;
ru uid iophores arising frorn th e substratum and rom aerial hyphae , smoo th
wullcd 01' partially deli cately roughen ed, scpta te , ranging frorn 300 to 500 11m
lu lcn gth by 2.0-3 .0 11m in diameter when borne directly from the substratum ,
11\( 1 not cxcecding 100 11m in length when borne a s branches from a eri al
hyphae; simple penicilli (appearing monoverticillate) a re not unfrequently
uhsc rved : metulae commonl y wedge shaped wh en arranged in verticils 01' 3- 6
r-l cmc n ts in th e wh orl , sho rt, abo ut 10-13 11m in len gth by 3 .0 -3.5 11m in
rli.u uoter a t th e base , a nd 3 .5- 5. 0 11m in di amet er at th e apex; wh en arranged
in grou ps 01' two per ve rticil: m etulae are longer a nd narrower, about 25-30
/ UII by 2.0 11m with a pices enla rge d in a vesicl e up to 4 .0 11m in di ameter;
ph ialid es bottle shaped , in cro wde d clusters 01' 2 -1 O 0 1' m or e eleme n ts in th e
vcrt icil, 11-1 4 11m by 2.8-4.0 11m, tapering to a conspicuo usly long narrowed
ncck ; conid ia globose to su bg lobose with one pointed end , sm ooth wall ed,
nbout 3.0 - 6. 0 11m by 2 .5 - 4 .2 11m in diameter (fig . 122 , B and SE M plat e 21 ,
rop: right); eon id ial ehains forming loosely div ergent eolum ns (fig . 122, C ).
Colonies upon CYA (eolour plate 122 , B a nd b) spreading broadly , att a in ing
:1 diameter 01' 60 mm in 14 days at room temperature (20 - 25C ), similar in
Ic xture to th e preeeding, but with th e dense network 01' fIoeeose white hyphae
ovcrl ayin g th e velvety bas al felt m or e eve nly, ob seuring gre a tly th e othe rwise
a bunda ntly sp oring a reas, whieh develop reddish brown shade s with th e
ripen ing al' eonidia ; eolony sur faee sulc at ed by a few shallow furrow s in a radial
pattern , a nd centre depressed ; undulating m argin surrounded by a fringe 01'
subme rge d hyphae extending about 1 mm beyond th e ae r ial gro wth ; exuda te
ubundantly produeed as amber-eoloured drops ; odo u r none; a m ber-eolou red
pigm ent diffusing int o th e su rrou nd ing agar ; eolony reverse (eolour plat e 122,
h) in reddish sienna brown shades, a nd strongly pulling th e agar away fro m th e
I'etri di sh an d cracking it ; penieilli essentially as deseribed aboye .
C olonies upon MEA (eolour plate 122, C a nd e) spread in g broadly ,
atla ini ng a di amet er a l' 70 mm in 14 da ys at room temperature (20 -2 5C),
cons isting 01'a loos er textured basal feIt , appearing deeply velvet y a nd ove rla id
by a thin, closely woven network 01' fIoeeose aerial h yphae infIu en ein g greatly
t he eolony appearanee; ve ry th in , eve n margin surrounded by a frin ge a l'
submerge d h yphae extend ing a bou t 5 mm beyond th e aerial gro wth ; heavily
sporing throughout in pal e ye llow-gre en a nd gray gree n shades (Sgu y , PI.
XXVI , 380, a nd PI. XXVII , 404 , resp eeti vely) depending on th e degr ee 01'
ripe ness 01' eon id ia; exuda te a nd od our laeking; no eolou re d pi gm ent diffusing
int o th e su rrou nd ing ag ar; eolony revers e (eolour plat e 122, e) dev eloping
yellow-gree n to brownish shades .
The speeies deseription is ee nte re d on th e typ e stra in C BS 409 .69 = VKM

F-I ,072 - C M I 110,:HI

desert soil near Kabun,
strains were isolated by
G ermana , in th e sa me

- 1.lIIM 7061 , isolntcd hy V. C h. Bughd adi rom Syria n

Dam ascu s rcgion, Sy r iu, M arch , 1964. Two add itiona l
th e same a u thor rom Sy ria n desert so ils ncar lroa and
D am ascu s region , Sy ria, March , 1964.
Penicillium kabunicum is a quite di stincti ve spec ies cha rac tc rizcd particul arly
by its typi call y divaricat e penicilli deli cately ro ug he ne d o nly a t th e base and its
smooth wall ed metulae , by its variously colou re d colon ies in yellow, pink ,
reddish brown 0 1' gray -gree n shades, dep end in g upon th e subs tra tu m; a nd by
its rather com pa ra tively large , su bglobose to globose , smooth-walled conid ia,
conspic uo us ly pointed at one end .
It appe a rs to be mo st sa tisfactorily ass ignable to th e Asymmetricadiv arica ta su b -sec tion, but it ca n no t be placcd in a ny 01' R ap er a nd Thom ' s
( 1949) reco gn ize d series, hen ce its inclus ion here.

123. Penicillium moldavicum MiIko el BeIjak ova : fig. 123, coIour plate

123, and SEM pIat e 21, (cen tre: left) .

oo

~O O

r!?

00

~O~ g
B

yei '1
~

100

jrrn

l?

Fig. 123. Camera lucida

drawings of Penicillium
moldavicum Milko et
Beljakova, CSS 627.67. A,
Detailed drawings of the
penicilli . B, Mature conidia.
C, Habit sketches . D. Coiled
hyphae.

nicillium mo/dovicum Milko et Beljakova - Nov. Syst . Niz. Rast . Acad. Sci.
U.S .S.R. Inst. Bot. Nom. Kom arovii. 22: 255256. 1967.
M ilko and Beljakova's original diagnosis follows :
' Colonies on Czapek's agar growing rapidly, consisting of a close -textured basal
1011, more or less funiculose 2-3 mm deep, at first straw yellow coloured , becoming
it or 2-3 months pale lead gray to drab in colour, with the velvety felt shading to
polo pink orange tints, poorly sporing , brittle; colony reverse developing cinnamon ,
uru nge, to brown shades . Conidiophores not ramified or w ith 2-4 scattered
brunches (rnetulae 7), smooth walled, about 15-50 Jlm (up to 100 Jlm ) by 2-3 Jlm ,
w ith apices slightly enlarged , arising from aerial hyphae or f rom ropes of hyphae.
Tho bran ches (rnetulae) are cylindrical or club shaped, mostly 10-14 Jlm by 3-4 Jlm .
Phialides bottle shaped , about 6-10 Jlm by 2.5-3 .0 Jlm, in clusters of 3, 6 to 9 , or
ingle. Conidia are elliptical or ovoid , 3-4.5 Jlm by 2.5-3.5 Jlm, echinulate, showing
pule olive shades, arranged in short chains. Conidial structures are scattered over
11 10 colony surface . Old cultures showing sometimes brown crusts , consisting of
hrow n thick-walled cells .'
(Translated from the Russian by Prof. L. Vilas).

Ou r notes follow :
C olo n ies u po n CA (colour plate 123, A a nd a ) sp rea d in g broadly , a tta in in g a
rIi.une ter o f 65 mm in 14 d a ys a t room temperature (20- 25 C) , consisting of a
..losc-tex tu red b a sal felt, d eeply ve lve ty, overla id b y a d eeply floccos e network
11 1' h yph a e and ro pes of h yphae in flu encing greatly the co lo ny su r face, whi ch
rrm ains persist ently white , n ot furrowed, and slightly umbonate in the ce n t re;
" l'I'Y thin , eve n m arg in surrounded b y a fringe o f submerged hyphae ex te nd ing
about 5 mm be yond th e ae ria l gro w th; poorly sp oring throu gh out , not
in llu enc ing th e colo n y a ppearance ; ex udate a nd od ou r la cking ; no co lou re d
pi ~m ent d iffus in g into th e surrou n d ing a ga r; co lony reverse (co lour pl ate 123,
a) in ve ry pa le cre a m shade s; penicilli (fig. 123, A) typica lly divaricate ,
ron sisti ng of sm all clusters of m etula e of 2-3 ele me n ts in th e ve rtic il, about
7 - 13 {tm b y 3.0 - 4. 0 {tm a t the tip o f short co n id iop ho res with w a lls sm oo th,
lcw scattere d branches a t different nodes of the co n id iop ho re s, be aring
phial id es in small clu st ers o f 2- 6 eleme n ts in the ve rt icil, so me times single,
Iio rne a t different leveis from th e m etul a or th e co n id iop hores; co n id iop hores
vc ry short, n o t exceed ing 50 tut: in len gth b y 2 - 3 {tm in d ia me te r; phia lid es
ho ttle sh aped , a bou t 10-1 3 p.u: b y 3. 0-3.5 tutv, sometimes lo nge r , up to 15
I~ m , tapered to a cons p icuo usly long , narrowed n eck ; numerous co iled subm crged hyp ha e , su gge stive of as comata initia ls or nematode tr a ps , have been
obse rved by u s , not reported in the or ig inal diagnosis (fig . 123, O ) ; conidia
a l first st rongly ellip tical, w ith co ns p icuous co n nective s ad he r ing a t the tip s,
lending them a lim oni form p attern , becomin g la ter m ore or less globose,

wi th wall s stro ng ly cchin ula te (Iig. J 2 :~ , B, a nd SE M plute 21, ce nt re : lc),

abo ut 4, 5 - 5,0 j.tlll by 2.5 j.tm in thc limonif orrn co nid ia, and about :~ , O j.ttn in
th e globos e on es; conid ia l cha ins very short ami stro ng ly div ergent.
Co lon ies llpon C YA (co lour plat e 123 , B and b), gro wing Icss rapidly than
th e preceding , a ttaining a diameter of 45 mm in 14 days at room tcmpera tu re
(20-2 5C) , cons isting of a closer textured basal felt overla id by a closely wovcn
netw ork of floccose ae rial hyphae lending a cottony appe arance tu th e colon y
surface which is radiately sul cat ed by a few shallow furrows ; margin ent irc;
spa ring ly sporing underneath th e cottony overgrowth of white aerial hyph ac;
exudate abu ndantly produced as yellow drop s; odo ur lacking ; no colourcd
pi gm ent diffusing in to th e surrounding aga r; colony reverse (colour plate 12 :~ ,
b) developing pale ora nge to cinnamo n shades; colon ies strongly pulling thc
agar away fr om th e Petri dish a nd crac king it ; pen icilli essen tially a s des cri bcd
aboye .
Colonies upon M EA (colour pla te 123 , C a nd c) spreading broadly,
attain ing a di ameter of 70 mm in 14 days at room temperature (20 -25 C) ,
co ns isting of a looser textured ba sal felt , de eply lanate , overl aid by a loose
netw ork of floccose ae rial hyphae influ en cing th e colony appearance; medium
sporing throughou t in pale brownish gree n shades (S guy, PI. XXVII , 405)
partially obsc ured by th e floccose netw ork of white aerial hyphae ; very thin
m argin largely su bme rge d, extend ing abo ut 5 0 1' m or e mm beyond the a eri al
gro wth; ex uda te and odo ur lacking ; no coloure d pigment diffusing into thc
surrounding agar; colony reverse (colou r plate 123, c) developing pale orangc
to cinnamo n sh ad es; penicilli esse ntially as described aboye .
T he spec ies descr iption is ce nte red upon th e typ e strain CBS 627. 67 = C M I
129,996 = ATC C 18 ,355 = V K M F-922 = IJFM 7047, isolat ed from vin eyard soil
near Kl iskinev , S.R. M old avi a , U .S.S.R , 1958 . It is kn own only from the ty pe
strain .
Penicillium moldavicum is clearly a di stinctive species charac terized by its vcry
sim ple altho u gh typi cally d ivaricat e penicilli , its broadly spre ad ing floccose to
slightl y fun iculose colon ies ; its peculi a r coiled, submerge d , hypha e suggestivc
of asco mata initials 0 1' nematode tr aps , a nd its lim oniform a nd strongly
ec hinula te conid ia .
It appears to be m ost satisfactorily assignable to the Asymmetrica di varicat a su b-section, but it ca nno t be placed in any of Raper and Thom'x
(1949) recognized series, hence its inclusion in th e newl y erecte d P. brasilianum
senes.

124. Penicillium onobense Ramrez el M artnez: fig . 124, colour plate 124
and SEM plate 21 , (centre: right).
Penicillium onobense Ramrez et Martnez - Mycopathologia. 74. 35-49, 1981.

:,:<

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Fig. 124. Camera lucida

drawings of Penicillium
onobense Ramrez et
Martnez, ATCC 42,225. A,
Detaled drawings of the
pencilli. B, Mature condia.
C, Habit sketches.

Colonies upon CA (colour plate 124, A and a) spreading broadly, attaining a

diameter of 60 mm in 14 days at room temperature (20- 25C), consisting of a
tough, rather leathery basal felt, velvety in margin , radiately wrinkled, dirty
white coloured, with margin evenly yellow, surrounded by a fringe of
submerged hyphae extending 2-3 mm beyond the aerial growth ; poorly
sporing throughout ; exudate and odour lacking; no coloured pigment diffusing
into the surrounding agar; colony reverse (colour plate 124, a) in cream
shades, with margin yellow; penicilli typically divaricate, borne terminally on
ascending stipes arising from trailing hyphae , up to 300 Ilm in length by
2 .5 -3 .5 Ilm in diameter, with walls delicately roughen ed , or on short branches
arising from aerial hyphae, commonly 10-50 Ilm by 2.5-3.0 Ilm (fig . 124, A),
varying in complexity from simple verticilis of phialides (appearing then
monoverticillate) to verticils containing both metulae and phialides , or verticils
of metulae of unequal length, and occasionally larger structures with metulae
and phialides borne upon one or more branches in addition to the main axis ;
branches variable , ranging from 15 to 20 Ilm in length by 2.0- 3 .5 iut: in
diameter; metulae in verticils of 2 - 3 elements in the whorl , finely roughened ,

mostly 10-1 :3 j,tm by :3.0 - 5 .0 j,tm wit h a pices ITlOI'C 0 1' lcss vesiculoso: phi alides
div er gent in crowdc d clus ters 01' :3- 6 clemc n ts pcr vcrti cil , conspicuo usly
tapering to a definite short narrowed ncck , bottl c sha pcd , common ly
measuring 8 .0-1 2 j,tm by 2.8- 3.5 j,tm ; conid ia stron gly elliptical to subg lobosc ,
with wall s roughen ed, with roughness a rra nged in spiral bands as Penicillium
brasilianum Bati sta ; con id ial chains a t first loosely diver gcnt , bccoming tanglcd
with ag e (fig. 124, C ).
C olonies up on CYA (colour plat e 124 , B and b) spreading broadl y , a tta in ing
a diamet er 01' 65 mm in 14 days a t room temperature (20 - 25C) , finc1y
velve ty, simila r in texture to the preceding bu t more irr egul arl y wrinkled by
numerous furrows; thin eve n margin with sub me rged hypha e extend ing about
2 mm beyond the ae r ial growth; spor ing ab unda ntIy in marginal zones in
brownish shades (Sguy , PI. XLVII , 694) ; non- sporulating areas in palc
pinkish sha des; exudate limited in a mo u nt, colourless; odour " mo uldy"; no
colou red pigment diffusing into the su rround ing agar; colony reve rse (colou r
plat e 124 , b) devel oping yellow ochre to pale ora nge sha de s; penicilli
essentially as described abo ye .
Co lon ies up on M EA (co lour plat e 124 , C a nd c) at ta ining a di ameter 01' 60
mm in 14 days at room temperature (20-2 5C) , velvety, plane, surfaec
appearing gra nular owing to the sea tte red tufts 01' white floceose ae r ial hyphac
a rising from th e basal felt; ve ry irregul arly dissected margin surrounded by a
frin ge 01' stol on -like subme rge d orange hyphae ex tend ing 10 01' mor e mm
b eyond th e ae rial gro wth; heavil y sporing throu ghout in blue-gray shades
(Sg uy, PI. XL , 598) : exudate and odour laeking; conid ial structures ar ising
eithe r from a netw ork 01' aerial hyphae 01' from the stolon -like submerged
hyphae, otherw ise, essentially as described aboye .
Penicillium onobense description is cente red up on th e typ e strain IJ FM 3026 =
AT C C 42,225 an d several othe r strains, isolat ed fro m a n a ndosol under beech
forest , in th e provi nce 01' N av a rr a , Spain, by DI' A .T. M a rt nez, April , 1974.
Penicillium onobense is quite a distineti ve spee ies character ized by its typ ically
di varicate pen icilli , by its bottIe-sh aped phi alides with cons picuo us tap ering
sho rt a nd narrow necks, by the production 01' ora nge stolon- like subme rge d
hyphae upon MEA , by its spirally rou ghen ed elliptical con idia , and by its
broadl y spread ing colon ies up on all substrata . The div aricat e charac ter 01' its
fru iting structu res warrant placem en t near the Asy m me tr ica - d iva ricata
altho ug h it ca nnot sa tisfactorily be ass igne d to any 01' R ap er and T ho m' s
(1949) well-recognized series, hen ce its inclu sion here .

125. Penicillium castellonense R amrez el M artnez : fig. 125, colour plate

125, and SEM plate 21, (bottom : left).

Penicillium castellonense Ramrez et Martnez - Myeopathologia. 74. 35-49, 1981.

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,

,
i
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Fig. 125. Camera lucida

drawings of Penicillium
castellonense Ramrez et
Martnez, ATCC 42,229. A,
Detailed drawings of the
penicilli. B, Mature conidia .
C, Habit sketches .

C olonies upon CA (colour plat e 125 , A and a) a ttaining a diamet er of 45 mm

in 14 days at ro om temperature (20 -25C), co ns isting of a closc-textu rc d basal
Ielt , ove rla id by a closely woven net work of wh ite ae ria l h yphae lending a
lanat e a ppe a rance to th e colo ny su rface , radiat ely sulca ted with shallow
Iur rows; undulating m argin su r ro u nded by a fringe of sub me rge d hyphae
cxtend ing a bo u t 3 mm beyond th e aerial growth; heavily spo ring throu gh out in
blu e-gr een shade s (Sguy, PI. XXX , 438), but with co nid ial are as totally
obscu red b y the dense overgrowth of ae ria l hyphae ; exuda te ab u nd a n tly
prod uce d a s large pale amber-coloured drop s; odo u r lacking; pale ambercolou red pi gm ent diffusing into th e surrounding agar; colony reverse (co lou r
plat e 125 , a) d eveloping a t first pale yellow shades, becoming reddish in age;
penicilli typi call y divaricat e, variously branched , rangin g from sing le terminal
vertic ils of phialides (appearing m onoverticill at e) to asym me tric str uc tures
composed of ver tic ils of metulae bearin g clu sters of phialides (fig . 125 , A);
conid iophores smooth walled, commonly arising fro m th e su bstratu m, sep ta te ,
up to 300 tut: in length by 1.8-2.5 tut: in di amet er , or as sho rt branch es from
ae rial h yphae ; m etulae in clusters of 2 - 4 elements in th e ve rt icil, m ostl y 10-15

2.0- :U) /lm, with apices e nla rge d in a vesicle up lo 4 .5 /lrn in di arn ct er:
ph ialides div ergenl , in clu ster 01' :3- 10 c1ern ents in the vcrticil , enlargcd al thc
b a se th en consp icuo us ly tapering abruptly to a sho r t na rrow ed neck, mo stly
7- 9 /lm by 2 .5 -4 .0 iuu ; co n id ia globo se, cons pie uo us ly sp inulose (fig . 125, B,
a nd SEM plate 2 1, b ottorn : left ), co m rno nly 3. 0 -3.5 /lm in di amet er; eon id ia l
chains typicall y adhe r ing in loose di vergent well -defined colu m ns (fig . 125 , C ).
Colon ies upon CYA (co lo ur plat e 125, B and b) a ttain ing a di ameter 01' 50
m m in 14 d ays a t room temper ature (20-25 C), co ns isting 01' a simila rly
tex tured b a sal felt as described a boye , but with d eep er radial furrow s;
remaining persistently white ow ing to th e deep close ly wove n net work 01' whi tc
ae rial hypha e overl aying th e basal felt; poorly spo r in g ; m argin e nti re; exu date
as pal e ye llow drops; odo ur lack in g; a m be r-co lou re d pigment diffusing into th e
su rrou nd ing a gar ; colony reverse (co lou r plate 125 , b) developing pale ye llow
to cream shade s; penicilli esse ntially as described a boye .
Colo n ies upon M EA (co lour plat e 125 , C and e), sp read ing b ro adly ,
a ttain ing a di ameter 01' 50 mm in 14 d ays a t room temperature (20 -25 C ),
co ns isting 01' a looser textured basal felt , ve lve ty to loosely lanate in m arginal
a reas which rem a in persistently wh ite; su b margin al a nd ce n tral a reas typi cally
ve lve ty, zo na te, n ot fu r ro we d, plane , h eavily sporing through out in deep bluegreen shades (Sguy, PI. XXX , 447 ); ex u date a n d odo ur lacking ; reve rsc
co lou rless w ith a gr een cast sho w ing throu gh (co lou r pl at e 125 , c); n o colou red
pi gment diffusing into the surrou n d ing agar; penicilli esse nt ia lly as described
aboye b ut with con id ia l ch ai ns form in g m ore compact di vergent colu m ns .
Species description is ce n tere d upon the typ e strain IJFM 5 144 = AT C C
42, 229 , isolated fro m th e a ir 01' M adrid , Spain, in May, 1977, by th e au th or
(C . R .). It is at present known on ly fro m th e typ e strain .
Penicilliu m castellonense is a distin ctive species charactc r izcd by its typi cally
di va rica te pen icilli , sho r t-poin te d phialides, b ro adly sp read ing colon ies upon
m ost su bstrata, spin u lose glob ose co n id ia, a nd by its both floccose a nd velve ty
characte r 01' th e colon ies upon MEA .
It a ppears to be m ost sa tis fac to r ily ass ignab le to the Asymmetricadi varica ta su b-sec tion , but it ca n no t be placed in a ny 01' Raper a nd T ho m ' s
( 1949) recogn ized series, hence its in clu sion in th e P. brasilianum se ries .

/l m b y

Species of dubious position

Penicillium glauco-cinerascens Chalabuda - Nov. Syst. Niz. Rast. Aead. Sei.
USSR. Inst. Bot. Nom. Komarovii. 6: 165. 1950. See also M.M. Kulik . ' A Compilation of Deseriptions of New Penicillium Species'. Agrie. Handbook No. 351.
Agrie. Res. Ser. U.S. Dep. Agrie. p. 41.1968.

Penicillium gla uco-cinerascens C halab u da is a spec ies descr ibed in 1950 th at is

hclicvcd to he closcly rc1at ed to spec ics included in thc ASYlIllIlc lric a .Iivnricat a sub -sec tion. H ow evc r, it is pr escnt cd hcre as a spec ics of dubi ous
posit ion and ha s not been inc1uded in th e keys lo spcc ies sincc the au tho r of th e
prcsc nt study was not able to exam ine anc1 st udy th e type strain of th e spec ics
in qu estion because a11 a tte rn p ts to obtain it from Internati on al C u lture
(lollcctions failed .
Chalabuda's original description follows:
'Colonies upon Czapek's agar tanate . growing rather restrictedly, developing
live-gray tints; exudate as crystal c1ear yellow drops; colony reverse developing
orange colours.
Colonies upon sugar beet root extract agar lanate, more or less circular, at first
creamy white, becoming ashen-olivaceous; exudate as numerous minute crystal
clear uncoloured droplets; colony reverse orange -brown; conidiophores 90-100 JLm
by 2 JLm , with walls commonly delicately roughened, w ith terminal divaricate
branches; phialides 7 JLm by 1.6-2 JLm ; conidia ellipsiod, smooth walled, 2.5-3.0 JLm
by 1.8-2.0 JLm in diameter, appearing yellow olivaceous in mass. The type strain
was isolated from soil in the environs of Kiev.'
(Translated from the Russian by Prof. Or. L. Vilas).

Penicillium impar Von Szilvinyi - Zentbl. fr Bakt. Parasitenk . Infek . Hyg. Abt. 11.
103: 164. 1940-41. See also M.M . Kulik, 'A Compilation of Oescriptions of New
Penicifium Species'. Agric. Handbook No . 351. Agric. Res. Ser. U.S. Oep.
Agric. p. 44. 1968.

Penicillium impar vo n Szilvin yi is a spec ies described by A. vo n Szy ilviny i in

1940 that is b elieved to be clo sely re lated to th e spec ies inc1ude d in the
Asy mme tr ica - d iva rica ta su b -section . H owever , it is prese nted here as a
spec ies of dubiou s positi on a nd ha s not been in cluded in the keys to spec ies in
t he curre nt study for the sa me reasons as above .
Sylvinyi's original diagnosis follows:
' Colonies upon Czapek's agar are circular after 7 davs, delicately lanate to
velvety, developing dark green colours, in comparison with the margin which is at
first white , soon becoming gray-green, abruptly raised. Exudate abundantly
produced, as large crystal c1ear drops; the oldest areas beco me yellow-green to
bright brown coloured in 10 davs, overlaid by a loosely woven network of floccose
fine hyphae . Exudate golden yellow in places; after 14 days the colonies are
delicately lanate to deeply floccose , dark to bright yellow-green coloured, with
white margino Exudate abundantly produced. Colony reverse pale yellowish.
Somewhat zonate . Colonies attaining a diameter of 10 mm in 7 days , 25 mm in ten
days. Conidiophores smooth walled, hyaline, 3-4 JLm in diameter, with penicilli

divaricata; rnetula e 5 by 2 .tm. Phialidas 5 by 2'Lm. Conidia smooth, globosa, 2.5 .tm
in diametor, producinq chains loosely divergent . Minimum temperature of growth
goC, optimum temperature 26C, and the maximum temperature is 26C to 36C.'
(Translated from the German by the author).

Penicillium vitale Pidoplichko et Bilai - In. V .1. Bilai's 'Antibiotic-Producing Microscopic Fungi', Kiev, p. 44. 1961. See also M.M. Kulik, 'A Compilation of
Descriptions of New Penicillium Species'. Agric . Handbook No . 351. Agric. Res.
Ser. U.S . Dep. Agric. p. 79. 1968.
Penicillium vitale Pidoplichko et Bilai is a sp ecie s described in 1961 whi ch is
believ ed to be closel y rel ated to th e species bel on ging to th e Asymmetricadi varicata sub-sect ion . However , it is presented here as a sp ecies of dubiou s
p osition a nd has not b een included in th e keys to species of th e cu r re n t study
for rea sons id entical to th ose for th e two p receding species .
Pidoplichko and Bilai's original diagnosis follows:
'Colonies upon malt extract agar at first white, becoming then blue-green ,
velvety, c1ose-textured and furrowed, with a rather broad white margin, developing
greenish dirty gray shades in age. Colony reverse at first straw yellow, turning to
yellowish cinnamon in the centre and yellow in the margin, shading to cinnamon in
age. With agar turning green in colour with the growing culture. Exudate as
numerous yellow drops on the surface of young colonies.
Vegetative hyphae white. Coniophores are smooth walled, 2-3.5 .tm in diameter,
with penicilli in c1usters, with phialides forming whorls at the tips of the
conidiophores (appearing monoverticillate) or more commonly in verticils of 2-3
metulae at the end of the conidiophores. Phialides are bottle shaped, 6-g .tm by
2.0-2.5 .tm, usually in more or less compact c1usters of 2-5 elements in the verticil.
Metulae measuring g-10 .tm by 2-3 .tm. Sometimes penicilli arising from a single
divaricate branch borne at a different level on the conidiophore, being then
frequently longer. Conidia elliptical or more often ovoid when young, 2.5-3 .tm by
2.0-2.5 .tm; becoming more or less globose, broadly elliptical, round-ovoid ,
2.5-4.4 .tm by 2.6-3.0 Lm , smooth, green in mass, forming loosely parallel to
tangled chains.
The maximum temperature growth on malt agar is 37-3goC .
Colonies on Czapek's agar plus 15% gelatin and glucose growing more slowly,
greenish blue, with green shades predominating, velvety, with rather wide white
margin, usually with abundant exudate as golden yellow drops in conidial areas,
turning later on dirty gray in colour; colony reverse in olive cinnamon shades,
becoming darker in age. The gelatin is similarly coloured in olive shades and
Iiquefies with difficulty. The gelatin starts to Iiquefy during the second week of
growth (at 23-25Cl. The Iiquefied gelatin has an alkaline reaction.

."".1
olonies upon Czapek's agar plus suerosa are at first white , then blue-green,
velvotv, with wide white margin, turning dirty gray in age. Colony reverse at first
traw yellow, turning to yellow-cinnamon shades. Green-coloured pigment
:lIHusing int o the surrounding agar. Exudate abundantly produced as yellowish
rops in young colonies .'
(Translated from the Russian by Prof. L. Vilas).

nici//ium velutinum 'Terui et Shibazaki (not P. velutinum van Beyma) - J.

erment. Technol. 26: 257. 1948. See also M .M . Kulik , 'A Compilation of
Descript ions of New Penicillium Species ' . Agric. Handbook No . 351. Agric. Res.
Ser, U.S. Dep. Agric . pp . 77-78. 1968.

Penicillium velutinum T erui el Shibazaki is a spec ies described in 1948 whi ch is

l u-li cved to b e closely related to sp ecies incl uded in th e As ymmetrica .Ii vur icata sub -sec tion . H ow ev er, it is presented here as a spec ies of dubious
pI/sil ion

a nd has not been included in th e keys to sp ecies of th e cu r re n t st u dy

lur the sa me reasons as for the 3 precedin g species .

Terui and Sh ibazaki's diagnosis follows :

'Colonies upon Czapek's agar are perfectly circular, showing green colours in 2-3
dDYS at 30C. The felt is velvety becoming ashen blue-green in zones with age .
Colony surface mostly flat with wide and white margin 1-2 mm in width. Closelext ured; colony reverse at first yellow, becoming reddish brown. Exudate at first
colourless, shading to yellow, becoming gradually peach to brownish yellow . Culture basic colours change about in the same way. Odour lacking . Sclerotia not
observed.
Penicilli are of two types , either monoverticillate or asymmetrical ; the latter are
more abundantly produced. Conidiophores smooth walled, up to 200-300 .m by
2.5-3.0 .m, seldom branched, with apices enlarged in a vesicle up to 3.5-4.5 .m .
Phialides in clusters of 3 elements in the verticil (seldorn 2 or 4), either direct ly at the
tip of the conidiophores (appearing monoverticillate) or borne at the apex of
metulae, generally 2 elements per metulae (seldom 31. Metulae are of unequa l
length in the verticil, usually those measuring 15 .m in length are the most abundant
on es. They usually have a diameter of 2.5-3.0 .m at the base and 3-4.5 .m at the
apex. Phialides mostly 8-12 .m (tne middle is wider, 2.4-3 .2 .m , than the t ipl, not
tapering to an abruptly pointed neck . Atypical ph ialides are differently shaped and
mostly asymmetrical atta ining a length of 20 .m . Also sometimes a metula may bear
a single phialides, and sometimes the single phialides may become a secondary
metula, bearing secondary phialides .
Matu re conidia are globose, smooth walled , pale ash green coloured . They are
2.7-3.0 .m in diamete r.
This fungus produces cit rinin and was isolated from t he air. It shows penicilli

either typically monoverticillate or typically of the Asymmetrica -divaricate type .'

(Translated from the Japanese by Miss Masae Yamamotol.

T he a bo ye description was published without providing a Lat in d iagnosis .

haptcr 5

Asyrnmetrica-velutina sub-section

'1'111 5 SUB-SECT lN INCLUDES TH SE SPECIES F PENICILLIUM WITH ASYMMET RIC

pcni cilli developing, ch a rac te r istically , into a dense and eve n st and so as to
produce a velvet y appearance , in wh ich th e vegetative mycelium is largely or
torally sub m er ged in th e substratum , a n d the aerial portion of the colon ies
run sists of erect conidiophores arising directly from the substratum like a field
01' cereal. Actually, th e term velvety is broadened enou gh here so as to include
1he m any gradations that m ay be enc ou n te red, encompassing those species
closely related but with conidiophores arising from a basal network of interwoven aerial hyphae. Flocco se , lanate, funi culose , a nd fasciculate or
xynnem a tou s forms are exclu ded from this sub-secti on .
Six series were recognized by Raper and Thom (1949) in th e Velutina and
maintained in the present st u dy as follows : Penicillium citrinum se r ies, ch a r acicri zed by as ymmetric penicilli con sistin g usually of terminal verticils of
metulae , bearing phialides producing con id ia l chains typically ad he r in g into
d ivergent columns . Penicillium chrysogenum series , ch a racte r ized by penicilli that
are often ramified several times below th e level of the m etulae , with con id ial
cha ins com m on ly adhering into well -defined colu m n s, a n d by colo n ies typica lly producing bright ye llo w pi gments in ex udate a nd colony rev erse. It is
represented actually by the sin gle species of th e same name , because th e
members of th e series recognized b y Raper a n d T ho m (1949) , namely : P.
meleagrinum Biourge , P. notatum Westling , and P. cyaneo-fuloum Biourge, h ave
been reduced to synonymies with P. chrysogenum Thom (Samson et al. 1977) .
The species is well kn own for the production of th e a n tib iotic penicillin .
Penicillium oxalicum series , ch a r acter ized by the str on gly elliptic a l conidia which
in new isolates ofte n form heavy crusts , which ch aracteristically fall away en
masse if th e cu ltu re dish is tapped . Penicillium digitatum series , actuall y
represented b y two species : P. digitatum Sa ccardo a n d P. [aponicum Smith . The
former sp ecies b eing responsible for th e so-called 'green rot' of citrus fruits .
Penicillium roquejorti series , including th e moulds u sed for th e production of
Roquefort-type cheeses, ch a r ac ter ized by colo n ies with a rach noid m argin and
by con id iop h ore s with walls conspicu ously roughened . Penicillium breuicompactum ser ies, ch a r ac te r ized b y its unusu all y co m pact p enicilli in which a ll
ele men ts a re closel y compacted and appres sed , b y th e rather restricted gro wth

01' th e colon ies up on certain suhs tra ta , ami by th e develop mcnt 01' stolon -lik..
submerged hyphac along th e colony m argin 01' most forms up on ccrta in substrata .

K ey to series and species

Paga
1. Penicilli seldom ramified below the level of metulae, with metulae often
more or less divergent.
P. citrinum series . ... 349
A. Metulae typically in verticils of 2-3 elements in the verticil , often of
unequal length; conidial chains not forming well-defined columns;
conidia elliptical to subglobose.
P. corylophilum Dierckx .. .. 353
B. Metulae typically in verticils of 3-5 elements in the verticil, bearing
phialides in crowded c1usters and conidial chains usuallv in welldefined columns; conidia globose to subglobose.
1. Conidial areas in bright blue-green shades; colony reverse
developing bright yellow to orange-pink shades; usually producing
citrinin.
P. citrinum Thom . . .. 351
2. Conidial areas developing dull blue-green or yellow-green shades;
colony reverse showing dull yellow to buff shades; not producing
citrinin .

P. steckiiZaleski ... . 355

3. Conidial areas developing gray-green shades; colony reverse in
reddish brown (sienna) shades; not producing citrinin.
P. matriti Smith .... 357
C. Metulae typically in verticils of 5 to 8 elements in the verticil, closelv
compacted; conidial chains not in well-defined columns.
P. paxilli Bainier . ... 403
(See P. brevi-compactum series)
11. Penicilli typically rebranched below the level of metulae, with the main
axis and branches terminating in verticils of metulae.
A. Penicilli commonly long, with elements loosely arranged and often
divergent .
1. Conidiophores smooth-walled; margin not appearing arachnoid.
a. Colonies typically producing abundant yellow pigment in exudate
and colony reverse; conidial chains often adherent into well
defined columns; conidia less than 5 ,tm in long axis; elliptical
to subglobose.
P. chrysogenum series
360
P. chrysogenum Thom
360

," 1' /

b. Colonies not producing yellow plgment In exudate or colony

reverse; conidia commonly 5.0 ~m or more in long axis.
1', Conidia elliptical or subglobose, fairly uniform in size;
common in soil.
P. oxalicum series .. .. 366
aa. Conidia smooth walled.
1". Conidia elliptical; colonies plane or nearly so, with
conidia often forming deep crusts, conidial chains
appearing 'silky' when viewed under low power.
P. oxalicum Currie etThom . . . . 367
2". Conidia subglobose; colonies radiately furrowed, with
conidia not forming crusts; colony reverse upon CYA in
deep cherry red, also diffusing into the surrounding agar.
P. atramentosum Thom .... 369
bb. Conidia conspicuously roughened ; colony reverse
developing deep orange-red shades.
P. fennel/iae Stolk . ... 371
2' . Conidia strongly elliptical to cylindrical, varying greatly in size
and often very large; penicilli very irregular often fragmentary.
P. digitatum series . ... 374
aa. Colonies upon CA growing but a little, 1 or 2 mm in diameter, mostly submerged; colony reverse not developing
deep purple red shades; produces 'green' citrus fruit rot.
P. digitatum Saccardo . . .. 375
bb. Colonies upon CA growing restrictedly but attaining at
least 15-20 mm in diameter in 14 days; colony reverse
developing deep purple brown shades upon all substrata;
does not produce 'green' citrus fruit rot.
P. japonicum Smith .... 377
2. Conidiophores conspicuously rough-walled; margin of colonies
appearing usually arachnoid.
P. roqueforti series .... 381
a. Conidiophores coarsely roughened, usually appearing conspicuously warty.
1'. Colonies broadly spreading upon all substrata, with surface
plane or nearly so; margin thin, appearing arachnoid; conidia
comparatively very large, attaining a diameter of 6.5 ~m .
P. roqueforti Thom .... 383
2'. Colonies growing restrictedly upon CA; conidia not exceeding
4.5 ~m in diameter.
P. farinosum Novobranova ... . 391

,r n l

b. Conidiophores not coarsely roughened but with walls appearing

finely granular; conidial areas in pale yellow-green to artemisia
blue-green shades; conidia comparatively small, not exceeding
2.5 tm in diameter.
P. fagi Martnez et Ramrez .. . . 385
c. Conidiophores not coarsely roughened but with walls appearing
finely spinulose.
1'. Conidial areas upon M EA developing olive green shades;
conidia large, up to 5 tm in diameter.
P. mali Novobranova . . . . 389
2'. Conidial areas upon MEA developing deep blue-green shades;
conidia not exceeding 4 tm in diameter.
P. cordubense Ramrez et Martnez .... 387
B. Penicilli comparatively short, with all elements closelv appressed.
P. brevi-compactum series . . . . 394
1. Penicilli typically showing one or more side branches below the
level of metulae.
a. Conidia smooth-walled or nearly so.
P. volgaense Beljakova et Milko . . . . 405
b. Conidia conspicuously roughened.
1' . Conidiophore walls conspicuously roughened.
P. brevi-compactum Dierckx . . . . 396
2'. Conidiophore walls smooth or nearly so .
aa. Conidia strongly echinulate; conidial areas developing dark
brown colours.
P. brunneo-stoloniferum Abe ex Ramrez ... . 412
bb. Conidia delicately roughened; conidial areas developing
ochre-yellow shades.
P. arenicola Chalabuda . ... 410
2. Penicilli typically consisting of a single, crowded, terminal verticil of
5-8 metulae.
a. Conidiophore walls conspicuously and coarsely roughened, at
least at the base; conidia elliptical , conspicuously roughened in
spiral bands.
P. skrjabinii Shmotina et Golovleva .... 407
b. Conidiophore walls smooth or nearly so .
1'. Colonies growing restrictedly upon CA and MEA; conidia
mostly globose.
P. stoloniferum Thom . ... 400
2' . Colonies growing more rapidly upon CA and MEA ; conidia
elliptical.
P. paxilli Bainier .... 403

,l " ':}

l'euicillium citrinum seri es

UIJ 'J'S'J'A NDl NG F EATU R':S

C olonics grow ing rathcr restrict cdl y upon C za pek's agar, thin bu t va ry ing
in dcpth anel tcx tu rc dep cn ding upo n th e spcc ies and strain, with conid ial areas
uppca ring velve ty , m or e or less heavily sporing , developing blue-green ,
yellow-green to olive gree n shades, often prod u cing exud ate more or less
ihu nd antl y , ra nging fro m co lou rless to brigh t yellow; colony reverse u suall y in
ycllow , ora ng e to redd ish b ro wn shades, in one species becom ing almost black.
C onid iopho res erec t , com paratively short, u su all y not exce eding 200 Jlm in
k llgth, ar ising d irectly fro m th e su bs tratu m or the basal felt in a dense sta nd,
lcss requen tly fro m aerial hyphae in more floccose strains, sm ooth walled,
usu ally not ra m ified, excep t a t th e ti p o
Pcnicilli typ ically cons isting of a n ap ica l cluster of 3 - 5, or occas ionally more
di vcrging m ctula e , eac h su p porti n g a crowde d cluster of p hialides taper ing to a
xho rt narrowed neck . Conid ial cha ins com monly adhering in to we ll-d efined
:l I I div erge nt colu m ns.
Conid ia com para tively small , not exceed ing 3.5 Jlm in d iameter, globose to
r-lliptical de pendi n g upon the species and st ra in, with wall s smooth or nea rly
so .

K cy to species
Page
1. Penicilli seldom ramified below the rever of metulae , with metulae often
more or less divergent.
P. citrinum series . . .. 349
A. Metulae typically in verticils of 2-3 elements in the verticil, often of
unequal length; conidial chains not forming well-defined columns;
conidia elliptical to subglobose; colony reverse developing almost
green-black shades in certain areas.
P. corylophilum Dierckx .... 353
B. Metulae typically in verticils of 3-5 elements in the verticil , bearing
phialides in crowded elusters and eonidial chains usually in welldefined columns; conidia globose to subglobose.
1. Conidial areas in bright blue-green shades; colony reverse developing bright yellow to orange-pink shades; usually producing eitrinin.
P. citrinum Thom . . . . 351
2. Conidial areas developing dull blue-green or yellow-green shades;
colony reverse showing dull yellow to buff shades; not producing
citrinin.
P. steckii Zaleski . .. . 355

3. Conidial areas developing gray -green shades; colony reverse in

sienna reddish brown shades; not producing citrinin.
P. matriti Smith ... . 357
C. Metulae typically in verticils of 5 to 8 elements in the verticil, closelv
compacted; conidial chains not in well-defined columns.
P. paxilli Bainier . .. . 403
(See P. brevi-compactum series)

T he princip al m embers of thi s seri es are abu nda nt a nd world wid e in

di stribution . They are fre que n tly isolat ed from so ils and frorn deteri orating
military eq u ipme nt, particul arly in tr opi cal an d subtro pica l zones.
C olonies gro wn on Czapek ' s agar vary in texture, colour, and in th e
abu ndance of coni d ial structu res, dep endin g upon th e species and strain. T hey
are generally restricted in Penicillium citrinum Thom and P. steckii Zaleski; more
or less ra d iately furrowed with th e exce ption of P. matriti Smith , which presents
a n irregularly wrink1ed su rface . Upon m alt ex trac t aga r th e colonies vary
greatly dep ending upon th e spe cies : in P. citrinum Thom, th e colon ies are
cons iste ntly restricted a nd gene ra lly smaller th an on C za pe k ' s agar, not
furrawed, heavily spo ring in very deep blue-green colou rs (Sguy , Pi. XXX,
446), com mo nly up to 300 p.m or m or e deep , and with reverse typi call y in
orange-yellow sha des . In P. corylophilum Dierckx , th e colonies a re broadly
spreading , thinner , with reverse in ligh t brown-gr een to very dark shades,
almost black in local ized areas and secto rs. In P. steckii and in P. matriti , the
colonies tend to be int ermediate bet ween the preceding , both in rat e of gr owth
and in cultu ral aspecto In P. steckii , colony reverse develops light yellow shades
to orange -ye llow in localized area s, while in P. matriti , th e development of
sienna reddish brown colou rs in th e colony reverse is cha racteristic .
H owever, th e outst anding di agn ostic characte ris tic of th e series is ce ntere d
on th e di stinctive charac te r of the penicilli. These cons ist ofterminal clusters of
di verging m etuale , arising from unbranch ed conidio pho res . The phialides are
com paratively sho rt, tapering to a short narrowed neck ; th e conid ial chains
tend to adhere in fairly com pa ct colum ns.
Penicillium matriti Smith has been added in th e present study to th e three
species recognized in this seri es by Raper an d Thom (1949) , namely :
Penicillium citrinum Thom , P. stecki i Zal eski , and P. corylophilum Di erckx.
Penicillium citrinum is by far the m ost abundant and th e mo st variable of the
four species .
The P. citrinum series is beli eved to be formed by a gr oup of transiti on al
spe cies , intermediate between th e monoverticillate forms of th e Monoverticillata-ramigena on th e one hand and Penicillium chrysogenum Thom on th e
other.

12G . Penicillium citrinum Thom: ligo 126,

CO IOlll'

platc 12G, am i SEM

plate 21, (bottorn : lct ).

000
B

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Fig. 126. Camera lucida

drawings of Penicillium
citrinum Thorn, CBS 139.45.
A, Detailed drawings of the
penicilli . B, Matu re conidia.
C, Habit sketches.

Penicillium citrinum Thom - Bull. Bur. Anim. Ind. U.S . Dep. Agric. 118: 61-63.
1910. Emmended by Thom in 'The Penicillia', pp. 256-257. 1930; also Raper and
Thom , 'A Manual of the Penicillia ' , pp. 345-350. 1949.

Synonym
Penicillium aurifluum Biourge - Monographie, La Cellule. 33: 250-252. 1923.

Colonies on CA (colour pIate 126, A and a) growing restrictedly, attaining a

diameter of 20 mm in 14 days at room temperature (20-25C), consisting of a
close-textured basal felt , ranging from velv ety in most strains, more or less
floccose in sorne, to almost lea thery in others , radiately furrowed towards the
central area in crateriform pattern; thi n even margin surrounded by a fringe of
submerged hyphae extend ing about 2 mm beyond the aerial growth; more or
less heavily sporing, depending upon the strain, with zonation more or less

cvidc nt in so rne strni ns , not in o thcrs; conid ia l a rcas a l firs: in pal c blu c-grcen
shaclcs (Sguy ; PI. XXVII , 4(0), bccoming a rtc m isia grce n al maturi ty
(Sguy, PI. XXVll , 41B), bccorning 1lI 0U SC gray lo dce p olivo gray (S guy , PI.
XXIX , 428) in age; ex udate a bu nda n t as pale yellow lo stra w-colo u red
d ropl et s; odou r strong, ' mouldy' in sorne stra ins, faint in othe rs: ycllow to
ora nge -colou re d pigmcnt di ffu sin g in to the su rrou nd ing ag a r; eolo ny reverso
(colour pIa te 126, a) in ora nge shades 01'ten assu m ing a pinkish tin t; penicilli
typi eall y eo ns isting 01' a terminal who rl 01' 3, 4 , or occasionally m ore , some what
di vergent m etulae , mostly 15-28 p.m b y 2.5- 3 .0 p.m , with a pices enlarged in a
ves icle up to 4 p.m in di ame ter (fig . 126, A) ; eon idiopho res smooth walled ,
a rising m ostIy from th e su bstratu m or from a erial hyphae in th e colony ce n tre
or in /loeeose strains, usu all y 50-200 p.m in length by 2.5-3 p.m in di ameter;
phi alides more or less erowde d a nd pa rallel , in clu sters 01' 6- 10 or more
elemen ts in th e ve r ticil, a bo u t 13-1 7 p.m b y 3-3 .5 p.m in sorne st ra in s, or 8 - 11
p.m by 2-2 .8 p.m in others, usu all y tape ring to a co ns picuo us sho r t n arrow ed
neck; eon id ia globose to su bg lobose, mostly 3-3 .8 p.m , smooth -wa lled or
nearl y so (fig . 126, B a nd SEM plate 21, bottom : ri gh t) .
Colonies upon C YA (co lou r plat e 126, B and b) gro wing more rapidly th an
aboye, a tta in ing a dia meter 01' abou t 35 m m in 14 d ays at room temperature
(20 - 25 C ), eo nsis ting 01' a tough, close-textured basal felt, typieall y velvety,
eons pieuously zo nate, strong ly radi at ely wrinkled an d raised in th e co lony
cen tre in a erate riform pa ttern ; white eve n m argin surro u nded b y a narrow
fringe 01' su b merged hyphac exte nd ing abo ut 1 mm beyon g th e ae ria l grow th;
heavily spo ring in zo nes , vario us ly coloured in blue-green (Sguy, PI. XXX,
442), blu e-gray (Sguy, PI. XXXII I, 493) or brow ni sh shades, d ep ending
up on th e d egree 01' m aturity 01' eonidial areas ; ex udate a b u ndan tly produced ,
as light yellow d ro ps; odou r fain t , aro matic; ambe r-coloured pigment di ffusi ng
into th e su rro u nd ing agar; eolo ny reverse (co lo u r plate 126, b) in deep ora ngered shades; penieilli esse ntially as deserib ed aboye .
C olon ies upon MEA (co lour pla te 126, C and e) gro wing m ore restrictedly
th an up on CA , a ttain ing a d iamet er 01' less th an 20 mm in 14 d ays a t room
temper a ture , not furrow ed , velvety, with cen tre sligh tly urnbonat e; with
undulati ng, thin m argin surrou nde d by a narrow fringe 01' submerged hyphae
ex te nd ing abou t 1 mm beyond th e aeri al grow th ; heavily sporing th rou gh out in
deep blu e-green shades (Sguy , PI. XXX, 446) ; ex udate laeking; odo ur faint ,
aromatic; golde n yellow pigment diffu sin g in to th e su r ro u nd ing agar ; col on y
reverse (eolour plate 126, e) in bright ora nge-yellow shade s; penicilli esse n tially
as described above ; co nid ia l chains in eom paet colu m ns (fig . 126, C ).
T he spec ies description is cen te re d upon th e typ e stra in CBS 139.45 = C M I
92, 196 = NRR L 1,841 = IJFM 3755, isolated by Ch. T ho m in 1910, and many
othe r strains isolated by th e author from Sp anish soils .
R ap er and Thom' s eonce p t 01' Penicillium citrinum is broad in seope and

includcs lorrn s whi ch va ry su bstantially in pa rt icu lar cha rac teristics, yet
posscss ccrta in basi c featurcs in cornrno n ( 1949) . In thcir opin ion , it seemcd
Hall: l o bclievc th at 75 % 01' thc strains tha t produce sorne citrinin and show
pcni cilli 01' the P. trinum-t y pc com ply full y with the spec ies description .

I 'D. Penicillium corylophilum Dierckx : fig . 127, colour plat e 127, and
SE M pl ate 22, (top: left) .

i.w.J.lJ.wJ

00

10 pm

00 0
0 8 0 0
00

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100 prn

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Fig. 127. Camera lucida

drawings of Penicilli um
corylophilum Dierckx , CBS
312.48. A, Detailed drawings
of the penicilli. B, Mature
conidia . C, Habit sketches.

Penicil/ium corylophilum Dierckx - Ann . Soc oSci. Bruxelles. 25: 86. 1901 . See also
Biourge's Monographie, La Cellule. 33: 266-267. 1923; also Ch. Thom , 'The
Penicillia', pp . 254-255. 1930; also Raper and Thom, 'A Manual of the Penicillia' ,
pp. 341-345. 1949.

Synanymies
Penicil/ium cloro-Ieucon Biourge - Monographie , La Cellule. 33: 270-271 . 1923.
Penicil/ium coeruleo-viride Smith - Trans. Brit. Mycol. Soco 48: 274. 1965. See
also M. M. Kulik, ' A Compilation of Descriptions of New Penicil/ium Species'.
Ag ric. Handbook No. 351. Ag ric. Res. Ser. U.S. Dep. Agric . pp . 29-30. 1968.

Penicillium corylophiloides Abe - J . Gen. Appl. Microbiol. Tokyo. 2: 89-90. 1956.

See also M. M. Kulik , 'A Compilation of Descriptions of New Penicillium Species'.
Agric. Handbook No. 351 . Agric. Res. Ser. U.S. Dep. Agric. pp . 31-32. 1968.
Penicillium obscurum Biourge - Monographie, La Cellule. 33: 267-269. 1923.
Penicillium sumatrense von Szilvinyi - Arch. fr Hydrobiologie. Suppl. Bd. XIV;
Tropische Binnengewasser . Bd. VI, pp. 551-552. 1936.

Co lonies up on C A (colour plate 127 , A and a) att a ining a di amet er of 40 mm

in 14 days at room temperature (20- 25C), con sisting of a close-textured basal
feIt , typi call y velve ty in appeara nce, with flat margin surrounded by a wide
frin ge of sub me rge d hyphae exte nding ab out 5 mm beyond the aerial gro wth ;
colony su rface stro ngly a nd deeply radiately furrowed in submarginal zones
and irregul arly fold ed and raised in th e centre, cons picu ou sly zonate owing to
the un equal degree of ripe ness of conidial zones ; heavily sporing throughout in
blue-green shades (Sguy, PI. XXXI , 462) with th e rip ening of conidia;
exudate lim ited in amo u nt, as small , colou rless dropl et s; odou r lacking;
reddish a m be r-colou red pigm ent diffusin g into th e surrounding agar ; colony
reverse (colour plat e 127, a) in reddish b rown shad es; peni cilli var iable in
pattern and dimen sion s (fig. 127, A) , typi cally biverticillate and asy m me tric
but with simple struc tu res (appeari n g mon overticillate) some times
pred ominant ; with conid iopho res smooth walled , arising mostly from th e sub stratu m , particul arl y in m argin , but elsewhe re as bran ch es fro m th e network of
a eri al hyphae overlaying th e basal felt , ranging fro m 40 to 200 tm in len gth by
2.2- 3.0 titt: in diameter, u su all y not ramified ; penicilli typi cally cons isting of
clusters of 2- 3 metulae per ve rt icil, variable in len gth , ran gin g from 15 to 30
tm by 2.0 -3. 0 tm , with api ces slightly enla rge d in a vesicle up to 4.5 tm in
diamet er; phialides parallel , in clu sters of 3-8 elem ents in th e ve rticil, abo ut
8-12 tm by 3. 0 -3.8 tm abru ptly tap ering to a very sho rt narrowed neck ;
con idia usually subglobose to elliptical, com mo nly 3.0 -4 .0 tm by 2. 8 - 3 .5 tm
in diameter, smooth wa lled or nearl y so (fig. 127 , B, a nd SEM plate 22 , top :
left) ; conidial chains loosely adherent or tan gled (fig. 127 , C) .
Colonies upon CYA (co lou r pIate 127 , B a nd b) attaining a di amet er of 45
mm in 14 days at room temperature (20-25C), simila r in textu re as aboye but
with conid iu m production beginning lat e , appeari ng first in th e mor e cro wde d
margin al areas of adjacent colonies but soo n developing in zones ove r the
whole colony surface; m argin entire; exudate abu nda n tly produced , as yellowgree n coloure d drops; odour fa int, indefinite; a mber yellow -coloure d pi gm ent
diffusin g into th e surrou nd ing agar; colony reverse (colour plat e 127, b) in
reddi sh orange shades; penicilli esse ntially as describ ed aboye.
C olonies up on M EA (colour plat e 127, C a nd e) spread ing broadl y ,
attain ing a diamet er of 50 mm in 14 days at room temperature (20 -25C),
flat , not furrowed , cons isting of a den se, stri ctly velvety feIt , with a very thi n

. J . J. J

murgin surroundcd by a wide fringc 01' submcrged hyphae extend ing about 5
more bcyond thc ac r ial growth; hcav ily sp or in g thro ug ho u t in deep
blu c-grcen shades (Sgu y , PI. X X X , 447); exudate and odour lackin g ; no
colourcd pigment diffu sin g int o th e surrou nding agar; colony reverse (colour
pla tc 127, r ) parti ally colou rless with locali zed zo nes, pa rticu la rly in the centre
dcvclopi ng very dark green, almos t black colou rs . Penicilli esse ntially as
dcscribed above.
T he spec ies descript ion is centered on th e ty pe strain ATCC 9, 784 = C BS
:112.48 = CM I 39, 754 = N R R L 802 = QM 7,5 10, isolat ed by L. P . D ierckx in
1901 , on strain CBS 281.36 = N RRL 779 = IJ F M 3754, isolat ed by A . von
Slviny i (as Penicillium sumatrense v . Szilvinyi) fro m tropical continenta l wa ters
i l\ 1936, and m any other strains isolated fro m soi1 in the Province 01' Navarra,
Spain , by DI'. A. T . M a rtnez .
111111 0 1'

128 . Penicillium steckii Zaleski: fig. 128, colour plat e 128, and SE M plate

22, (top : rig ht).

O O
O

80

00
00 O

O O

O O000 0O'
8

00

O00O00
00

10 pm

~~
L.-........J
jrrn

100

eR .
Fig. 128. Camera lucida
drawings of Penicillium
steckii Zaleski, CSS 260.55.
A, Detailed draw ings of the
penicilli . S, Mature conid ia.
C, Habit sketches.

.'''\1

Penicillium steckii Zalaski - Bul!. Acad. Polonaisa Sci. Math. Nat. Sr. B, pp.
469-471.1927. See also Ch. Thom, 'The Penicillia', pp. 255-256.1930; also Rapar
and Thom, ' A Manual of the Penicillia', pp. 350-352. 1949.

Colon ies upon CA (eolou r pla te 128 , A and a) growing rather restri etedl y,
attain ing a d iameter of about 30 m m in 14 days a t roo m tem perature
(20- 25C), eons isting of a tough , close- tex tured basal felt , appearing velvety,
za nate, ra ised a nd radiately furrowed , with white eve n marg in su rro u nded by
a narro w frin ge of su bmerged hyp hae extend ing abo u t 1 mm beyond th e aerial
growth ; medium to heavy sporing in zones, depend ing of the degree of ri peness
of eonidial zo nes; conidial areas in blue-green shades (Sguy, Pi. X XX , 443)
with th e ripening of eon idia; ex udate lim ite d to abu ndant depend ing upon the
strain, as pale yellow -eoloured droplets; odour at first laeking or fain t, with age
beeoming somewhat mo uldy; very pa le amber-eo loured pigme nt d iffus ing in to
the surrou nding agar; eolony reverse (eolour plat e 128 , a) in pink ish orange
shades; penieilli typieally bive rtieillate asymmetrieal (fig. 128, A), with
eonid iophores arising from the substratum or th e basal felt, smooth wa lled ,
variable in length , but eommonly eomparatively short, seldom exceeding 200
iuu, generally less, by 2.5-3 .5 iuu , u sually not ramifi ed ; penieilli eonsisting of
a ter m inal vertieil of 3-5 m etul ae bearing eompaet whorls of phialides
produeing well-defined d ive rgent eolumns (fig. 128 , C) of eon id ia u p to 150 J-m
in len gth ; m etulae ra ngi ng from 12 to 18 tui: by 3 - 4 J-m; phialides in erowded
clusters, parallel , about 8 - 12 J-m by 3.O- 4 .5 usn, bottle shaped, ta pe ring to a
conspieuous, short , narrowed neek; eonid ia comparatively small, globose to
subglobose, abo ut 2.0-2 .5 J-m in d iameter, with wa lls smooth or del iea tely
ro ughened (fig. 128 , and SEM plate 22, top : right) .
Colonies u pon CYA (eolour plate 128 , B and b) attaining a d ia mete r of 35
mm in 14 days at room tem pera ture (20 -25 C), similar in text ure and eolony
appearanee to the preeedi ng, but more deeply radiately furrowed a nd less
ra ised; heavily spori ng in zo nes in bl ue -green (Sguy , Pi. XXX , 443) to graygreen shades (Sguy, Pi. X X X V , 519), depen di ng on the degree of ripeness of
eonidia; exudate very abundan dy prod uced, as la rge yellow -eoloured drops;
odour laekin g; pale amber-eoloured pigme nt diffusi ng in to the surroundi ng
agar; colony reverse (eolour plate 128, b) in yellowish orange shades ; peni eilli
essentially as deseri bed aboye .
Colonies u pon M EA (co lour piate 128, C and e) attaini ng a diameter of 30
mm in 14 days at room temperature (20- 25 C) , eonsisting of a looser textu red
basal felt, fla t , with seetors of aerial, floeeose , vegetat ive, whi te h yph ae overlaying th e basal felt, with very thin margin surrou nded by a wide fri nge of subm er ged h yphae extendi ng abo ut 5 m m beyond the aerial growth; heavily
spo ring throughout in deep a rtemisia green shades (Sguy , Pi. XXI, 308) ;
exudate and odour laekin g ; no eoloured pigment d iffus ing into the su rrou n-

:1;17

d ill!{ agur; co lo ny reve rso (co lour plat e 1211 , e) in paj e yellow sha des, bccoming'
oru ngc-ycllow in th e ce n tre ; pcni cilli casc nt iully as dcscribed a hoye.
T he spcc ies description is ccntc red llp on stra ins CI\S 260.55 = CM140,583 =
\'['cc 10,499 = NRR L 2, 140 = Q M 6,41 :~ = I./FM 7211 , isolat ed by Prof, Wm .
11. W cston fro m a sa m ple 01' fabric ex pos cd on Barro Colorado Island ,
l'uu ama , 1948 , a nd from othe r stra ins isolatcd from so il in th e Province 01'
Nav arra by Dr. A.T. Martnez , 1975. (Colou r pl ate 236 presents a representa tiv e strain 01' P. steckii isol at ed fro m sea sa lt. )

129. Penicillium matriti* Smith: fig. 129 , colour pl at e 129 , and SEM plat e
22 , (ce ntre : left),

Lw.L!.wJJ

10 urn

o
0) 0
O O O
0
0 00 0 0 0
00 00

." o/ o;""~';f1;,?
00

,,~

100,JJm

;;::k'

"\

('l. .

~'"l~'" ,

'",\,

Fig. 129. Camera lucida

drawings of Penicillium
matriti Smith, CBS 347.61.
A, Detailed drawings of the
penicilli. S , Mature conidia.
e, Habit sketches.

Penicillium matriti Smith - Trans. Brit. Myeol. Soe. 44: 44-45. 1961. See also M.M.
Kulik, 'A Compilaton of Deseriptions of New Penicillium Species' . Agrie.
Handbook No. 351. Agre. Res. Ser. U.S. Dep. Agrie. p. 52. 1968.
N ote of the a u thor: The bi no mial P. madriti has been fou nd to be orthogra phically incorrect o
T he correc t design ati on , namely P. matriti , is ther efore prop osed.

Smith's original diagnosis follows, since this adequately presents the species in
question when first isolated:
'On Czapek agar grows fairly rapidly, pale blue-green , turning gray-green, with
white edge about 2 mm broad during the growing period, velvety but tending to
become somewhat heaped in the centre; drops numerous, small, yellowish; reverse
peach coloured, indistinctly zonate ; on malt agar grows very rapidly, grayish bluegreen, thick velvety, with numerous yellow drops and reverse very reddish brown;
penicilli with metulae each bearing a compact cluster of phialides, branching below
the metulae not observed ; conidiophores arising from the substrate, long, smooth,
2-3 m in diameter; metulae slightly swollen at the apex, 10-11 m by 2.5-3.5 m;
phialides tapering rather gradually, but not acuminate as in the Biverticillata, 7-8
m by 1.5-1.8 m; conid ia globose, finely roughened, 2-3 m, mostly 2-2.5 m in
diameter, conidial chains forming a single fairly compact columnar rnass, often bent
or slightly twisted.
This species appears to form a transition between the P. citrinum and the P.
chrysogenum series. In the lack of any branching below the metulae it is nearest to
P. steckii Zaleski, but the latter shows slower growth, thinner and more bluish
colonies, and spore chains in several distinct columns, one to each metula. P.
matritiwas also grown in parallel cultures along with P. cyaneo-fu/vum Dierckx, the
member of the P. chrysogenum series which it most closelv resembles . P. cyaneofu/vum has thicker, almost floccose, colonies, penicilli which frequently show
branching below the metulae, and much larger spores than P. matriti.
The type strain, maintained as Cato No. BB 389, was isolated in 1954, by G.
Agosti in this Department (London School of Hygiene and Tropical Medicine), from
garden soil from Madrid. A second strain BB 389 A, was isolated from soil from a
different garden in Madrid. On Czapek agar the latter forms slightly thicker colonies
than BB 389, and rather more brownish colour in reverse; on malt agar the two are
indistinguishable. '

Our no tes follow :

C oloni es upon CA (colour pIat e 129, A and a) spreading broadl y attaining a
diameter of 55 mm in 14 days at room tem perature (20 - 25C) , con sisting of a
close-t extured basal felt , appearing velvety, stro n gly and irregul arl y wrinkled
throughout , with a very thin flat margin , su rrou nde d by a wide frin ge of submerged hyphae extendi ng ab out 5 mm beyond th e ae rial gro wth; medium
sporing in bluish gray sh ade s (Sgu y, PI. XXX, 449) with the ripening of
conidia, elsewhere in sandy shades; od ou r faint , aro matic; exudate limited in
amount, as crys ta l-clear, colourles s droplet s; pale a mbe r-coloure d pigm ent
diffu sin g into the su rround ing agar ; colony revers e (colour plat e 129 , a) in
reddish brown shades ; penicilli (fig. 129, A) esse n tially as described aboye,
cons isting of terminal whorls of 2- 3 metulae per verticil , ab out 10-1 6 /lm by
2.5 - 4.5 tui, with apices enlarged in a vesicle up to 4.5 tut: in diameter;

phiu lide bot tle sha ped , tapcring lo a sho rt narrowcd neck , lIIostly B-IO /-UIl by
:1.0 - :i .5 JlIII; co n id ia globos e , about 2. :> - :i. O JllII in di a mctcr, a p pcr u -ing
IOlloo lh wall ed 01' ncarl y so wh cn obsc rvcd lind el' (he light microscop e ( fig . 129,
B) 0 1' dclicately rou gh en ed whe n obse rved undcr th c SE M (pIa te 22, ce ntre :
lef't ); co nid ial chains in well-defined di vcr gcnt colu mns (fig . 129, C).
C olonics upon C YA (eolour plato 129 , B a nd b) attain ing a diameter of 50
111111 in 14 da ys a t room te m pe ra tu re (20 -25 C ), eons isting of a sim ila rly
ic xtured ba sal felt bu t mo re regula rly wrinkled in a radial and conce nt r ical
pnt tcrn , with a narro w flat margin su rrou nde d by a frin ge of su bme rge d
hyph ae extend ing abou t 1 mm beyo nd th e ae rial growth; abundantIy sporing
in conspicuous zo nes in m ou se gray (Sgu y , PI. XXIX , 430) to pale pinki sh
': I'a y shades, dep ending on th e degree of rip eness of eon id ia; odour stro ng,
sllggesting am mon ia; exu date not obse rve d; arnbe r-colou re d pigment diffu sin g
i nro the sur rou nd ing agar ; penieilli esse ntially as deseribed aboye .
Colonies upon M EA (colour plate 129, C and c) gro wing rather restrietedl y ,
a llaining a di ameter of 45 mm in 14 days at room temperature , flat , not
lurrowed , eon spieuo usly zonate, whi te th in m argin surrounde d by a frin ge of
subrnerged hyphae extend ing about 2-3 mm beyond th e ae rial gro wth; eolony
surface typi call y velvety , ove rlaid in sectors by a thi n closely woven network of
white ae rial h yphae obse u ring th e ot he rw ise heavy spor in g are as; heavily
spo ring throughout in blue- gr een shades (Sguy , PI. XXXI , 462); exudate an d
od our laeking ; no eoloure d pigment diffu sing into th e surrounding agar;
ro lony reverse (colou r plate 129 , e) developing sien na reddish brown colou rs in
loealized seetors eorres po nd ing to th e heavily spo ring are as, and pale yellow
sha des in seeto rs eorres po ndi ng to th e ove rgrowth of vege ta tive floceose
hyphac ; penicilli essentially as described aboye .
The spee ies deseription is een tere d up on th e typ e strai n C BS 347.6 1 = CMI
B6,563 = ATC C 18,223 = Q M 7,959 = IJ FM 7128 , isolat ed from ga rde n soil in
M adrid , 1954 , by G. Agosti .
M embers of th e Penicillium citrinum series occu r upon a wid e va riety of substrata su bject to soil conta m ination including cott on a nd othe r fabrics, dairy
and food product s, tobacco, leather goo ds, and vegetable m at eri als undergoing
slow decomposition. They are com mo nly isolat ed from soil. They have been
cncou ntered more frequently th an any othe r Penicillia upon deteri or ating
m ilita ry equ ipm ent in tropical and su b- tro pical cou ntries. They m ay be
isolat ed from len s mountings in photogr aphic ca me ra s, with th e mycelia
exte nd ing ou t upon the glass surfaee and etching it sufficientIy to render it
praetically us eless .

Penicillium

dlly ,\'o,l.tt:7lUm

seri es

O U T STANDIN G FEATU R ES

Colon ies typi cally velvety, more 0 1' lcss clo se textured , with co nid iopho res
arising frorn the subs tra tu m in a dense stand ; in sorn e strain s more 01' less
floccose , with conid iopho res borne as bran ches from ae rial hyphae ; com m only
radiately furrowed .
Exudat e com mo nly abu ndantly pradu ced as bri ght yellow drops.
C olony reverse usually in th e same b right-yellow tints, becoming brown in
age, with the su rra u nd ing agar sim ila rly colou re d .
Peni cilli typi cally asy m metrical, with con id iophores smo oth walled,
irregul arly ramified in larger stru ct ures, a l' u su ally cons isting of a terminal
wh orl of m etulae in smaller pen icilli; with con id ial chains formin g well-de fined
compact colu m ns eac h arising from individual ve rticils of phialides.
Conidia sm oot h- walled a l' nearl y so, va ry ing from globose, su bglobo se to
definitely elliptical, dep ending on th e strains, and rangin g in size from abou t
2.5 to 4.5 flm in di ameter.
The series is a t present represented by a single species, Penicillium chrysogenum
Thom , since th e three othe r spec ies recognized by R ap er a nd Thom (1949 ),
namely: P. meleagrinum Biourge , P. cyaneo-fulvum Biourge , and P. notatum
W estling have recently been reduced to syno ny m ies of th e former species
(Sa mso n et al. 1977 ).
Penicillium chrysogenum Thom is espec ially commo n in natu re, occ u rrin g fairly
regul arly in soil and up on a wide variety of org an ic su bstra ta. It has received
particul ar atte n tion during th e past few decades after th e di scovery of th e a ntibiotic penicillin by P rafessor Alexander Flemin g in 1928, wh ose interest was
revived in 1940 thraugh th e demon strati on of its pot ential th erapeutic value by
Professor s E . C ha in, H.W . Florey , and others at Oxford University , UK.
The species is quite va riable in pigmentati on , exudate production , colony
texture , rat e of grawth , etc . , dep ending upon th e strain, yet all show th e
gene ra l charac teristics indicat ed aboye and ca n be readily recognized.
In th e m orphology of th e peni cilli , th e va ria tion m ay range fram stra ins in
which th e penicillu s cons ists of a conidiophore bearing a serie s of branch es ,
with eac h praducin g a m on overticillat e structu re, to a com plex a nd typi cally
asymmetrical appa ra tus. The conid ia may vary from globose, su bg lobose to
defin itely elliptical, and fro m quite small , with di ameters m ostly less th an 3 .5
tui, to strains with conid ia up to 4.5 flm in th e long ax is.

130 . Penicillium chrysogenum Thom : fig. 130 , colo ur plat e 130 , and SEM
plat e 22 , (centre : ri ght) .
Penicillium chrysogenum Thom - Bull. Bur. Anim . Ind. U.S. Dep. Agric. 118:

00 o

000
O

000
B
O 00Wul=J
10jJm

Fig. 130. Camera lucida

drawings of Penicillium
chrysogenum Thom, CBS
306.48. A, Detailed drawings
of the penicilli . B, Mature
conidia . C, Habit sketches .

58-60.1910. See also Westling, Arkiv. Bot. 11 : 54,107-108.1911. Also Biourge,

La Cellule. 33: 170-172. 1923. Also Thom, 'The Penieillia', pp. 261-262.1930; also
Raper and Thom, 'A Manual of the Penieillia', pp. 359-373. 1949.
Synonymies
Penicil/ium aromaticum Sopp forma microsporum Romankova - Sci. Papers
Lenin Univ. Biol. Ser. 40,191: 102. 1955. See also M.M. Kulik, ' A Compilation of
Descriptions of New Penici//ium Species' . Agric . Handbook No. 351. Agric . Res.
Ser. U.S . Dep. Agrie . pp. 16-17. 1968.
Penici//ium baculatum Westling - Svensk Botanik Tidskrift. 4: 139-145. 1910; see
also Arkiv Bot. 11 : 53, 79-83. 1911. Also Biourge's Monographie, La Cellule. 33:
186-188.1933. Also Ch. Thom, 'The Penicillia', pp. 268-269.1930; also Raperand
Thom, 'A Manual of the Penicillia', p 363. 1949.
Penici//ium brunneo-rubrum Dierckx - Ann . Soco Sei. Bruxelles. 25: 88. 1901.
See also Biourge's Monographie, La Cellule. 33: 176-179. 1923; also Ch. Thom,
'The Penicillia', pp. 267-268. 1930; also Raper and Thom, 'A Manual of the
Penicillia', pp. 372-373. 1949.

Penicltlium cemerunense Heim, Nouvel et Saccas - Bull . Acad. Roy. Belgiqua

(CI. sei .i. Sr. 58, 35: 42-49. 1949. See also M.M . Kulik, 'A Compilation of
Descriptions of New Penicillium Species'. Agric. Handbook. No. 351 . Agric . Res .
Ser. U.S. Dep. Agric. p. 23. 1968.

Penicillium citreo-roseum Dierckx - Ann. Soc o Sci. Bruxelles. 25: 86. 1901; sea
also Biourge's Monographie, La Cellule. 33: 182-184. 1923; also Ch. Thom, 'The
Penicillia' , pp. 265-266. 1930; also Raper and Thorn, 'A Manual of the Penicillia',
p . 373. 1949.
Penicillium chlorophaeum Biourge - Monographie, La Cellule. 33: 271 -273.1923.
See also Ch. Thom, 'The Penicillia', pp . 262-263.1930; also Raper and Thom, 'A
Manual of the Penicillia', pp. 363-364. 1949.
Penicillium cyaneo-fulvum Biourge - Monographie, La Cellule. 33: 174-176. 1923;
See also Ch. Thom, 'The Penicillia', p. 267. 1930; also Raper and Thom, 'A
Manual of the Penicillia', pp. 371-372. 1949.
Penicillium fluorescens Laxa nomen nudum (?) - It was cited but not described by
O. Laxa, Zentbl. Bakt. Parasitenk. Inf. Hyg. 11 Abt. 86: 164-165. 1932. See also
Raper and Thom , 'A Manual of the Peniciltia', p. 364. 1949.
Penicillium griseo-roseum Dierckx - Ann. Soco Sci. Bruxelles. 25: 89. 1901. See
Biourge's Monographie, La Cellule. 33: 168-170. 1923; also Ch. Thom, 'The
Penicillia', pp . 263-264. 1930.
Penicillium meleagrinum Biourge - Monographie, La Cellule. 33: 147-149. 1923.
See also Ch. Thom, 'The Penicillia', pp. 266-267. 1930; also Raper and Thom, 'A
Manual of the Penicillia', pp. 364-367. 1949.
Penicillium notatum Westling - Arkiv. Bot. 11: 55, 95-97. 1911. See also
Biourge's Monographie, La Cellule. 33: 179-181. 1923. Also Ch. Thom, 'The
Penicillia', pp . 264-265. 1930; also Raper and Thom, 'A Manual of the Penicillia' ,
pp. 367-371. 1949.
Penicillium roseo-citreum Biourge - Monographie, La Cellule. 33: 184-186. 1923.
See also Ch. Thom, 'The Penicillia', p. 323. 1930; also Raper and Thom, 'A
Manual of the Penicillia', p. 364. 1949.
Penicillium rubens Biourge - Monographie, La Cellule. 33: 265. 1923. See also
Ch. Thom, 'The Penicillia', pp. 249-250. 1930; also Raper and Thom, 'A Manual
of the Penicillia', p. 364. 1949.

C olonies upon C A (colour plat e 130 , A a nd a) growing rapidly , attain ing a

d ia mete r of about 55 mm in 14 day s at room temper ature (20 - 25C) ,
cons isting of a fairl y close-t extured basal felt, comparatively thin , commonly
1- 2 mm de ep , stro ngly furrowed in a radial pattern but with central area usu ally
irregul arly buckled and wrinkled , in sorne strains closely velvety, in othe rs
deepl y velve ty and ra the r loose textured , usu all y azonate, with gr owing
m argin flat , 1- 2 m m wide, surro unde d by a fringe of submer ged hyphae
extend ing abo ut 1- 2 mm beyond th e ae rial gro wth; heavily sporing

th rou ghour in most stru ins, in o thcrs o tcn sho wing sorne tcndcn cy lo rc mn in
srerilc in centra l arcas with veget ati vo hyph ae ycllowish to crc a m colou rcd,
Iu:collling dcc p bl ue grcc n (S guy , PI. X X X , 447 ) with the ripc ning 01' con id ia;
cxuda te ab unda nt ly produ ced in most stra ins, as golde n yellow co ns picuo us
dro ps ; odou r lacking 0 1' ind efinitc ; bright yellow-coloure d pigme nt diffusin g
l l lo (he surro und ing agar; colony reverso (colour pla te 130 , a) in bri ght yellow IIl'ange shades; conid iophorcs arisin g mostly fro m th e su bstratum in a den se
xtund , smoo th wall ed , variable in leng th, us ually rangin g from 150 to 400 J-t m
111' mor e in length by 3.0-3 .5 J-tm in di am eter, occasionally bo rne fro m aerial
hyph ae a nd shorte r, abo ut 50- 75 J-tm in leng th, with ap ices somewhat inflat ed ;
pcni cilli asy mmetric (fig . 130 , A) , commonly branched , with branch es a nd
nurin axes each usu all y bearing 2 -4 metulae to form a fairly large peni cillu s,
k ss com mo n ly cons isting 01' a simple terminal vertic il 01' m etulae only, bea rin g
rl us ters 01' phialides producing con id ial chains up to 50 - 75 J-tm in length ,
uu igled 0 1' adhe re nt int o fai rly well-defin ed colu m ns; branches va riable,
ro m mo nly 15- 25 J-t m by 3.0 -3 .5 J-tm ; me tulae u su all y 10-1 3 J-tm by 3 .0 - 4. 0
tIJl, occasionally lon ger; phialides bottl e shaped, in fairly com pact cluste rs 0 1'
'1- 6 ele me n ts in the vertici l, m ostly 8 - 10 tut: by 3.0-4. 0 J-tm but va riable in
d ilferent strains, abru ptly ta pered to a sho rt narrowed nec k; conidia ellipt ical,
su bglobose 01' globose, dep ending u po n th e stra in, mostly 3.0 -4.0 J-tm by
L.B- 3 .5 utt , occasionally larger, smooth wall ed 0 1' nearly so wh en observed
lindel' th e light microscope (fig . 130 , B) but del ica tely ro ughe ned wh en
observed under th e SEM (piat e 22, ce ntre: right) ; coni dial chains in more 0 1'
lcss defin ed colu m ns.
Co lonies u po n C YA (colour plate 130, B and b) spread ing as upon C A,
altaining a d iame ter 0 1' abo u t 50 mm in 14 days at roo m temper ature
(20- 25C), sim ilar in tex ture to the precedin g but heavier sporing throu gh out
in mos t stra ins a nd conspicuo us ly zonate, in th e same deep blue-gr een shades
with the ri pening 0 1' coni d ia (Sguy, PI. X XX, 447); with thin margin
surrou nde d by a fringe 01' su bmerge d hyphae exte nd ing abou t 3 mm beyond
the ae rial growth; strong ly radiat ely furrowe d and pulling the agar away and
crac king th e colony su rface in th e centre as deep crevices along th e ra d ial
Iurrows; ex udate abu ndantly produced as large , b right golde n yellow drops;
odo ur lacking ; bright golde n yellow-coloured pigment di ffus ing abu ndantly
int o th e surrou ndi ng agar; colony reverse (colou r plate 130 , b) develop in g deep
ora nge -yellow shades; peni cilli esse ntia lly as descri bed a boye bu t cons istently
heavier spo ring .
Colonies upon M EA (colour piate 130 , C and c) attaining a di ameter 01'
55- 60 m m in 14 days at ro om temperatu re (20-25 C), cons isting 0 1' a looser
tex tured basal felt , flat , never fu rrowed, strictly velvety, zanat e, with thin
m argin surrounded by a fringe 0 1' su bme rge d hyphae exte nd ing abo ut 4 mm
beyond th e ae riaJ gro wth; heaviJy spo ring throu ghout in gr ay-gr een shades

(S gllY, PI. XXIX , 429); cx uda tc a nd odo u r luckin g; no COl0 11I'l: d pi gm en!
dillu sing in to th c su rro u nd ing aga r; co lo ny rev erso (co lour plat o 130, e) in dul l
yellowish, tu vinaceous or brownish drab shadc s: pcni cilli cssc ut iully as
described ab oye but tending to dc vclop better defin ed colu m ns 01' co nid ia (Ilg.
130, C).
The spec ies description is cen tered up on th e typ e stra in C BS = 306 .48 = C M I
24, 314 = AT C C 10,10 6 = Q M 7,500 = NR R L 807 = IJFM 7063, isolated by C h.
Thom , and up on com parative exa m ination al' many add itiona l stra ins
representing th e sa me basi c cu ltu ra l a nd morphologic al charac te ristics, a rno ng
th em , strain C BS 355.48 = C M ! 39 ,759 = ATCC 10,108 = NRRL 821 , isolat cd by
R. W estling frorn bran ch es of Hyssopus in Norway , 1910 , is the formerly typ c
stra in of Penicillium notatum W estling, actually syno ny mo us with P. chrysogenum
Thom . (Co lou r plat es 237 and 238 present two representati ve stra ins of P.
chrysogenum, isolat ed frorn th e a ir by th e autho r. )

131. Penicillium chrysogenum Thom muto ful vescens Takashima , Arima el

Abe ex Ramrez : fig . 131 , co lo ur pl ate 131 , and SEM plate 22 ,
(bottom : left) .

lJ.J.J..WwJ

10jJm

O O

000
00 0
0 800 0
O

00

00 O
000

1~ 11

O
0
Fig . 131. Camera lucida
drawi ngs of Penicillium
chrysogenum Th om mut o
fulves cens Takashima, Ar ima
et A be ex Ramrez, CBS
319.59. A , Detailed draw ings
of the penicilli. B, Mature
conid ia. e, Habit sketc hes.

nicillium chrysogenum Thom muto fu/vescens Takashima, Arima et Abe - J.

Gen. Appl. Microbiol., Tokyo. 2: 92.1956. See also M.M . Kulik, 'A Compilation of
Descriptions of New Peniclllium Species'. Agric. Handbook No. 351. Agric. Res.
Ser. U.S. Dep. Agre. pp. 25-26. 1968; also G. Smith. Trans. Brit. Mycol. Soco
46: 332. 1963.

T his strain wa s obta ine d by T ak a shima a nd Arima a s a mutant from a

norm al strain of Penicillium chrysogenum Thom FAT 5 14 by ultra-violet light
irrad iation . It was described by S. Ab e in 1956 as a new va riety , prop osin g th e
llam e of Penicillium chrysogenum Thom mut. Jul vescens, but without providing a
I.atin di agn osis. To va lida te th e variet y a Latin diagn osis of it is give n below ;
I he names of th e original autho rs are m a int ained under R ecommendation 46 C
01' the Internation al C od e of Bot anical Nomenclature (Sta fleu (Ed .), 1972).
Latin di agn osis of Penicillium chrysogenum Thom m ut. Julvescens T ak a shima ,
Ar ima el Ab e ex R amrez:
Mutatio a Penicillium chrysogenum Thom differt propter colonias cynnamomeas
colorant ur, nunquam coeruleo-vrdem.

Colonies of th e m uta n t upon C A (colou r pIat e 131 , A a nd a) differ fro m th e

ryp e spec ies in th e entire abse nce of gree n colou r , a nd usu ally sho wing various

shades of cin namo n in heavily sporing conidial areas; attaining a di ameter of

40 mm in 14 da ys at room temperature (20 -25 C); exudate abu ndantly
produced in b ri ght yellow shades; od our lacking; colony reverse (colour pIate
131 , a) in reddish brown sha des ; reddish brown coloured pi gm ent diffusin g
int o th e surro unding agar ; penicilli (lig. 131 , A a nd C), essentially as in th e
type spec ies .
Colonies upon C YA (colour plat e 131 , B a nd b) a tta ini ng a di amet er of 55
mm in 14 days at roo m temperature (20-25 C) , similarl y colou re d as aboye
but in pal er cin na mon shade s; exuda te limited in amo unt; odo ur lacking ;
golde n yellow-coloure d pigm ent diffu sin g into the su rrounding agar; colony
reverse (colour pIate 131 , b) developing ora nge shades; penicilli esse nt ially as
aboye .
Colonies upon M EA (colou r plate 131 , C and c) attaining a di a m eter of60
mm in 14 days at room temperature (20 -25 C ) , th in, strictly velvety, flat ;
heavil y sporing thro ugho ut in pale cinna mo n shades; no exudate produced ;
odo ur lacking ; no coloured pigme nt d iffusin g into th e su rro u nding agar ;
reverse (colou r plat e 131 , c) developing pale ye llow ish cin namo n shades ;
peni cilli as aboye .
T he mutant description is ce ntered up on strain CBS = 3 19.59 = CMI68, 23 1
= ATCC 18, 226 = IF 6,223 = QM 7,296 = IJFM 720 3, ob ta ine d by U .V . light
irrad iation of stra in FA T 5 14 of Penicillium chrysogenum isolat ed from J a pa nese
soil by S . Ab e , 1956 .

Strains of Penicillium chrysogenum T ho m OCCUI' rcgula rly in soil a nd are

com rnonly isolatcd Irom l wid c variety 01' or ganic substrata, including: chces c
a nd othe r dairy products , bread and pastries , fruit s, vegetables, rneat and rnca t
products, improperly ca nned foods , a nd decayin g vegetation 0 1' all kind s
(Rap er, Alexander, and Coghill, 1944) . The species is unquestionably world
wid e in di stribution.
The principal interest in thi s spec ies is derived fro m th e ca pac ity 0 1' thi s
mould to produce penicillin .

Penicillium oxalicum se r ies

O U T ST A NDING FEATU RES

C olonies spreading broadl y , velutinou s, flat 01' irregularl y 0 1' radi at ely
furrowed; heavily sporing throu ghout in dull to deep blue- green shad es ;
velve ty ; with conid ia 01'ten formin g deep crusts whi ch break off wh en th e
culture vessel is trapped .
Co nid iophores typi cally arising from th e subs tra tu m in a den se a nd eve n
sta nd, va r iable in length but u su ally not excee ding 200 Lm ; smo oth wall ed.
Pcni cilli typi call y biv erticillat e asym me trical, cons isting 0 1' 2 0 1' more
metulae , 0 1' irregularl y branch ed , with branch es and metulae some time s
arising a t th e sa me level , 01'ten closely appres sed.
Phi alides closely parallel , com monly producing chains 0 1' conidia adhering in
loose to com pact colu m n s up to 500 Lm 0 1' m or e in len gth .
Con idia usu ally elliptical, sm ooth-walled 0 1' conspicuo usly spinulose.
K ey to spe cie s
Page
1. Penicilli typically rebranched below the level of metulae, with the main
axes and branches terminating in verticils of metulae. Penicilli commonly
long, with elements loosely arranged and often divergent. Conidio
phores smooth walled; margin not appearing arachnoid. Colonies not
producing yellow pigment in exudate or colony reverse; conidia
commonly 5 ,tm or more in long axis; elliptical or subglobose , fairly
uniform in size; common in soil.
P. oxalicum series . .. . 366
A. Conidia smooth walled.
1. Conidia elliptical; colonies plane or nearly so, with conidia often
forming deep crusts; con idial chains appearing 'silky' when viewed
under lower power.
P. oxalicum Currie et Thom . ... 367
2. Conidia subglobose; colonies radiately furrowed, with conidia not
forming crusts; colony reverse upon CYA developing deep cherry

red colours, also diffusing into the surrounding ager.

P. etrementosum Thom . ... 369
B. Conidia conspicuously roughened; colonv reversa developing deep

orenge-red shades.

P. fennelliae Stolk . . . . 371

Th c series as recogni zed by R aper and 'I'horn has been broaden ed eno ugh as
include in it one species: Penicillium fen nelliae Stolk, add itional to th e two
wcll-marked , vclvety spec ies with elliptica l, smooth-wa lled conidia already
incluclcd in it (Rape r a nd T ho m 1949). It is a rtificial in characte r and was
introduced primarily as a m atter of conve n ience .
Alth ou gh a cla se relat ion ship is not presumed between th ese three species,
thcy do possess su fficient cha racters in com mo n to locat e th em here m or e
n -ad ily tha n any whe re else.
O ne of th e spec ies , Penicillium oxalicum Currie el Thom is world wid e in
clist ribution a nd one of th e most ubiquitou s of all th e Penicillia . It has long
I leen known to represent a normal com po ne nt of th e soil m ycoflora .
The seco nd spec ies , Penicillium atramentosum T ho m is prob abl y not close ly
rclated to P. oxalicum , and unlike it , P. atramentosum appears to be
ro rnpa ra tively rare .
lo

1:12. Penicillium oxalicum Currie el Thom : fig . 132 , co lo ur pl a te 132 ,

a nd SEM pl ate 22 , (botto m: ri ght).

Penicillium oxa/icum Currie etThom - J. Biol. Chem. 22: 289.1915; also Ch. Thom,
'The Penicillia', pp. 247-248. 1930; also Raper and Thom, 'A Manual of the
Penicillia', pp. 378-381. 1949.

Colonies on C A (colour plate 132, A and a) spread ing braadl y , attaining a

d iarneter of 60 mm in 14 days at room temperature (20- 25C), gene ra lly flat
but in sorn e strains irregularly furrawed , str ictly velvet y in typi cal strains ,
cspec ially in new isolates, with thin m argin surro unde d by a wide fringe of
su bme rged hyphae exte nd ing abou t 4 mm beyond th e ae rial gro wth ; heavily
sporing th rough out from pale blue-gr een (Sgu y , PI. XXX, 438) to deep blue
gree n (Sguy, PI. XXX , 447 ) with th e rip ening of coni d ia; exudate and od our
lacking ; colony reverse colou rle ss in sorne strains but gene rally in yellowish,
orange , brown (colour plat e 132 , a) , or m ore com monly pink sha de s; peni cilli
typically bi verticillat e asy m metrica l (fig. 132, A) , produced in grea t
abundance; con id iopho res smooth wa lled, 100- 200 p.m by 3 .2 - 4 .5 tui, m ostly
a rising fram th e subs tra tu m in a close sta nd as to produce a velvety
appearance , occ a sionally simple (appearing monoverticillate) but gene ra lly
biverticilla te asy m me trical, consisting of ve rticils of 2- 3 metulae , 0 1' branches

CJo
00

O Ooa

00
o

1i ,
L--...J

100 ).1m

at.

Fig. 132. Camera lucida

drawings of Penici/lium
atramentosum Thom , CSS
291.48. A , Detailed drawings
of the penicilli. B, Mature
conidia. e, Habit sketches .

a nd m etulae a risin g from th e same lev el , with all elements a pp re sse d a n d

producing co n id ia l chains in well-defined colu m ns up to 500 uru or m ore in
length (fig. 132 , C), a p pear ing silky when obse rve d under ve ry low
magnification , becoming ta ngled in age a nd ad he r ing with adjacent co lu m ns
forming a deep la yer or crust of con id ia whi ch break s off as indicat ed aboye;
branches, when present , are b orne sin gly, usu all y 10- 35 p.m by 3 .5 - 4 .5 p.m ;
me tulae co m monly in grou ps of2 -3 ele men ts in th e ve rticil, m ostly 15 - 30 p.m
by 3 .5 -4.0 utn , with a pices en la rge d in a vesicle up to 7. 0 tui: in di amet er;
phialides parall el , in clu sters of 6- 10 ele rne n ts in th e verticil, 13-19 utt: b y
3.5 -5 .0 usn , tape r ing to a co ns p icuous short n arro wed neck ; co n id ia stro ngly
a nd persistently ellip tical, smooth wa lled or n early so (fig. 132 , B , and SE M
pIa te 22 , bottom: right), with co ns picuo us con nectives between th em , mostly
5 - 7 p.m b y 3 .5 -4 .5 tuu : Amber colou re d pi gm ent diffusing in to th e
surrounding agar (colour pl ate 132, A a n d a) .
Colo n ies upon CYA (co lour plat e 132 , B a n d b ) spread ing broadly , a ttai ning
a di amet er of 60 mm in 14 d ays at room temperature (2 0-25C), cons ist ing of

close-rcxturcd basal Iclt , typ ically velvcty, stro ng ly rnd iatcly urrowed and
rlcprcsscd in th e ce n tre , ov erl aid hy a thin , closely wo ven network 01' aerial
hyph ae inllucncin g somewhat th e colony appcarancc ; with a thin m argin
su rr ou nded by a fringc 01' submcrged hyph ae cx te nd in g about 3 mm beyond
h ae rial gro wth; heavil y spo ring throughou t , dev eloping gray-gr een shade s
(SI:gUY, PI. XXIX , 428 ) instead 01' the deep blue-green sho w n by th e
pr cccding ; ex udate a nd odo ur lacking ; no colou re d pi gment diffusing into the
su rr ou nd ing aga r ; colony reverse (col our plat e 132, b) devel oping reddish
h rown shades in th e ce n tre, else whe re cin namo n colou rs; p eni eil1i esse n tia l1y as
. lcscr ibe d above .
Colon ies upon M EA (co lour plat e 132 , C a n d e) sp read ing b ro adly ,
a tta ining a diamet er 01' 60 mm in 14 days a t room temperature (2 0- 25 C ) ,
l'llns isting 01' a looser textured b asal felt , very hea vily sporing forming deep
rru sts as not ed a boye, partiall y obsc u re d, es pecia lly in margin al a reas, by an
.ibundant ove rg rowt h consisting 01' a loosely woven net work 01' deeply floccos e
nc rial hyphae, influencin g greatly th e colo ny ap pearance; not furrow ed; od ou r
.m d cxu da te lacking; pale a m ber-colou re d p igment d iffu sing in to th e su rro u ndin g agar; colony rev er se (colour plate 132 , c) d ev eloping d eep purple brown
shades in th e ce n tre , elsewhere sh owing cin n a m on tints ; penicil1i essen tial1y as
dcscr ibed ab oye .
T he spec ies d escription is ce n te re d upon typi cal cu lt u res suc h a s C BS 219 .30
= NR R L 787 , isolat ed by C u rr ie a n d Thom in 191 5 , strain C BS 358 .48 = C M I
:19 ,750 = ATC C 10 ,476 = N R R L 790 = Q M 6 ,747 , isolat ed in 1934 from Blu e
Danish cheese b y S . H all , a n d upon m any othe r strains isol at ed from soil in th e
Pr ovin ce 01' N av arra by DI'. A .T. Martnez .
Penicillium oxalicum wh en long m aintained in laborat ory a rtificial cu ltu re
sho ws a m arked tendency to produce grad ual a n d progressive va r ia tion away
from th e or igin al, espeeial1y an in clination to be come more lanat e and less
hea vily spo rin g . It is cosm op olita n in its habitat and wo rld wide in distribution .
I t is espccial1y frequently isolated from soil a n d from plant residues undergoing
slow decomposition .
1I

133. Penicillium atramentosum Thom : fig. 133, colour plat e 133, and SEM

plate 23 , (to p : left).

Penici//ium atramentosum Thom - Bull. Bur. Anim. Ind. U.S. Dep. Agre. 118:
65-66.1910. See also Raper and Thom, 'A Manual of the Penicillia' , pp. 381-383.
1949.

Colonies upon CA (colour pl at e 133, A and a) attain ing a diamet er 01' abo u t
45 -50 mm in 14 days at room temperature (20 -25 C) , consisti n g 01' a typi cal1y vel vety b a sal felt , co m para tive ly thin and flat, not furrowed, excep t 1'01'

-,

!! JI
e ~"""'
~l

~ ' '

L...--.....I

100pm

1t,

R.

Fig. 133. Camera lucida

drawings of PenicJllium
atramentosum Thom, CBS
291.48. A, Detailed drawings
of the penicilli. B, Mature
conidia. C, Habit sketches .

th e presence of limited ov ergrowth of white floccose hyphae in localiz ed secto rs

of th e colony surface ; margin ver y thin , surrounded by a wide fringe of
submer ged h yphae exte nd ing a bo u t 5 mm or more beyond th e ae rial gro wth;
heavily spo ring th roughout in deep gree n shades (Sguy, PI. XXX , 44 1);
exuda te limited in amount , as minute crys tal clear droplet s en me she d in thc
mycelial felt; odo ur faint, inde finite; pale a m be r-coloured pi gm ent diffu sing
in to th e su r ro u nd ing agar; co lony reverse (co lour plat e 133 , a) at first
developing light cin namo n shade s, be coming deep orange-r ed to m ahogany in
colony cen tre in age ; penicil1i ab u nda n tly produced , with con id iophore s
arising primarily from th e su bs tra tu m in a close and even sta nd; penicil1i
typically bi verticil1ate asymmetrical (fig . 133. A), va riable in size and pattern ,
com mo nly co ns isting of te r m inal ve rticils of 2 - 4 m ore or les s div ergent
metulae , occasional1y branch ed with one or m ore metulae borne upon each
branch , seld om single (appearing monoverticillate); con idiopho res va r iable in
len gth , co m m only 50 - 75 /lm up to 125 tux , mo stl y 3 .0 - 4. 0 /lm in di amet er,
with walls sm oo th; b ranch es wh en p rese n t , 13 -20 /lm by 2.5 - 3.5 /lm ; m etulae
1O-17/lm b y 2.5- 4 .5 /lm, with apices sligh tly enla rge d in a ves icle up to 5. 0
tui: in di ameter; phialides parallel , in crowde d clu sters of 3-10 eleme n ts in the

vc rt icil, moxtl y
nnrrowed ncck ;
:111 <1 SEM pl at c
rh u in s ad he ring

ll -1 2 tlll by :1. 0 - 1'.0 UlI, ah ru ptly tape red lo a vel'Y s ho rt

co nid ia su bglobosc, smoo rh-wa llcd 01' ncarl y so (Iig . l:n, B,
23, top : len) , :1 - 4 UlI by 2.8 - 3.5 tm in d ia rne te r ; co n id ia l
in to poorly defined colum ns (Iig . 133, C) .

C olon ice upon C YA (colour pia te 133, B a nd b) spread ing broadl y , attain ing
a diam eter (JI' 60 mm in 14 days a t roo m temperature (20 - 2S0C), cons isting 01'
a iypi cally velvety basal felt , with wid e flat m argin (ab ou t 10 mm in width)
su rrou nded by a frin ge 01' subm erged hyphae exte nd ing abo u t S mm beyond
II\(: acrial growth; su bmarg ina l and central are as deepl y and stro ngly sulc ated
Iy nu merou s radial furrow s and ove rla id by a thin overgrowth 01' fine ste rile
hyph ae influ en cin g th e colony appearance; heavily sporing th rou gh out in deep
grccn sha de s (S guy, PI. XXX , 441) particul arl y cons picuo us in m a rgin al
arcas , elsewhere partially obs cured by th e thin overgrowth 01' pinkish coloured
xtcr ile hyph ae ; odo ur lackin g ; exudate limited in amo u nt, as crys ta l-clear
.lroplets enmes he d in th e myceli al felt ; purple re d to vinaceous coloured
pigment diffusing in to th e surround ing aga r; colony reverse (colour plate 133 ,
h ) develop ing deep purple red to che rry red shades; penicilli es sent ially as
.lcscribed aboye.
Colonies upon M EA (co lour plate 133, C a nd c) at ta ining a di amet er 01' 55
111m in 14 days at room temperature (20- 2S0C), cons isting 01' a looser textured
basal felt , typi call y velve ty, azo nate, flat , not furro wed, with very thin m argin
xurrou n de d by a frin ge 01' submerged hyphae exte nd ing abo ut 5 mm 01' more
bcyond th e ae rial growth; heavily sporing throu gh out in blue-gr een shades
(Sguy, PI. XXXI , 462) wi th th e ripe ning 01' co ni d ia; colony surface showing
locali zed secto rs 01' sterile whi te hyphae slightly obsc u ring th e heavily spo ri ng
ar cas, lending grayish green tin ts to th ese sec to rs; no coloured pigm ent
d iffusin g in to th e surrou nd ing agar; colony reverse (colour pia te 133 , c) at first
colo urless becomin g ora nge in age with a green cast sho wing throu gh ; pe nicilli
cssentially as described aboye .
T he species description is ce nte re d upon th e typ e stra in CBS 29 1.48 = C M I
:19 ,752 = AT C C 10,104 = N R R L 795 = I FO 8, 137 = Q M 7,483, isolat ed by C h.
Thom from Fren ch Came m be rt cheese a t Storrs, Con necticut, U. S .A . in
1905. T he cultu re has rema ined remarkably sta ble during more th an 7S years
that it has been under labor atory cultivation .
Penicillium atramentosum does not appear to be wid ely d istributed in natu rc
out re prese nta tives 01' the spec ies are occasionally isolat ed from soil.

134 . Penicillium fe nnelliae Stolk: fig. 134 , colour plat e 134 , and SEM

plate 23 , (top : right).

Penicillium fennelliae Stolk - Antonie van Leeuwenhoek. 35: 261-264. 1969.

~(

I.~

1
te
1,
tM
,
~

u..w..4:J..J

10

jrrn

I/?
fi< (Ji
~~ \?)

<\l>: !6~~
/?.

Fig. 134. Camera lucida

drawings of Penicillium
fennelliae Stolk, CBS 711.68.
A, Detailed drawings of the
penicilli. B, Mature canidia.
e, Habit sketches .

Stolk's original diagnosis follows, since this adequately presents the species in
question when first isolated:
'Colonies on Czapek agar growing restrictedly, attaining a diameter of 30 mm
within 2 weeks at 25C, consisting of a thin, buff to orange, ultimately pinkish basal
felt bearing abundant conidial structures, velvety, azonate , commonly buckled in
central areas and furrowed in a radial pattern, sporulating throughout the colony
but heaviest in submarginal areas. Conidiaf areas showing blue-green shades
approximating artemisia green, with the very narrow growing margin light celandine
green (Ridgway, PI. 47); when young surrounded by small orange zone, approximating orange cinnamon (Ridgway, PI. 29), consisting of submerged vegetative
mycelium studded with orange granules. Sectors which sporulate less abundantly
show orange brown shades approximating cinnamon when young, becoming
pinkish near Iight corinthian red with age (Ridgway, Pis. 29, 27). No macroscopically visible exudate drops are produced. Reverses of colonies showing orange

brown colours, ranglng fram cinnaman-rufous at tho margln thraugh hay's russet to
hnornat ite red in the centre, becaming dark purpla near hay's marcan with age
(Rldgway, Pis. 14, 27, 13) with the surroundlng agar caloured in Iighter tints of the
lino shades but at length becoming intensely coloured.
Colonies on malt agar growing more rapidly, attaining a diameter of 55-60 mm
w lthin 2 weeks at 25C, plane, velvety, very thin, azonate; conidial structures
produced throughout the culture but generally in greater abundance in central than
In marginal areas, showing blue-green shades near artemisia green (Ridgway, PI.
47) against a background of the orange coloured, largely submerged vegetative
mycelium. Marginal zone broad, showing conspicuous orange colours near
i lnnarnon-ruf o us (Ridgway, PI. 14). and consisting of radiating hyphae with the
jreen conidia l areas following the hyphae in unevenly radiating lines. Exudate
luckinq. Reverse of colonies show orange shades near zinc orange and cinnamonrufous (Ridgway, Pis. 15, 14).
Vegetative hyphae rather coarse, studded with orange granules uneven in diameter, 4.5-6.5 .m wide, often showing conspicuous swellings, especially on malt
Iga r. Conidiophores arising from the basal felt as well as directly from the substrat um, hyaline, septate, smooth-walled and fairly long, ranging from 250-500 .m
in length by 2.5-4.5 .m in diameter. Penicilli typically biverticillate and asymmetric,
vmying in complexity, but usually showing one or very rarely two rami in addition to
the main axis, each bearing like the main axis verticils of two to five metulae
torminating in clusters of phialides. Occasionally metulae and phialides arise from
the same level. Branched metulae may occur. Smaller heads, only consisting of a
verticil of metulae with phialides are also presento AII elements of the penicillus are
slight ly appressed, hyaline and smooth walled. Rami variable in length, ranging
from 14-35 .m by 2.5-4 .m in diameter, usually slightly enlarged at the apex up to
3- 5 .m in diameter. Metulae measuring 10-16 .m by 2.5-4 .m, enlarged at the
apex to 3.5-5.5 .m in diameter. Occasionally secondary metulae develop agreeing
in shape with the primary ones. Phialides occurring in fairly compact clusters of four
to six, with their axes roughly parallel, consisting of a large nearly cylindrical basal
part tapering distally to a small conidium -bearing tube (neck). Phialides usually
measuring 9-14 .m by 2-3 .m , occasionally larger up to 20 .m in length. Conidia
varying from cylindrical to ellipsoid with their bases sometimes flattened, hyaline,
extremely variable in size, commonly 3.5-5 .m by 1.5-3.0 .m, but occasionally
larger up to 10 .m, showing finely roughened walls and producing tangled chains
up to 150 .m in length. Neither ascocarps nor sclerotia were observed.'
Our not es follow:
Colonies u po n CA (colour plat e 134, A and a) in our observatio n co nform
Iairly well with Stolk's orig inal d iagnosis ei the r in rat e of growth or in ge neral
cultural characte ristics and in detail s of morpho logy (fig. 134, A, B and C), bu t
with conid ia co ns isten tly larger , ranging u sually from 5-7 tut: by 2.5 -4 .5 tut ,

co n sp icu o u sly spinulose when obscrvcd lindel' th e light mi croscope (Ii g. 1:H ,
B) and granular wh cn ohscrvcd undcr th c S EM (plat c :,n , IOp: righr).
Colonies upon CYA (colour plate 134, A and a) growing fast cr, auaining a
diameter of 45 mm in 14 days at room temperature (20-25C), con sisting 01' a
clos e-textured basal orange felt, typically velv ety, strongly wrinkled in a radial
pattern, conspicuously zanate and umbonate in the centre; with thin margin
surrounded by a fringe of deep orange-red coloured submerged hyphae
ext ending about 1 mm beyond the aerial growth ; sporing abundantly in central
areas in deep blue-gray shades (Sguy, PI. XXXI, 461) in conspicuous zoncs
with th e ripening of conidia, remaining persistently orange elsewhere; exudate
and odour lacking; reverse of colonies (colour piate 134, b) developing very
dark purple-red vinaceous shades ; similarly coloured pigment slowly diffusing
into the surrounding agar; penicilli essentially as described aboye .
Colonies upon MEA (colour piate 134, C and c) in our observation are
duplicating those described in the original diagnosis and heavily sporing
throughout in blue-green shades (Sguy , PI. XXX, 437, 438).
The species description is centered upon the type strain CBS 711.68 = CMI
151,747 = ATCC 22,050 = IJFM 7120, isolated by DI'. R . Bouillenne , Institut
Botanique, Liege, Belgium from Congolese soil in 1956 . It is at present known
only from the type species .
Penicillium Jennelliae appears to be most satisfactorily assignable to the
Asymmetrica-velutina subsection near the P. oxalicum series , on account of its
typically biverticillate and asymmetrical penicilli and its velvety cultural
appearance . It clearly differs from both P. oxalicum Currie et Thom and P.
atramentosum Thom in the shape , size and roughness of its conidia and in its
cultural appearance .

Penicillium digitatum series

OUTSTANDING FEATURES

Colonies upon Czapek' s agar growing very restrictedly , characterized in

Penicillium digitatum Saccardo by a limited submerged vegetative mycelium
bearing few and very irregular, asymmetric conidial structures, and in P.
japonicum Smith by a strongly umbonate and floccose colony not exceeding 20
mm in diameter.
Colonies upon Czapek's yeast extract and malt extract agars growing
luxuriantly, spreading broadly , loose textured, velvety to floccose , heavily
sporing throughout in dull yellow-green, becoming grayish olive in age ; odour
pronounced , strongly aromatic . The principal species has be en variously
described as the 'gray-green rot ' of citrus fruits .

lo uid iop ho rc s coa rsc, co m para tively sho rt , ari sin g lrom su b mc rgc d hyphac
111'

lrom the basal Iclt, with walls srn ooth.

Pcnicilli asymmctric, very irregular in siz e and pau crn, co rn para tivc ly vcry
IlII'ge, with id cntity 01' branchcs ami m ctulue olt cn poorly csta blishc d , bcaring
phialidcs a nd con id ial cha ins al vari ou s lcv els in th e co n id iopho re .
Co n id ia a t Iirst cy lind rical, usu all y becoming clliptical 01' occasionall y su b~ l () l>o sc , extre mcly variable in size but larger than in a ny othe r ser ies.
'I'h c seri es is ac tually represented by two sp ecie s, Penicillium digitatum
Saccard o a nd P. j aponicum Smith . The former is we ll kn own 1'01' being responsib lc 1'01' ser iou s losses 01' citrus fruit s in com me rce. In its natural habitats , thi s
mould produces a n olive-green colou red rot th a t ca n gene rally b e easily
disl in guished from th e 'blue-green rot ' 01' citrus fru its produced by m embers 01'
lite Penicillium italicum series. The latter spec ies isol at ed a nd fir st d escribed b y
Al: without indicating its origin is known only from th e type strain . It
rx -scmble s P. digitatum only in th e size a nd th e shape 01' th e co n idia .

1:1 5. Penicillium digitatum Saccardo: fig. 13 5 , colour pl at e 13 5 , and SEM

plate 33, (centre: Icft).

lu..u..W..uJ

10 }lm

\J

c:000
) o

fR .

Fig. 135. Camera lucida

drawings of Penicillium
digitatum Saccardo, CSS
319.28. A, Detailed drawings
of the penicilli. B, Mature
con idia .

Peniclllium digit8tum Saccardo - Sylloge Fungorum, Vol. IV: 78. 1886. Sea also
Ch. Thom, Bull . Bur. Anim . Ind. U.S. Dep. Agric. 118: 31-33. 1910; also Ch.
Thom, 'The Penicillia', pp. 242-245. 1930; also Raper and Thom, 'A Manual of the
Penicillia', pp. 386-390. 1949.

Synonym

Penicillium olivaceum Wehmer - Beitrqe zur kenntnis einheimisher Pilze. 11.

Untersuchungen ber die Fulnis der Frchte, pp. 1-84. Jena. 1895. See also Ch.
Thom, 'The Penicillia', p. 245. 1930. Also Raper and Thom, 'A Manual of the
Penicillia', p. 390. 1949.

Coloni es upon C A (colour plat e 135, A) grow ing ve ry res trictedly, from
punctiform to 10 mm in di ameter in 14 days at room temperature , consisting
mo stly of subme rged vegetative mycelium from whi ch a few conidial stru ctures
a rise of sm all dimen sions and irregular pattern , presenting without any doubt
th e gen eral aspec t of moulds suffering from sorn e typ e of nutritional deficiency
(a vitamin deficiency in th is particul ar case).
Colonies up on C YA (colour plate 135 , B a nd b) growing lu xuriantly ,
broadly spreadi ng, attaining a diameter of abo u t 40 mm in 14 day s at room
temperature (20-25 C) , cons ist ing of a den sely textured basal felt, typically
velve ty, flat in submarg inal and m argin al a reas and irregul arI y rai sed ancl
wrinkled in th e centre in a crateriform pattern , with undulating thin margin
surrounde d by a frin ge of sub me rge d hyphae extending abo ut 5 mm beyond
the ae rial gro wth ; heavily sporing th roughout in yellow- green to olive green
shad es (Sguy, PI. XXV , 373), show ing sorne zonation; exudate lacking;
odo ur pronounced , aromatic; no colou red pigment diffusing in to th e su rrou nding ag ar; colony reverse (colour plate 135, b) developing redd ish brown
shades; pe nicilli cha racte rized by great irregul arity in th e number and
dimension s of parts (fig. 135, A), usu all y cons isting of an indefinite number of
bran ch es a nd/or m etulae su ppo rting a limited but va riable seri es of phi alides,
eac h producing a cha in of co nidia that a re typi call y elliptical bu t showing wide
individual variation ; con idio pho res very sho rt , com mo nly rangin g from 30 to
100 Jlm in length by 4. 5-6 Jlm in di am eter, smooth walled , arising from
submerged hyphae or from th e basal fe1t; penicilli typi call y biv erti cillate
asym me trica l but va ry ing greatly in dimen sion s and in com plexity, with
branch es, metulae and phialides often poorI y defin ed ; cons isting of bran ches
vari abl e in len gth , bearing eithe r metulae and ph ialides, or phialides alone;
metul ae variabl e , com mo nly rangin g from 15- 35 Jlm by 5-7 uit, and bearing
small clu sters of phi alides eq ually va riable, rangin g from 10 to 30 Jlm by 4- 5
iuu , more or less cylindrical or bottle shap ed, terminating to a broad en ed sho rt
neck ; con idia at first cylindrical, becoming elliptical, sm ooth and thick wa lled
(fig. 135 , B, and SEM plat e 23, centre: left ), va ryi ng greatly in shape and

,J I I

t1 iI1II: IlS iO IlS ,

I'I'OIll

:L5 lo 5.0 un by

:LO -:~.5 /.tlll al

Iirst , thcn 6- B .t1l1 by

1.11 -6 .0 .t lll , and occasion ally up to 10-1 2 .tlll by 6 -8 .tm ; conid ial cha ins
v/ll'ying grea tly in lcn gth , up lo 150 .tlll , usuull y di vergent a nd com mo nly
I w('(lllling cnta ng lcd with other cha ins Irom [he sa rne on adjacen t pen icilli.
Colonies upon M EA (colour plat e 135, C an d c) sp read ing rapidly, attaining
1 d iamctcr of 40 -60 mm in 14 da ys a t room ternperature (20 - 25C ), cons isl ill f.{ 0 1' a looser textured basal felt , thin , flat , not fu rrowed , with very thin ,
irrc gula rly di ssected m argin surrau nde d by a fringe of su bmerged hyphae
l'xlc ndi ng abo u t 4 mm beyond the ac rial growth ; heavily spo ring throu gh out ,
uppoa rin g den sely velve ty ; conid ial areas developing pale yellow-green shades
(SI;gUY, PI. XXV , 366); cxudate lacking ; odour pronounced , a ro matic ; no
I'II lourcd pigment d iffus ing int o th e surroundin g agar; colony reverse (colour
plat c 135, e) uncoloured 01' in d ull tan shade s with a pale grc en ca st showing
hrou gh: pen icilli essentially a s describe d aboy e .
T he spe cies description is cc nte red up on strain C BS 3 19.48 = I MI 39, 748 =
"TCC 10,030 = NRRL 786 = I F a 7,006 = Q M 7,552 = IJ FM 722 1, isolat ed by
( :11 . Thom in 1904 at Storrs, C onnecticu t , USA , fro m Citrus sp . , and
nume rous strains mor e rccently isolated by th e author in Spain, from Citrus
uurantium Ri sso , Cit rus vulgaris Ri sso , Citrus limonium Risso , Citrus medica Risso ,
a lid from Citrus deliciosus Risso .
T he description as presented ade quately covers all but a n occasion aI
in cmber of thi s spec ies a s th ey a re isolated from fruits collected frorn packing
plants, sto rage wareh ous es, a nd market s. OccasionaI variants have been
r-ncountered . O ne suc h form producin g wh ite conid ia and showing no green
ro lour was isolat ed by DI'. H .S . Fawcet t, Ri verside , C aliforn ia , U SA in 1924 ,
.md was citc d in Thom ' s M onogr aph ( 1930 , p . 245) as a new va riety ,
l 'enicillium digit atum Saccardo va l'. calijornicum Thom . It has been reported as
dcstructive of ora nge s in th e u sual olive green form . The strain was soon lost
lrom Thom ' s collection and ha s not again bee n rep orted . T he occ urre nce of
white 0 1' ta n varia nts has been obs erved in m any spec ies, and the strain upon
which Thom based his va riety was undoubtedl y of such or igin o

136 . PenicilliumJaponicum Smith : fig . 136 . colou r plat e 136, and SEM

pIat e 23 , (cen tr e: ri ght ).

Penicillium japonicum Smith - Trans . Brit. Mycol. Soco 46: 333. 1963.
Synonym
Penicillium digitatum Saccardo var./atum Abe - J. Gen. Appl. Microbiol. Tokyo.
2: 97-98. 1956.
Smith's original diagnosis follows, since this adequately presents the species in
question when recently isolated:

:17B

ll..uWJ.uJ
jirn

10

.f?

Fig. 136. Camera lucida

drawings of Penicillium
japonicum Smith, CBS
327.59. A, Detailed drawings
of the penicilli. B, Mature
conidia . C, Habit sketches,

'Colonies on Czapek agar spreading rapidly and broadly, velvety, rather pale bluegreen, with a very broad margin consisting of a loose web of interlacing hyphae,
almost white; reverse reddish brown except that there is often a central spot which
is almost colourless; penicilli typically asymmetric, either consisting of a compact
cluster of metulae and phialides or having two to several rami bearing metulae, or
occasionally very large and complex with four stages of branching, the totallength
being 50-80 p'm; conidiophores mostly arising from the submerged mycelium,
smooth, often sinuous, 3-4 p.m in diameter; metulae variable but mostly 10-16 p.m
by 3-4 p.m; phialides remarkably uniform in length, 12 p.m by 2.8-3.0 p.m; conidia
very variable in size and shape, mostly ovoid to elongate, occasionally pyriform,
smooth or with a slightly uneven surface, 3.5-7 p.m by 3-4 p.m, in chains, roughly
parallel at first, then tangled.
The varietal epithet proposed by Abe refened to the spreading growth of the
isolate . However, P. digitatum, on a suitable culture medium, spreads extremely
rapidly, so the epithet is an unfortunate one. More serious is the fact that the variety
resembles P. digitatum in the size and shape of the conidia. It differs in surface

;17!}

i olour. reverse col our, and ty pe of penicillu s, also in it s ability to grow well on
ipe k agar. It is difficult to prepare slides of P. digitatum, the penicilli being frag ile
'II1d rarely found unbroken in f luid mounts. P. jeponicum , on the cont rary, is easy to
rnoun t, and , provided that the culture is not too old , readily gives slides showing
ores of intact penicilli . The varietal epithet used by Abe is unsuitable as a specific
oplthet , so it has been cons idered adv isable to provide a new name for the fungus.'

S . Abe (1956) , in a n ex te n sive paper on the tax onomy of th e genus

l'micillium , d escribed 30 new sp ecies, varieti es and mutants , none of which
wr.rc va lidly published , a s th ey were without L ati n diagnoses . 26 cu lt u re s of
IIlese stra ins were sent by Abe to th e Commonwealth M ycological Institute of
Kc w , Surrey, UK, where th ey were exam ine d by G . Smith ( 1963) . The names
1'1' t he new species bestowed by Abe , where th ese were judged to represent
hil he rto u nde scr ibed species, were validated b y G . Smith ' s Latin di agnoses
iI rn wn u p fram his own observ a tio n s. On th e othe r h an d , two of th e varietal
e pilhct s were judged by Smith as unsu itable to serve as sp ecific ep ithe ts , a nd
I he spec ies were redescribed under new names . Such is th e case of Penicillium
jojllmicum Smith , formerI y d escribed by Abe a s P. digitatum Sa ccardo var o latum ,
.md P. abeanum Smith , de scribed by Abe as P. trz ebinskii Z al eski var magnum
( S(T th e Monoverticillate) .
During th e more than 25-year period that Penicillium japonicum Smith has
I x-en m aintained in la bora to ry cu lt u re, th e sp ecie s h a s ex h ib ited a gradua l and
progressive va r ia tion a way from th e o r iginal, particularIy co ncerning th e ra te
01' grow th upon Czape k' s agar a n d th e cu ltu r al a p pea ra nce on the same subst raturn .
Note s drawn up frorn our ow n observations foll ow :
C olo n ies u po n CA (col our plate 136 , A and a) growing restrict edly instead of
ra pidly a n d braadly , as stated by both Abe a nd Smith in the ir respective
c1cscriptions of th e sp ecies , a ttain ing a di ameter of 10- 20 mm in 14 d a ys a t
room temperature (20- 25 C) , co n sisting of a rather clos e-textured b as al felt ,
slrangly co nve x , not furrow ed , with thin margin su rrou nde d by a frin ge of
rcddish brawn su b merged hyphae ex te nd ing about 5 mm beyond the ae ria l
growth , with colony su rface ove rl a id b y a d eep loosely wo ven network of whi te
lloccose ae ria l h yph ae influencin g greatly th e colo ny a ppe a rance, and len d in g
it a floccose to la n a te a sp ect ; poorIy spo r in g not infl u encing the co lo ny
a ppe a rance; odour faint, a ramatic; exudate lim ited in amount , as minute
colou rless draplet s en m eshe d in th e m ycel ia l felt ; no co lo ured p igment
d iffusin g into the surro u nd ing a gar; colo ny reverse (colour plate 136 , a)
develo p ing d eep reddish b raw n co lou rs.
C olonies upon C Y A (co lour plat e 136 , B a nd b) spreading b raadly , a tta in ing
a di amet er of 65 mm in 14 d a ys a t room temperature (20- 25 C ) , co ns isting of
a looser textured ba sal felt , overIa id in su b marginal a nd ce n tral a re a s by a

.J O\ I

dense nciwork 01' white Iloccosc vegetativ o hyph ac influ cn cin g gr eally th c
colony appearance ; with margin a ppearing gra n ular owing to thc production
of abu ndant synnem at a ; colony su r face sulcated by sha llow radial f'urrows a nd
slightly rai sed in ce ntral areas; heavily spo ring throughoul in pale blu c-gr ccn
sha des to blue-gray tints (Sgu y , PIs XXIX, 435, and XXX , 449) ; ex uda re
and odo u r lacking; dark a m ber colou re d pigm en t diffusin g int o the
su rrou nd ing agar; colony reverse (colour plate 136, b) stro ng ly pulling the ag ar
away fram th e cult ure dish a nd cracking it , dev eloping deep reddish brown
sha des; pe nicil1i esse ntially as described by G . Sm ith but with all elernents
la rger ; con id iopho res m ostly a risin g from the su bs tratum, m ore 0 1' lcss
ag gregated into erec t bundles 0 1' synnemat a , parti cul arl y cons picuous in
margin , e1sewh ere th ey are obscu red by th e den se ove rg rowth of white floccose
hyphae , measuring up to 200 tm in len gth by 4-6 uit: in di amet er; branchcs
whe n present m ostl y 20- 40 utt: by 4-5 tm ; m etul ae variable but m ostly 20 -25
tm by 3 - 5 tm ; phialides mostly 15- 18 tm by 3.5 - 5.O iut , tap ering to a
conspicuo us ly long narrowed neck; con idi a varia ble in size, usuall y strang ly
elliptical , m ostly rangin g from 4 .5 to 9.0 tm by 3.0 -4 .5 tut , srnoo th-wa lled or
nearl y so (fig. 136, B, and SE M plat e 23, ce ntre: right) ; con id ial chains at first
rou ghly parall el (fig . 136 , A) , th en becomin g tangled (fig . 136 , C) .
C olonies on M EA (colou r plate 136 , C and c) gro wing rapidly a nd braadly,
attaining a di ameter of 65 mm in 14 days at raom te m pe rature (20 -25C ),
cons isting of a loos er textu red b asal felt , flat, not fu rrowed, with wid e a nd thin
m argin su rrau nde d by a frin ge of reddish brown-coloured , su bmerged h yphae
exte nd in g m ore th an 5 mm beyond th e ae rial gra wth, cons picuo us ly zo na te,
with colony surface appearing gra n ula r owing to th e numeraus fas cicles 01'
synnem at a arrange d in co ncentric zones in su bmarg inal a nd m arginal areas,
with ce ntre ove rlai d by a loosely wov en netw ork of floccose h yphae lend ing a
lanat e appearance to it ; heavily sporing throu gh out in gray-green shades
(Sguy, PI. XXX, 449) with fascicles most abu ndantly praduced along th e
int ercolon y m argin s; od our strong, suggesti ng fennel (A nethum f oeniculum L.) ;
exu da te la cking ; no coloured pigment diffu sing in to the surra u nd ing a ga r ;
colony reverse developing deep reddish brown shades (colour plat e 136, e);
peni cilli (fig . 136 , A) esse ntially as described aboye .
The species description is ce nte re d upon th e type stra in C BS 32 7.59 = C M I
68 ,233 = ATC C 18 ,317 = IFO 5, 747 = Q M 7,298 = IJFM 7129 , isolated by S .
Ab e , prabably fram soil , J apan , 1956 . It is known only fram the typ e stra in .
Penicillium japonicum Sm ith, as received fram th e Centraalbu reau voor
Schimmelcultures, Baarn , The Netherl ands, in 1979 , ac tu ally exhib its noteworthy va riation away frorn th e original, p articul arly the restricted grawth a nd
poor sporulation upon C zape k's agar culture m ed ium (colou r pla te 136, A) ,
prob abl y due to a vitam in defi ciency suffered by th e strain in question al't er a
long peri od of lab oratory cultiva tion, as it ca n c1early be inferred from th e

lu xuri ant gro wth ami ubu nda n t sp orulation upon C za pck ' a gar plus a dr lcd
viuu nins (as Difco yca st cx trac t) (SCl: (colour plat c 136, B) . Furthermorc, in
11111' obse rva tio ns , P. [aponicum grow n UpOll malt ex trae r agar develops a strong
1I 1ld rha rac ter istic odour suggesting fenncl and whcn ~rown upon CY A and
M EA (colour plate 136, B and C) produces abundant fasciculation, features
111)1 observed by previous authors .
A close relationship between Penicillium japonicum Smith and P. digitatum
Saccardo is not presumed here, and they are con sidered together primarily as a
u uutcr of eonvenienee. They do , however, show eertain features in eommon,
p.ut icularly the eomparatively large asymmetrie eonidial struetures and the
sha pe and size of conidia . However, in our opinion, the produetion of
xyn nc m a ta suggestive of a eertain relationship to speeies included within the
Asym me triea - faseieu lata sub-seetion should not be overlooked .

Penicillium roqueforti series

( h JTSTANDING FEATURES

C olonies eommony not always broadly spreading, typieally velvety ,

gcne rally thin upon maIt extraet agar , with abundant short conidiophores
.uising from trailing 01' submerged hyphae just below the agar surfaee, with
growing margin usually appearing more 01' less a rachnoid, namely, formed of
dclicat e hyphae ; with conidial areas typically developing dark yellow-green or
da rk blue-green shades with the ripening of con id ia, often showing greenish to
ulmo st blaek shades in th e eolony rev ers e.
Conidiophorcs eonspicuou sly roughened 01' warty throughout , with the
cxception of phialides .
Penieilli two-stage branch ed 01' biv ertieillate as ymmetrical, bearing conidia
cither in tangled cha ins 01' adhering into loos e colu m ns.
Conidia comparatively thick-walled and smooth 01' nearly so .
K ey lo species
Page
1. Penicilli typically rebranched below the level of metulae, with the main
axes and branches terminating in a verticil of metulae. Conidiophores
conspicuously rough walled; growing margin appearing usually
arachnoid.
P. roqueforti series . .. . 381
A. Conidiophores coarsely roughened, commonly appearing conspicuously warty 01' tuberculate.
1. Colonies broadly spreading upon all substrata, with surface plane
01' nearly so; thin margin appearing arachnoid; conidia cornparatively very large, attaining a diameter of 6.5 .tm.
P. roquefortiThom . ... 383

.) 0

2. Colonies growing rather restrictedly upo n CA; con idia not

exceeding 4.5 /lm in diameter.
P. farinosum Novobranova
B. Conidiophores conspicuously but not coarsely roughened but w ith
walls appearing finely granular; conidial areas developing pale yellowgreen to artemisia green shades; con idia comparatively small , not
exceeding 2.5 /lm in diameter.
P. fag i Martnez et Ramrez
C. Conidiophores with wall appearing fine ly spinulose.
1. Conidia l areas upon MEA developing olive green shades; conidia
large, up to 5 /lm in diameter.
P. mali Novobranova
2. Conid ial areas upon MEA develop ing deep blue-green shades ;
conidia not exceeding 4 /l m in diameter .
P. cordubense Ramrez et Martnez

. . . . 391

.... 385

. . . . 389

.... 387

The Penicillium roquejorti series a s u nderstood by Raper and Thom ( 1949) was
represented by two species, namely , Penicillium roquejorti Thom a nd P. casei
Staub, whi ch is now regarded as synonymou s with P. verrucosum D ierckx , va ro
cyclopium (Westling) Samson , Stolk et H adlok (Samson et al. 19 76).
In the present study, the series has been broadened enough in concept to
accomodate four additional forms as follows: Penicillium jagi Martnez el
R amrez, P. cordubense Ramrez et Martnez , P. mali Novobranova, and P.
jarinosum Novobranova .
The principal species of th e series, P. roquejorti Thom has been studied
particularly with relation to its role in cheese manufacture . lt is the
predom inant mould in the who le group of cheeses charac te rized by streaks, or
'marbling ' , of green mo ulds . C losel y related strains of thi s form appear in thc
true Roquefort , or sheep's m ilk chee se of sou the rn F rance, in the ' from ages
bleus ' , or blu e cheeses of cen tr al France, m ade either from sheep' s m ilk or
cow ' s mi lk , in th e Gorgonzola che ese from northern Italy, in th e Stilton ch eese
as made in England, in the Gammelost cheese as made in Norway, in the Bluc
chee se as made in D enmark , or th e American Roquefort cheese as made in the
US , and in many varicti es of loose-textu red che eses whi ch obtain their
charac teristic flavour from th e moulds which invade th e ha les and crac ks
throughout th eir mass .
A clas e relationship between Penicillium roquejorti Thom a nd the four
additional sp ecies incl uded now in th e series is not presumed , and th ey are
considered together primarily as a matter of con ve nience. They do, however,
show certa in characte ristics in com m on , particu la rly th e co nspicuou sly roughwa lled conidiophores, th e co m m only thin arac h noid margins, particularly
upon malt extract agar , th e dark green colou rs develop ed in th e colony surface
and in th e rev erse , and th e globose, thi ck walled and smooth conidia .

I :~ 7 . Penicillium roqud orti T ho m: ligo l :n , colou r plat el :n , a nd SEM

plato 23 , (bo tto m : lcft ).

ll..I.J.J..WW

10J.lm

o~ o

O
B
O ;"
00
0 0
00 0

0
' 0
0

,.ctlR

1~,/1

Fig. 137. Camera lucida

drawings of Penicillium
roqueforti Thom, eBS
221.30. A, Detailed drawings
of the penicilli . B, Mature
conidia. e, Habit sketches .

Penicillium roqueforti Thom - Bull . Bur . Anim . Ind . U.S. Dep. Agre. 82: 35-36.
1906. Also Ch. Thorn, ibid. , 118: 34. 1910; also Ch . Thom , 'The Penicillia ', pp .
277-279. 1930; also Raper and Thom, 'A Manual of the Penicill ia' , pp. 395-401.

1949.
Synonym ies
Penicillium aromaticum Sopp - Monograph ie, pp. 159-161. 1912. See also Raper
and Thorn, ' A Manual of the Penicillia' , p. 400. 1949.
Penicillium etro-viride Dierekx - Ann. Soe. Sci. Bruxelles . 25: 87. 1901. See also
Biourge's Monographie, La Cellule . 33: 199. 1923; also Raper and Thom , 'A
Manual of the Penicillia ', p. 399. 1949.
Penicillium etro-viridum Sopp - Monographie, pp. 149-150. 1912. See also Raper
and Thorn, ' A Manual of the Penicill ia.' p . 400. 1949.
Penicillium biourgeii Arnaudi - BolI. 1st. Sieroterapieo Milanese. 6: 25-27. 1927;
also Zentbl. Bakt. Parasitenk. Inf. Hyg. 11 Abt. 73: 321-330. 1928; also Raper and
Thom , ' A Manual of the Penicillia' , p. 401. 1949.

Penicillium gorgonzolo Weidemann - In Biourge's Monographie, La Callule. 33:

204-206. 1923; also Rapar and Thom, ' A Manual of tha Penicillia.' p. 400. 1949.
Penicillium roquefort Sopp - Monographie, pp. 156-157. 1912. See also Rapar
and Thom, 'A Manual of the Penicillia.' p. 399. 1949.
Penicillium roqueforti Thom var. weidemannii Westling - Arkiv. Bot. 11: 52, and
71-73. 1911. See also Raper and Thom, 'A Manual of the Penicillia.' p. 399. 1949.
Penicillium stilton Biourge - Monographie, La Cellule. 33: 206-207. 1923. See also
Raper and Thom, 'A Manual of the Penicillia.' p. 400. 1949.
Penicillium suaveolens Biourge - Monographie, La Cellule. 33: 200-202. 1923.
See also Raper and Thom, 'A Manual of the Penicillia.' p . 400. 1949.
Penicillium vesiculosum Bainier - Bull. Soco Mycol. France. 23: 10-12. 1907. See
also Raper and Thom, 'A Manual of the Penicillia.' p. 400. 1949.
C olon ies upon C A (co lour pl at e 137 , A and a) sp read ing b roadly, a ttain ing a
diamet er of abo u t 50 -55 mm in 14 days a t roo m temperature, typicall y
velvet y, plane , not furrowed, with thin m argin a rach no id or co bwe bby with
hyphae radiating p artly on the surface a nd partly j us t b elow th e surface of th e
subs tra tu m , ex te n d ing about 5 mm beyond th e ae r ial growth in unevenly
radiating lines; hea vily spo ring throu gh out in d eep blue-green shades (Sguy ,
PI . XXX , 44 1) with th e ripening of con id ia; od our a nd ex u da te lacking; a m ber
colou re d pi gment di ffusing into th e su r ro u n d ing agar; colon y reverse (colo u r
pla te 137 , a) d eveloping sh ade s of green, bluish green, bl ack ish brown to
almost bl ack , dep ending upon the strain ; penicilli variable in pattern (fig . 137 ,
A), ranging from simple structures (a p pear ing m on ov erticilla te) to ve rticils of
m etulae a n d phial ides, or var ious ly ram ified , with on e or more lon g a pp ressed
or di verging rami ; co n id iop ho res com mon ly sho rt , usu ally a bout 50- 100 tui ,
less fre q ue n tly up to 200 tm in length by 4. 5- 5.0 tm in di amet er, arising fro m
submerged hyphae or from trailing ae r ia l h yphae , co ns p icu ous ly warty
(occasionall y with wall s sm oo th) or gran u la r from enc rustmen ts of var iable
size; rami 16 - 30 tm b y 4 .0-4 .5 psn , mostly with wall s co ns p icu ous ly tubercu la te ; m etulae 14- 20 tm by 4 .0 -4 .8 tui, with walls m ore or less rou gh ened ;
phial id es b ottle shaped , 10-1 5 tm by 4 - 5 iuu , tapering to a m ore or less
cons p icuous ly sho rt n arrowed neck ; con id ia co m mo nly large , m ostly 4 .5 - 7.0
tm in diamet e r , globose to su bglobo se, wi th walls smooth or nearly so (fig.
137, B, a n d S EM 23, b ottom : left) ; con idial chains tangled or adhe r ing into
loose colu m n s (fig. 137 , C) .
Colo n ies upon C YA (colou r plat e 137 , B a n d b ) growing more rapidly than
aboye, attain ing a di amet er of JO mm in 14 d ays at room temperature
(20 -25 C) , con sistin g of a closer textured b asal fe!t , a p peari ng velvety,
stron gly a n d deepl y furrowed in a radial p attern , with ve ry thin m argin
surrounded b y a frin ge of su b merged hyphae ex te nd ing abo u t 4 mm beyond
th e ae r ial gro wth; colo ny su rface partl y ove rlaid b y a floccose ove rgro wt h of

whit c acrinl hypha c pnrtially obs curing the oth crwisc hcavily sporin g co nid ia l
ureus; co n id ia l arcas d cv cloping gray -grccn sh ades with th c ripcnin g 01' conidia

(Sg llY, PI. XXIX, 4:n); exuda re limir cd in amount , as minute colou rless
droplcts cnrnc shed in th e my cclial fclt ; odour lacking; no colou re d pigment
dilTusing into th e su rrou nd ing agar ; colony rcvcrs e (colour plat e 137 , b)
ilevcloping brown to lilacinum shades in th e cen tre, almost colou rless in th e
margin ; peni cilli essentially as described abuve .
C olun ies upon MEA (colour plate 137 , C a nd c) attain ing a di ameter of 65
111m in 14 days (20 -2 5C), consisting of a very thin loose-textured basal felt ,
iypically velvet y , with a very wid e arachnoid margin, largely sub m er ged and
uxtcnd ing abou t 10 mm beyond th e aeri al growth ; colony surface flat , not
urrowed ; heavily spo rin g throughout in blue-green shade s (Sguy , PI.
XXVII , 403); exudat e and odo ur lacking; no coloured pigment diffu sin g into
lile surro und in g agar; colony reverse (colour plate 137, e), at first colou rle ss
with a gree n cast showing through , blackening in age; conid ial struc tures
usually exh ib iting increas ed roughness and con id ial chains tending to form
\on ger a nd more com pa ct colu m n s (fig. 137, C) , up to 300 tut: in len gth .
Species description is cen tere d upon th e typ e strain CBS 221. 30 = CMI24, 31 3
= ATCC 10,110 = ATCC 1,129 = NRRL 849 = IF0 5,459 = QM 1,9 37 = IJFM
7201, isolated in 1904 from french R oquefort chee se by Ch . Thom , and from
nurnerous other stra ins possessing similar cultu ra l a nd morphological characleri stics that hav e been isolat ed by th e author from different sub strata . Among
such cult ure s now m aintained in our collection strain IJFM 3706, isolated by th e
au tho r in 1976 from Cabralc s chee se of no rthe rn Spain , made from cow's milk ,
ma y be m entioned and regarded as entire ly typical of th e species as it is
understo od by R aper and Thom (194 9). (Colour plates 239 and 240 present
lwo representati ve strain s of P. roqueJorti, isolated by th e autho r from Cabrales
cheese a nd from th e air , respectively .)

138 . Penicilliumfagi Martnez el Ramrez : fig. 138, Colour plate 138, and
SEM plate 23, (bottom: right).
Penici/lium fagi Martnez et Ramrez - Mycopathologia . 63: 57-59. 1978.

C olonies upon CA (colour plate 138 , A and a) a tt a ining a diamet er of 45 mm

in 14 days at room temperature (20- 25 C) , consisting of a rather loos etextured basal felt , typicall y velv et y , with m arginal and submarginal area s
plane and cent re deeply and irregularly wrinkled ; thin margin irregularly
di ssected, typi call y arachnoid 01' cobwebby, largely submerged in unevenl y
radiating lin es; colony surface ov erl aid by a very thin network con sisting of
closely wov en fine aeri al hyphae influ en cin g gr eatly th e colony appe ara nce;
heavily sporing throughout in yellow-gree n shade s (Sgu y, PI. XXVII, 395)

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Fig. 138. Camera lucida

drawings of Penicillium fagi
Martnez et Ramrez, CBS
689.77. A, Detailed drawings
of the penicilli. B, Mature
conidia. C, Habit sketches .
D, Submerged hyphae with
eonidial structures arising
from it .

with gree n co nid ia l areas fol1owing th e m argin al su b me rge d hyphae in an

unevenly radiating pattern ; odo u r and ex u date lacking ; colou re d pi gm ent
diffusing into th e surro u nd ing agar no t produced ; colony reve rse (co lou r plat e
138 , a) colo u rless to pale pinkish cream coloured; pe n icilli (fig. 138, A)
typi call y bi verticillat e and asym me tric va rying in com plex ity , bu t usu ally
showing on e or two rami in ad d itio n to th e m ain axis, and verticils of 3- 7 or
more m etula e beari ng clusters of phiali des , with all ele me n ts m ore or less
a ppressed; smaller co nid ial struc tures co ns isting of only a ve rt icil of metulae
bearing verticils of phialides, or simple penicilli (a ppearing m onoverticill at e)
are not unfrequently enc ou ntered; co n id iop hores mo stl y short, usuall y not
exceed ing 150 p.m in len gth b y 3.0-4.5 p.m in di amet er, a rising prim aril y fro m
sub me rge d h yphae and from th e basal felt , not septate, with wall s coarse ly
ro ug he ne d ; rami va riable, ra nging from 35 - 40 p.m in len gth by 3-4 p.m in
di amet er , u su all y slightly enla rged at th e a pex up to 3-4 p.m in di amet er, with
wal1s simi la rly rou gh ; m etulae rou gh-walled , m ostly 10-1 5 p.m b y 3-4 p.m ,
with a pices enla rge d up to 3 .5 -5. 0 utt: in di amet er; phialides bottle shape d,
occ urring in fairly com pact cluster s of 4- 12 elements in th e ve rticil, with th eir

.1111

nxcs rou ghly parall cl, a bo u t 7-10 J.t11l b y 2.:i -:L:i ,tlll , tapcrin g abruptl y l o a
sho rt narrowcd ncck ; co n id ia co m pa ra tivc ly vc ry small , 2 -2 .5 ,tm in di amct cr ,
globo sc, with wall s srnoot h 01' ncarl y so ( lig. 138 , B, a nd SEM pl at e 23,
ho uo rn: r igh t); eo n id ia l chains loosel y parall el beco m ing tangled in age (fig .
l :lS , C) .

Colo n ies upon C YA (co lo ur pl at e 138, B a nd b ) attainin g a d ia mete r of

a bo ut 55 mm in 25 days a t room te mpera ture (20 -25 C), consisting ofa close r
rcxtured basal fel t , deep er than a boye . Strongly and uniforml y wrinkled in a
radial pattern , with ce ntre irregularly folded a n d rai sed , with m argin en tirely
su rrou nded by a na r ro w fringe of su b merged hy p hae ex te nding abou t 1 mm
hcyond the aeria l growth; hea vil y sporing th ro ughou t in p al e blue-g reen al'
gray -g reen shades (Sguy, PIs X X X , 438 , a nd XXIX , 429) ; ex u d a te limited
in amou nt , as amber-co lo u red d ro ps; odo u r lack in g ; ve ry pal e amber-c olo u red
pigrn ent slow ly d iffus ing in to the surro u ndi ng agar ; colo ny reverse (co lo u r
plat e 138 , b ) developing ve ry d ark green shades (Sguy, PI. XXX , 446) ,
!>ecomin g almos t bl a ck in age ; penicilli esse nt ia lly as descri bed a b oye .
C olon ies upon MEA (colour pl ate 138 , C a n d c) a ttain ing 45 mm in
diamete r in 14 days a t room temperatu re (2 0 - 25 C ) , co ns ist ing of a loose r
textured b a sal felt , thin an d flat , not furro wed , with a rach no id m argin largely
su b m erged , ex ten d in g a bou t 3 mm beyond th e ae ria l growth ; hea vil y spor ing
th ro ugho u t in blue- green shades (Sguy, PI. XXX , 44 3) ; odo ur a n d ex u date
lac king ; no co lou red pi gmen t diffusing in to the surro u n d ing agar; colo ny
reverse (colour pl ate 138 , c) at fir st ye llow ish, th en blackening in age ; penicilli
essen tially as de scribed aboye but with wall s m ore coarsely rou gh en ed ,
alth ou gh smooth -walled m etula e m a y occa sionall y be ob serv ed (fig. 138 , A).
The spe cies description is cen tered upon th e ty pe strain IJ F M 3049 = CBS
689. 77 = ATCC 36 ,956 , isolated in 197 6 by DI' A .T . M artnez fro m Fagus
silvatica L. fall en leaves on an a n d osol in a b eech forest located n ear th e
O ro q uieta pa ss (800 m a lt itud e) , Province of N avarra , Spain .
Penicillium fagi is a ve ry di stinctive spe cies whi ch is bel ieved to be closely
related to species a ss igne d w ith in th e P. roqueforti series o n a ccou n t of its
coa rsely rou gh ened co ni d io pho res , th e a rach no id m argin sho w n by the
colon ies upon m ost su b strata, a nd th e thin , typicall y ve lve ty colo n ies b ecoming
bl ack in th e reverse. It is kn own o nly from th e typ e strain .

139 . Penicillium cordubense R amrez el M artnez : fig. 139 , colou r plate

139 , a nd SEM piate 24 , (top: left).

Penici/lium cordubense Ramrez et Martnez - Mycopathologia. 74:163-171. 1981.
C olonies upon C A (colour pl ate 139 , A a nd a) attainin g a d iamet er of 45 mm
in 14 da ys a t ro om temperature (20 - 25 C), co ns isti ng of a rather loose-

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Fig. 139. Camera lucida

drawings of Penicum
cordubense Ramrez et
Martnez, ATCC 42,238. A,
Detailed drawings of the
penicilli. B, Mature conidia.
C, Habit sketches.

textured b asal felt , typically ve lve ty , consp icuo u sly zonate, radiately su lcated
with shallow furrows, and with centre rais ed; with thin undulating margin
surrounded by a fringe of submerged hyphae ext ending about 2 mm beyond
th e aeri al gro wth; with colony surface ov erl aid by a thin , closel y wov en network of ae rial hyphae influencing greatl y the colony appe a ra nce ; heavily
spo ring in blue-green shades (Sguy, PI. XXVII , 402) with con id ial areas
partially obscured by th e thin overgrowth of aerial hyphae shading to
ochraceou s tints; exudate abundantly produced, as large dark amber- coloured
drops ; od our faint , ' m ouldy ' ; no coloured pigment diffusing into th e
surrounding ag ar; colony rev ers e in reddish brown shade s with a gree n cas t
showing through (colour plat e 139, a); penicilli (fig. 139, A) typically
bi verticillate asym me tric, charac teristically de veloped in a dense and eve n
stand to produce a velvet y effect, variously ramified , with con id iophores
/
conspicu ous ly spinulose, arising primarily from th e su bstra tum, consisting of
two or three sta ge branched struc tures ; con id iophores short up to 150 p.m in
len gth by 2.5-3.5 p.m in di ameter; rami ranging from 15 to 20 p.m by 3.0-4 .0
p.u , some what appressed , rou gh wall ed ; m etulae more or less cylind rica l,

. J U ;J

usually rough wall cd, mosrly 7 -14J.ull by :1.O - !Ul/UII with a p iccs c n la rged in a
vcsicle up to 4 -6 1t1ll in di um et cr, smoo th-wa lled metulae are occasionally
c nco un te red (fig. 139, A); phialidcs bottlc sh aped, 7-121tm by 2.7 -4.0 1tm,
lapering abruptly to a short-pointcd nc ck; co n id ia globose to subglobose ,
~ . 8 - 4 . 0 tut: in long axis, smooth wall ed 01' nc arly so (fig. 139 , B, and SEM
plato 24, top: left) ; co n id ia l cha in s tending to produce long, loosely parallel ,
c1i vergent colu m n s (fig. 139, c).
Colonies upon CYA (colour plate 139, B a n d b) growing m ore rapidly than
aboye, attaining a diameter of a bo ut 50 mm in 14 days at room temperature
(20- 25C ), co ns istin g of a c1ose-textured felt , typically velv ety , stronglyand
dcepl y wrinkled in a radial p attern , with ce n tre raised and fold ed in a
cra ter ifo rrn pattern , with thin , flat , undulatin g m argin surrounded by a frin ge
01' su bmer ged hyphae extending about 3 mm beyond th e aerial growth; very
heavily sporin g throughout in deep blue-green shades (Sguy , PI. XXX , 447)
with the ripening of con id ia, show in g ochra ceous shades in ce n tr al areas;
odour faint , indefinite ; exu da te limited in amount, as minute a m be r coloured
droplets enmeshed in th e m yeelial felt ; no colou re d pigment diffusing into th e
su rrou nd in g aga r ; colony reverse (colour pl at e 139 , b) developing reddish
cin na m on shades; penicilli esse n tia lly as d escribed aboye .
Colonies upon MEA (colour pi ate 139 , C and c) attaining a diameter of
a bou t 45 mm in 14 days at room temperature (20 - 25 C ) , con sistin g of a looser
tex tu red bas al felt, a p pea r ing vel vet y to powdery in heavil y spor in g areas , with
margin thin , a rac h noid, largely submerged a n d extending up to 5 mm b eyond
(he . a erial growth ; heavily sporing throughout in deep blue-green shades
(Sgu y , PI. XXX , 447) ; exu d a te and odour a b se n t ; colony su r fac e flat , not
Iurrowed; no colou red pi gment diffusing into th e surrounding a ga r; colony
rev ers e (col our pl ate 139 , e) developing very d ark reddish brown shades;
penieilli ess entially as des cribed a boye.
The sp ecies description is ce n te re d upon th e type strain IJFM 7030 = ATCC
42,238, isolated by th e author of the current stu d y from sand y soil, Madrid,
Spain, M ay 1979 . It is at present known only from th e type strain .
Penicillium cordubense is a distinctive species ch a rac te rized by its typically
velvety colo n ies, with ara chnoid margins, especially upon MEA , b y th e very
dark-coloured reverse upon MEA and its co n sp icuo u sly rough-walled
con id iopho re s. The correet plaeement of this sp eeies appears th en , to b e m ost
sa tisfae tor ily ass ign able to th e P. roqueforti series in the Asymmetriea-velutina
sub -seetio n .

140 . Penicillium mali N ovobranova: fig . 140 , colou r pIate 140 , and SEM
pIate 24 , (top: right).
Penici/lium mali Navabranava - Bial. Nauki. 15: 103-108. 1972.

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Fig. 140. Camera lucida

drawings of Penicillium mali
Novobranova, C8S 500.73.
A, Detailed drawings of the
penicilli. B, Mature conidia .
C, Habit sketches .

C olonies upon C A (colou r plat e 140 , A and a) att a ining a di amet er of abo u t
40 mm in 14 days at room temperature (20 -25 C) , cons isting of a rather loo setextured basal felt , typi cal1y velv ety , som ewhat zanat e, flat , with white undulatin g m argin surrounded by a narrow fringe of su bme rge d hyphae exte nd ing
abo u t 1 mm beyond th e aerial gro wth; heavily sporing throu gh out in deep
gre en sha de s (Sgu y , PI. XXVII , 401); with th e rip ening of conid ia , pa ssin g
through va r ious shades of blue-gr een (Sguy , PI. XXX , 446 , 447); odo ur
faint , ' mouldy ' , exudate abu ndantly produced , as dark am ber drop s; red dish
brown-coloured pi gm ent diffu sin g into th e surrou nd ing agar; colony rever se
(colour piate 140 a) in pale cin namo n to reddish brown shades; peni cil1i (fig.
140 , A) typi cal1y bivertic il1ate asymm etrical , cons isti ng of ve rticils of ram i a nd
metulae or verticils of m etulae alone, m or e or less appressed; con id iopho res
mostly a rising fro m th e bas al felt in a den se and eve n stand, com mo nly sho rt
up to 100-1 50 ,tm in len gth by 3- 4 ,tm in diameter , septa te, wit h wa l1s
conspicuo us ly a nd fine1 y spi n ulose , with apices enla rge d in a vesicl e up to
4.5 - 6 .0 ,tm in di ameter; rami wh en present in number of on e or two eleme nts
in th e vert icil, u su al1y 18-25 ,tm by 3 .5 tui , cons picuo us ly spinulose;metul ae

lormiu g who rls 01' 2- :i clc mc n ts in lhe verticil , lIIos11 y 11 -1 511-111 by :i.O - 1'.5
unv , wi th pi ces c n la rgc d in a vesiclc up l o :i.5 -5.0 tit u in di amet cr , wi th wall s
lind y spinulosc ; phi alid cs ve ry regul ar in sizc, bottl e sha pe d, com mo nly 8- 10
Itlll by 3. 5 - 4.0 1I-m, tapcring grad ua lly to a n inconspi cu ou s, com paratively
widc neck ; co n id ia globose to subglobo se, smooth walled or nearl y so whe n
observed under th e light mi cr oscop e (fig . 140, B), bu t delicately gra n ula r when
observed under the SE M (pl ate 24 , top: righ t), m ostly 4-5 11-m by 3 .5 - 4.5 11-m
in di ameter; co nid ial chains te ndi ng to adhe re in to loos ely parallel divergen t
colu m ns (fig . 140 , C).
Colonies upon C YA (colour plat e 140 , B a nd b) atta in ing a d iameter 01' 45
mm in 14 days at roo m temperature (20 -25 C), consis ting 01' a close r textured
basal felt, ve lve ty, a ppearin g finel y granu la r, stro ngly and deepl y sulca ted by
nu rnerou s radial fu rro ws and slightly raised towards th e ce nt re; with sorne
tloccose tufts 01' white ae rial h yph ae in th e colony cent re; with a th in wh ite
rna rgin su rrou nde d by a fringe 01' su bme rge d hyphae exte nd ing abo u t 1 m m
beyond th e ae rial growth; heavily spo ring th rou gh out in di fferent shades 01'
brown- gr een to deep gree n (Sguy, PIs XXI , 3 14, and X X VI , 386) with the
ripening 01' co nid ia ; odo u r suggesting Actinomycetes; exudate limited in
amount, enmeshed in th e m ycelial felt; pale amber-coloured pi gm ent d iffusing
into th e surrounding aga r; colony reverse (colour plate 140 , b) in reddish
cinnamon shades; peni cilli esse nt ially as described aboye.
Colonies upon M EA (co lou r plate 140 , C and c) attaining a di ameter 01'
abo ut 50 mm in 14 days at room temperature (20- 25 C), consisting 01' a loosetext ured basal fel t , appearing velvety to delicately gra n ula r, flat, thin , with
ma rgin ar achnoid largely submerged and exte nd ing abou t 3 mm beyond the
aerial growth, wit h small sectors 01' floccose wh ite ae rial hyphae u su ally
ove rlaying th e colony centre ; heavily spo ring throu gh out in yellow -green
sha des (Sg uy, PI. XXVI , 386); odo u r lacking; ex udate limited in a mo u nt, as
m in ute colourless d roplet s em be dde d in m ycelial felt ; no co loured pigm ent
d iffus ing into the surrounding agar; colony reverse (colou r plat e 140 , c) at first
colourless, th en blackening in age; peni cilli esse n tially as described ab oye but
with conid ial chains tendin g to produce m ore com pact co lumns,
T he spec ies description is ce nte red up on th e typ e strain CBS 500. 73 = VKM
F-1 ,751 = ATCC 24,72 7 = IJ FM 7130 , isolated from Pirus malus L. fruit, in 1964
by 1'.1. N ovobranova , Institu te 01' Pomology and Ampelogr aphy , SS R 01'
Kazachstan , in the Alma- At a region, U SSR .
Penicillium mali N ovobranova is a dis tinctive spec ies whi ch , in ou r opinion
appears to be m ost satisfac torily ass igna ble to the P. roqueforti series on accou nt
01' its cultu ra l characterist ics a nd detail s 01' m orphology .

141. Penicillium f arinosum N ovob ra n ova: fig . 141 , colo ur pl ate 141 , a n d

SEM pl ate 24 , (centre: left) .

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Fig. 141. Camera lucida
drawings of Penicillium
farinosum Novobranova ,
CBS 499.73. A, Detailed
drawings of the penicilli . B,
Mature conidia . C, Habit
sketches .

Penicillium farinosum Novobranova - Nov. Syst. Niz. Rast. 11: 232-233. 1974.
Novobranova's original description follows :
'Colonies upon Czapek's agar attaining a diameter of about 30-40 mm in 10-12
days , consisting of a loose -textured felt appea ring pruinose or flour Iike at the
colony surface, which is gray-green coloured. Colony reverse uncoloured . Exudate
lacking. Odour very pronounced , 'mouldy'. Penicilli typically asymmetric, with
conidial chains tangled . Conidiophores coarsely roughened , 200-300 p.m by 4.5-5.6
(6.0) p.m, rami roughened , 12.0-18.0 by 3.5-4.0 p. m, metulae in whorls of 3-4
elements in the verticil , 10-12 p'm by 3.2-3.5 p. m, phialides 6.0 -8.0 p'm by 2.2 -2.5
p. m . Conidia smooth walled , globose to subglobose, 3.5-4.0 p.m in diameter. It
belongs to the Asymmetrica section . Related species unknown.'
(Translated from the Russian by Prof . L. Vilas) ,

Colonies u pon CA (co lour plate 141 , A and a) growing rather restrictedly,
attaining a diamet er of about 25 mm in 14 days at room temperature , velvety
in appearance, almost flat , not furrowed, or sligh tly fold ed in th e cen tre,

cons isling 0 1' a dense busal Iclt , with whitc (hin murgin surro u ndcd by a ri ugc
0 1' sub me rged hyph ac extc nd ing up lo 1:> II1Ill bcy ond thc acri al gro wth ; colony
surlace ovcrla id in places by a loosc1y WOVl:n netw ork of Iloccose white aerial
hyphae , lending lo th c colony a lan at c appcara ncc ; heavily spo ring throu gh out
in dccp grcc n shadc s (Sguy, PI. X X XI, 46 1); odour pen ctra ting , ' mo uldy',
exud a re limited in amo u nt, as minut e dropl ets enrneshed in th e m ycelial felt;
110 coloured pigm ent d iffus ing int o th e su rround ing agar; colony reve rse
(co lour plat e 141 , a) in pal e cream sha des ; peni cilli typically asym me tric (fig.
141 , A), co ns isting mostly of verticils of 3- 6 m etulae and less fre que ntly of
hranc hcs and metul ae , with all ele ments exce pt the phialides coa rse ly tuberenla te 0 1' wa rty; conidio pho res lon g , up to 300 /l m 0 1' m or e in length by
'1.0 - 5.5 /lm in d iameter, slightly enlarged at apex up to 4.5 -6.5 /lm in di ameter; ra mi 15- 25 /lm by 4 .0 /lm ; m etulae more 01' less cylindrica l, 13- 23 /lm
by 3.5 -4 .5 /lm , in clusters of 3-6 01' m ore ele me nts in th e verticil, rath er
d osely appressed; phialides bottle shape d, taperin g in a m or e 0 1' less lon g and
na rr owed neck , commo nly 9- 13 /lm by 4. 0 -4 .8 /lm ; con id ia globose to su b globose, 3 .5 -5 .0 uit: in long ax is , with walls smooth wh en obse rve d under th e
light micr oscop e (fig. 141, B) and very finely gra n ula r wh en obse rve d under
the SE M (pla te 24, centre : left) ; con id ial chains tending to adhe re into long
loosely parallel colu m ns (fig. 141, C) up to 300 utt: in len gth .
Colonies upon C YA gro wing m or e rapidly and broadl y spread ing (colou r
plate 141 , B and b) a tta ining a di ameter of 60 mm in 14 days at room
tem perature (20 - 25 C) , cons isti ng of a loose-textured basal felt, some what
fur ro wed an d rai sed , appearing velvety , with m argin thi n and irregul arl y
di ssected , largely subme rge d , ex te ndi ng abou t 3 mm beyond the ae rial gro wth,
app earing some what arach no id; very heavily sporing throu gh out in blue-green
sha des (S guy , PI. XXXI , 46 1); exudate abu ndantly produced as crys ta l clear ,
un coloured drops ; odo ur stro n g, suggestive of the Beta vulgaris L. aroma; no
coloured pigment diffusin g into th e su rrou nd ing ag ar ; colony reverse (colour
plat e 141 , b) developi ng reddish bro wn shades wit h a green cast sho wing
thro ugh in m argin ; peni cilli essentially as descri bed aboye .
Colonies upon M EA (colour pIate 141 , C a nd e), spre ad ing broadl y ,
attaining a di ameter of 55 mm in 14 days at room temperature (20- 25 C),
consisting of a loose-t extured basal felt, typi call y velvet y , some what powdery
0 1' pr uinose, fla t , not fur rowed, with margin arach no id, largely sub me rged,
exte nd ing abo u t 3 mm beyond th e aerial gr ow th ; heavily spo ring throu ghout in
gree n sha des (Sguy, PI. XXVIII, 417); odo ur penetrating, aromati c; exudate
lacking; no coloured pigm ent di ffusin g into th e surro und ing agar; colony
reverse (colour pIat e 141 , e) in yellowish shade s with a gree n ca st .showing
th rou gh ; penicilli essentially as describe d aboye , with con id iopho res eve n
lon ger, m ostly arising primarily from th e su bs tratu m, and con idial chains
adhe ring in m ore com pact colu m ns .

"."1'

'I'hc spec ics dcscripti on is cc nrcrcd llpOIl rhc t y p c srrn in GIlS 199. 7 :~ - VI< M
F-1 ,746 = CMl 174 ,717 = ATCC 24 , ni = I.lFM 7 1:11, isolat cd from (he su rfac
0 1' 'Schasselas' u nd er sto rage, in Octo bc r 1967 , by T .I. Novobran ova ,
lnstitu te 0 1' Pomology a nd Ampelography , al' the SSR of Kazach st an , USSR ,
in th e Alma -Ata region . lt is kn own only fro m th e typ e st ra in.
Penicillium fari nosum N ovobranova , on accou n t 0 1' its cult u ra l charactc ristics
and details al' morp ho logy is beli eved to be m ost sa tisfac torily assign able to the
P. roqueforti series.
Penicillium brevi-compactum se r ies
O UT ST A NDI NG F E AT U R E S

Colon ies grow ing rather restrictedl y , typi call y co ns isti ng 0 1' a close-tex turcd
ba sal fel t, w ith su rface growt h velve ty to almost lanat e , more a l' less co ns picuo us ly fu r rowed, heavily sporing th rough out, in variou s shades 0 1' dull yellowgree n, gray-gree n, ave lla neous to brown ; colony reverse usu all y in dull yellow ,
gree nish gray, 0 1' b row nish shades .
Conidiop ho res variable in length , com paratively long whe n borne di rectly
from th e su bstra tum, 0 1' short wh en a rising as branches fro m ae ria l hyphae,
usu all y smooth wa lled 0 1' del icat ely ro ughe ne d .
Marg inal stolo ns typi call y p roduced in m ost me mbe rs 01' th e series whe n
cultivated upon m oist su bs trata 0 1' under very humid cond itions.
Penicilli asym metrical , ty pica lly ra mi fied , characterized part icula rly by th e
co m pa ra tive sho rt and very co m pact b ase , with ra m i a p pressed , m et ul ae and
p hialides numerou s a nd closely cro wded; not ramified in sorne stra ins,
cons isti ng a l' terminal clusters a l' metul ae bearing crowded phi alides .
Conid ia globose , elliptica l lo more 01' less pyri form , smooth 0 1' delicately
ro ug he ned , wi th co nid ial chains tending to ad he re in mo re a l' less loosely
parallel colu m n s a l' definitely ta ng led .
K ey to species
Page
1. Penicilli typically rebranched below the level of metulae, with the main
axes and branches terminating in verticils of metulae. Penicilli comparatively short, with all elements closely appressed.
P. brevi-compactum series . . . . 394
A. Penicilli typically showing one or more side branches below the level
of metulae.
1. Conidia smooth-walled or nearly so.
P. vo/gaense Beljakova et Milko ... . 405
2. Conidia conspicuously roughened.
a. Conidiophore walls conspicuously roughened .
P. brevi-compactum Dierckx ... . 396

b. Coni diophoro wall s smooth or noorly so .

1' . Conidia strongly echinulate; co nidiol areas developing dark
brown shades .
P. brunneo-stoloniferum Abe ex Ramrez . . . . 412
2'. Conidia delicately roughened ; conidial areas developing
ochre-yellow shades.

P. arenicola Chalabuda . . . . 410

H. Penicill i typically consisting of a single , crowded , terminal verticil of
5-8 metulae .
1. Conid iophore walls conspicuously and coarsely roughened , at least
at the base; conidia ellipt ical , conspicuously roughened in spirally
bands.
P. skrjabinii Shmotina et Golovleva . . .. 407
2. Conidiophore walls smooth or nearly so.
a. Colonies growing restrictedly upon MEA and CA ; con idia mostly
globose.
P. stoloniferum Thom ... . 400
b. Colon ies growing more rapidly upon MEA and CA; conidia ellip tical.
P. paxilli Bainier . . . . 403

The ser ies as under stood by R a pe r a nd Thom (1949) , was represented by

well-defined species , n amely: Penicillium breoi-compactum D ierckx , P.
stoloniferum Thom , and P. paxilli Bainier .
In th e current study, the series h as been broadened enou gh in concept to
accomodate the following four additional species which ca n n ot otherw ise be
sa tisfac torily placed; Penicillium oolgaense Beljakova et Milko , P. skrjabinii
Shmotina et Golov lev a, P. arenicola Chalabuda , a nd P. brunneo-stoloniferum Abe
ex R a m rez . A n ew variet y of P. breoi-compactum is also included he re: Penicillium
breoi-compactum Dierckx var . magnum var . n ov .
However, a close relationship between these additional forms a nd those
included in this se r ies by Raper and Thom (1949) is not presumed , and th ey
a re considered together primarily as a matter of conveni ence .
Members of this series are very commonly e ncou n te red in soil , and h av e
be en isolat ed freque ntly from d ecaying ve ge ta tion, nuts, fruits, paper sto cks ,
fleshy fun gi , etc .
T he series is characterized particularly by the comparatively short and very
com p act penicilli , typically consisting of com p act verticils of m etulae and
phialid es borne d irectly from th e stipe of th e con id iophore s a n d upon one or
more la te ral rami below th e level of th e met u lae , but with all eleme nts
remaining clo sely appressed a ga in st the m ain axis . In m ost members of the
series a considerable portion of th e penicilli a re not ra m ified , showing a single
1hr ee

terminal verti cil 01' lil e! II lac supporting a cluste r 01' crowdcd phi alidcs. In SOIll"
stra ins thi s t yp e 01' pcni cillu s occu rs almos t cxclus ivcly. C on id ia l cha ins a re
variable in lcn gth , a ppearing loosely parallel 0 1' div er gcnt , with liule or no
tenden cy to form colu m ns , becoming tanglcd in ag c (lig . 142, C ).
Vi ewed under high magnificati on , th e peni cilli 0 1' mo st strains may be
stro ngly sugges tive 01' Penicillium citrinum a nd relat ed spec ies, exce pt 1'01' thcir
com pactness a nd larger size; viewed under low magnifi cati on, they show
masses 0 1' tan gled 01' div ergent chains 01' con id ia rather th an well-defin ed
colu m ns .

142 . Penicillium breoi-compactum Dierckx: fig . 142 , colo u r pl ate 142 , a n d

SEM plate 24, (ce n t re: right) .

~L.

'i

(~~rl '1'\~{
~

'1

L...--..J

100 jirn

Fig. 142. Camera lucida

drawings of Penlcillium
brevi-compactum Dierckx,
CBS 210.28. A , Detailed
drawings of the penicilli. B,
Mature conidia. C, Habit
sketches .

Penicillium brevi-compectum Dierckx - Ann. Soc oSci. Bruxelles. 25: 88. 1901. See
also Biourge's Monographie, La Cellule. 33: 155-157. 1923; also Ch. Thom, 'The
Penicillia', pp. 295-296. 1930; also Raper and Thorn, ' A Manual of the Penicillia',
pp. 407-411.1949.

Synonymies
Peniciltiurn cressum Sopp - Monographie, pp. 147-148. 1912. See also Ch.
Thom, 'The Penicillia', pp . 297-298.1930; sea also Raper and Thom , 'A Manual of
the Penicillia.' p. 410. 1949.
Penicillium blelowiezense Zaleski - Bull, Acad. Polonaise Sci. Math. Nat. Sr. B,
pp . 450-451. 1927. See also Ch. Thom, 'The Penicillia', pp. 303-304. 1930; also
Raper and Thom , 'A Manual of the Penicillia', pp. 410-411 .1949.
Penici//ium hagemi Zaleski - Bull , Acad . Polonaise Sci. Math. Nat. Sr. B, pp.
448-450. 1927. See also Ch. Thom , 'The Penicillia' , pp . 298-299. 1930; also Raper
and Thom , 'A Manual of the Penicillia', p. 411.1949.
Penicil/ium pa tris-meiZaleski - Bull, Acad. Polonaise Sci. Math. Nat. Sr. B, pp.
496-498. 1927. See also Ch. Thom, 'The Penicillia', p. 303. 1930; also Raper and
Thom, 'A Manual of the Penicillia', p. 411 . 1949.
Penici//ium szaferi Zaleski - Bull . Acad. Polonaise Sci. Math . Nat. Sr. B, pp .
447-448. 1927. See also Ch. Thom, 'The Penicillia', pp. 299-300. 1930; also Raper
and Thom , 'A Manua l of the Penicillia', p. 411 . 1949.

Colonies upon CA (eolour plate 142, A and a) growing very restrietedl y ,

a tta ining a diameter of 15 mm in 14 days at room temperature (20-25 C) ,
consisting of a close-textured basal felt , velv ety to almost lan ate , depending
upon the strain, slightly raised in the centre, with thin margin surrounded by a
fringe of submerged hyphae extending 1- 2 mm beyond the aerial growth,
za nate or azonate; aerial stolon- like hyphae oeeasionally observed,
com pa ratively shor t and stout; heavily sporing throughout in blue-green
shades (Sguy, PI. XXX, 442) ; with surfaee sometimes radiately furrowed ;
exud ate limited, as minute eolou rle ss droplets enmeshed in th e myeelial felt, or
laeking; odour faint, indefinite, or laeking ; no eo loured pi gm ent diffusing into
th e surrounding ag ar; eolony reverse (eo lour plate '142, a) in pale pinkish
eream shades ; penieilli (fig. 142, A) asym metr ieal , eompaet, often eom plexly
ram ified , with all elements commonly swo llen or in flat ed, with eoarse , ereet
eonidiophores wit h walls smooth or definitely roughened; eonidial struetures
very abundant , borne primarily from the basal felt at the agar surfaee in a close
stand ; eonidiophores ereet, usually straight and appearing m ore or less rigid ,
mostly 300 - 500 fJ-m in length by 4-5 fJ-m in diameter, with walls either smooth
or definitely roughen ed (fig . 142, A) ; penieilli typieally eompaet, ir regularly
ramified with 1, 2, or more rami closcly appressed against th e main axis,
bearing erowde d clu sters of m etulae , eaeh terminating in a eom pae t whorl of
phi alid es; rami mostl y 20-30 tut: by 4.0- 5.5 fJ- m ; m etulae eom monly wedge- ./
sh ap ed , 12-1 5 fJ-m b y 4.0 -5 .0 uii: in a m edian pl ane , with a piees enlarge d up
to 5-6 fJ-m in diameter, smooth or definitcly rough wa lled ; phi alides bottle
sh ap ed, 8-1 3 fJ-m b y 3.5 - 5.0 pst , abruptl y tap ering to a n ineonsp ieuou s neek ;
eonid ia globose to subglobose , 3.0 -4.5 fJ-m by 3.0 -3 .5 fJ-m in diamet er ,

dclicat cly rou ghcn cd (Iig , 142 , B, aIHI SEM plnrc H, ce n tre: right); conid ial
chai ns becollling ta uglcd in age (Iig . 11-2 , C).
Colo n ics upon C YA (co lou r plat e 142 , B a nd b) all ainin g a diamc tcr 01' 15
m m in 14 d a ys a t roo m ternpcrature (20 - 25C), co ns ist ing 01' a close- tcx tu rcd
a nd tou gh basal felt , strongly radia tely urrowed a nd irregul arly rai sed , wi th
ab ru pt m a rgin sligh tly su b me rged, ex te nd ing less than 1 mm beyond th e aerial
growth; spori ng less ab u ndan tly than a boye in th e sa me sh ades; odou r lacking;
exudate lim ited 01' lacking; ye llow ish coloured pi gment diffusing in to th c
su rrou nd ing agar; colony reverse (colour piate 142 , b ) in reddish cin namo n
shades, strongly pulling th e agar a way fro rn th e cu lture dish a n d cracking it ;
penicilli esse ntia lly as describ ed aboye.
Colon ies upon M EA (co lour plat e 142 , C a n d c) a ttain ing a di amet er 01'
about 20 m m in 14 days at room temperature (20 -2 5C), con sisting 01' a
velvety felt, not furrowed, pla ne, with m a rgin la r gely su b merged, exte n di ng
abo u t 4 m m b eyon d th e aerial growth; ve ry slight ly rai sed in colony ce n tre ;
hea vily spo ring in the sa me shades as aboye; m argin m ore 0 1' less irregul arly
crene llate in old cu ltu res from the irregul a r develop me n t al' stolo nlike hyphae;
odo u r a n d ex udate lack in g ; no coloured pi gment diffu sing into the
su rrou nd ing agar; colony reve rse (co lour pia te 142 , c) in pale ye llow ish shades;
peni cilli esse n tially as described a boye but with the co n id iopho re walls more
definitely ro ughened (fig. 142, A).
T he species d escri ption is centere d upon strain CBS 2 10.28 = CMI92,266 =
IJFM 72 15, isolated by K. Zaleski in 1926 as the typ e strai n 01' h is Penicillium
patris-m ei, from soil under con ifers in Poland , a nd upon m any othe r strains
isolated b y DI' A.T. M a rtnez from a n andosol in th e Provin ce 01' N a varra ,
Sp ain , under Fagus silvatica L. (Colour pl a tes 24 1 a nd 242 presen t two
re p resen tati ve strains 01' P. brevi-compactum, isola ted fro m soil by A .T.
M art nez , 1977 .)
Penicillium brevi-compactum D erckx val'. magn um val' n o v . : fig. 223 (see
p . 616), colo u r plate 22 3, a n d SEM pl ate 24, (bottom : left).
Latin diagnosis of Penicillium brevi-compactum Dierckx varo magnum varo nov.
Coloniae in agaro Czapekii, post duas hebdomadas ad 20-25C, 35 mm in diametro attingunt, velutinae, laxo texto basali coacto mycelio, planae, cum margine
inaequaliter plerumque e hyphis (2 mm) submersis formatur, planae ad marginem,
umbonatae ad centrum; fortiter sporulantes, cum fructificationibus conidialibus
coeruleo-viridibus; exsudatio parco; odor abest; sine pigmento in medio diffluente;
reverso sine colore. Penicillus varietatis a Penicillium brevi-compactum Dierckx
differt in latitudine conidiophorum, ramorum et metularum; conidiophora laevia,
tenus 7.5-8.0 p,m in diametro; rami laevia, tenus 7 p,m in diametro; metulae laevia,
tenus 6.5 p,m in diametro; conidia globosa vel subglobosa, 3-4 p,m in diametro,
asperula , in intricatis catenis formantur.

Coloniae in agaro Czapekii cum ext racto lovodlnl s, po st duas hebdomadas ad

O-25 C, 55 mm in diametro attingunt, velutino-granulosae, zonatae, cum radiatlbus sulcis, cum albo margine e submersis hyphis (2 mm) formatur; fructificationibus conidialibus abundantibus coeruleo-viridibus; odor et exsudatio absunt;
pigmento succineo diffluente in medio; reverso pallido flavo.
Coloniae in agaro hordeaceo, post duas hebdomadas ad 20-25C, 30 mm in diametro attingunt, velutinae , zonatae , planae, cum margine inaequalit er plerumque e
hvphis submersis (2 mm) formatur; fortiter sporulantes, cum fructificationibus
onidialibus coeruleo-viridibus; exsudatio et odor absunt; sine pigmento diffluente
in medio; reverso aurantio pallido.
Varietas a Penicillium brevi-compactum Dierckx differt plerumque maioru
coloniarum diametro atque in latidudine et magn itude penicilliorum.
Typus cultura IJFM 5954 ex aere Matriti Hispaniorum in anno 1979 isolatus est.

Colonies upon Czapek's agar (eolour plate 223 , A and a) attaining a di ameter of 35 mm in 14 days at room temperatu re (20 -25 C), eonsisting of a
loose- tcxt u red basal felt , d eep ly velvety to su bla na te , not fur rowed , with thin
white margin su rrou nd ed by a fri nge of submerged hyph ae extending a bou t 2
111 111 beyond the ae ria l growth, wi th eolony su rface pla ne in marg inal and su bIlIargin al a reas, a nd abruptIy raised as a plateau in th e cen tre, overla id by a
loosely wov en ne twork of white floccose hyph ae len ding a lanate a p peara nce to
I he central a rea; heavily sporing th rou gh ou t in deep blue-g recn shades (Sg uy ,
1'1. XXXI , 46 1); wit h the blue predom inant ; exudate lim ited in a mo u nt,
rnmeshed in th e mycelial felt ; odour lacki ng ; no coloured pigme nt diffus ing
1I10 th e surrou nding ag a r ; eolony rev ers e (co lou r plat e 223 , a) eolourless;
I)('nicilli (fig . 223 , A) typ ically of the Penicill ium breoi-compactum type bu t
ons pic uously larger and sto uter, persistentI y smooth wa lIed, wit h conid iophores ereet , arising from the m ycelial basal felt a t the agar surface in a close
and even stand , usualI y straight a nd rigid, un scp ta tc , rangin g fro m 400 to 500
t m or more in lcngth by 7.5 - 8 .0 ;,m in diamcter, with a pices eom monly
swollcn u p to 10 ;,m in d iamete r; pen icilli usu all y consisti ng of cro wde d
vcrticils of closely a ppressed sto ut ram i, mostIy 10-1 8 tui: b y 5 -7 u u,
numbe ring u p to 10 elements or m ore in thc vc rticil, vert icils bearin g stou t
mctulae, com m only wedge shaped , measu ring 4 .5- 7.0 ;,m in dia m eter in a
med ian plane b y 9 -12 ;,m in length , enlarged a t the ap cx up to 7.5 ;,m in
d ia me ter; phialides bottIe shaped, mostIy in who rls of 3 -6 elc m ents in the
vcrtieil, us ually 9 - 14 ;,m by 4 .5 -5 .5 ;,m ; co nidia globose to subglobose,
3. 0 - 4 .0 ;,m in d ia me te r , wit h wa lls delicately ro ughened (fig . 223 , 8 , a nd
SEM plat e 24 , bottom : left); con id ial cha ins usu all y ta ng led (fig. 223, C) .
Colo nies u po n CYA sp read ing broadl y (colour plate 223 , B) , attai n ing a
d ia meter of 55 mm in 14 d ays at roo m tem peraturc (20- 25 C), consisting of a
deeply velvety felt, co nsp icuo us ly za nate a nd radia tely wrin kled, slightIy

ra iscd, with white thin m a rgi n surrou ndc d by a rin gc 01' su b me rged hyph ae
exte nd ing a bo u t 2 mm bcyond th e ac rial gro w th; co lo ny surf ace ovc rla id by a
loosely woven n etwork of'white Iloccose ae r ial hyph ae lendin g a co ttony a ppc arance to th e co lony; heavil y sporing throu gh out in th e sa me sh ades as aboy e;
ex u date limited in a mo u n t, as minute co lou rlc ss drops; odour lacking ; pale
a m be r colou red pi gment diffu sing in to th e su rrou n d ing agar ; colo ny reverso
(colour pIa te 223, b ) in pale orange sh ades; penicilli esse n tial1y as de scr ibcd
aboye.
C olonies upon MEA (co lour pIate 223, C a n d e) a tta in ing a di ameter 01' 30
mm in 14 d ays a t room temperature (20-25C), co ns isting of a deepl y velvety
elt, appear ing granu lar a nd co ns picuous ly zo nate, with irregul arly di ssectcd
m argin developing sto lon- Iike su bmerge d hyphae exte nd ing a bo u t 2 mm
beyond th e aeria l growth; hea vily spo ring throu gh out in th e sa me shades 01'
blue-green as a boye; colony su rface not fu r rowed a nd Ilat ; ex u date a nd od ou r
lacking; no coloured pigment diffusin g into the su rrou n d ing agar; colony
rever se (colou r pla te 223, c) in pale oran ge shades; pen icil1i esse ntia lly as
descri bed aboye .
The va riety description is ce n te re d upon th e typ e strain IJFM 59 54, isolated
by th e au thor as a cu ltu re co n ta m inan t, M adrid , Spain, May , 197 9 . It is
kn own only frorn th e typ e strain.
Penicillium breoi-compactum Di erckx va r. magnum d iffers m arkedl y fro rn the
ty pe species, particul arl y by its b ro adl y sp re ad ing colo n ies upon al1 subs tra ta
a nd by its sto u te r a n d larger penicil1i, but othe rw ise, it ap p rox imates th e ty pc
spec ies we l1 enou gh in th e characte r istics a ttr ib u te d b y R ap er a n d Thom to th e
spec ies to warrant recogn ition as a va riety a n d not as a sepa rate species in th e
P. breoi-compactum ser ies . Therefore , it is d escribed a nd p rop osed as a n ew
variety wi th th e spec ific n ame of Penicillium breoi-comp actum D ierckx va r.
magnu m . T he va r ieta l ep ithe t refers to th e broadl y spread ing colo n ies,
particularl y upon G YA (co lour plat e 223, B) a n d th e sto u t penicil1i (fig . 223,

A).
143 . Penicillium stoloniferum T ho m: fig. 143 , colour plate 143 , a nd SEM
plat e 24 , (b ottom : ri ght).
Penicil/ium stoloniferum Thom - Bull, Bur. Anim. Ind. U.S. Dep. Agric. 118: 68-69.
1910. Also Ch. Thom, 'The Penicillia', pp. 292-294. 1930; also Raper and Thom,
'A Manual of the Penicillia.' pp. 412-414. 1949.

Synonymies
Penicillium biourgeianum Zaleski - Bul!. Acad. Polonaise Sci. Math. Nat . Sr. B,
pp. 462-464. 1927. See also Ch. Thom, 'The Penicillia', pp. 296-297. 1930; also
Raper and Thorn, 'A Manual of the Penicillia' , p. 414. 1949.

0 0
0 80 O
O

ca 00 0 0

O 0 00 O
O 00
00
O

00

,~~

j~~~( ~" ;;~~\'

1~~~'\

~r~'
0:,'4

1~1 ~
100

jirn

fR .

Fig. 143. Camera lucida

drawings of Penicillium
stoloniferum Thom, CBS
227.28. A, Detailed drawing s
of the penicilli. B, Mature
conidia. C, Habit sketches.

Penicillium erectum Bainier - Bull. Soc o Mycol. France. 23: 13. 1907. See also
Ch. Tho m, 'The Penicillia' , p. 295. 1930; also Raper and Thom , 'A Manual of the
Penicillia', p. 413. 1949.
Penicillium griseo-brunneum Dierckx - Ann . Soc o Sci. Bruxelles. 25: 88. 1901.
See also Biourge's.Monographie, La Cellule. 33: 162-163. 1923; also Ch. Thom ,
'The Penicillia', p. 302. 1930; also Raper and Thom, 'A Manual of the Penicillia' ,
p. 413. 1949.
Penicillium tabascens Westling - Arkiv . Bol. 11 : 56, 100-102. 1911. See also Ch.
Thorn, 'The Penicillia' , pp . 300-301. 1930; also Raper and Thom , 'A Manual of the
Penicillia', pp . 413-414. 1949.

Colonies u pon CA (colour plate 143, A a nd a) growing moderately,

attaining a di ameter of 35 mm in 14 days at room tem perature (2-25 C),
typically consisting of a close -textured basal felt, with su rface deeply velvety to
lanate in most strains, but more or less floccose in others, obscuring in th ese
cases th e otherwise abundantly produced conidial structures (colour plate 143,
A) , with cen tral areas conspicuously raised ; rad iately sulcated by shallow
furrows; heavily sporing throughout in yellow-green shades (Sguy, PI.

XXIX, 12G), hu! lI\oslly ob scu red hy th c lloccos ovcrgrowth 01' whit e acrinl
hyphac; thin and Ila! m argin su r rou ndc d by a narrow fringe 01' submcrgcd
hyph ac ext ending less than 1 mm beyond th e ac ria l growth ; cxudate ami
odou r lackin g ; pale yellow pigment slowly diffu sing into the surrounding agar;
colony rev erse (col our pi ate 143, a) in pale cre a m sh ad es; penicilli (lig. 143, A)
typically co m p ac t with all elements clos ely a p p re ssed, forming loosely p arall cl
or divergent ch ains 01' con id ia , presenting th e pattern 01' short , co m pact
brushes, with con id iopho res a r ising eithe r from submerged hyphae at th e agar
su rface, or a s lateral ramifications from aerial h yphae , the forrn er rangin g from
100 to 350 flm in length by 3. 0 - 3 .5 flm in di ameter, the latter from 35 lO 100
flm by 3 .0 - 3 .5 tuts , smooth wall ed; larger penicilli co ns isting usu all y a l'
ve rticils 01' metulae a nd sometimes on e , or occasion ally more , sid e rami ea ch
terminated in a whorl a l' m etulae; smalle r penicilli com mo nly co nsisting 01' a
sin gle terminal ve rticil 01' closely a pp res sed m etulae each bearing cro wde d
phialidcs ; rami , when present usuall y 10-25 flm by 3. 0 -3 .5 flm ; m etulac ,
ran ging from 3 to 8 in number in th e verticil, mostly 11-1 5 iut: b y 2 .8-5 .0 flm ,
with a p ices e nla rged up to 4- 6 flm in di amet er; phialides in co m pac t cluste rs
4- 8 elemen ts in th e verticil , b ottl e sh aped , mostly 7-11 flm b y 3- 4 flm,
taper ing to a sho rt n arrowed neck ; con id ia elliptica l wh en you ng , but
becoming globose to subglobose a t m aturity , mostly 2.5-3 .5 flm in di amet er,
appear ing smooth walled 0 1' nearly so when obs erve d under th e light
mi cr oscope (fig. 143 , B) but fin ely roughened when obse rve d under th e SEM
(pl at e 24 , bottom : right); co n id ial cha ins forming loosel y parallcl or divergent
colu m ns (fig. 143, C) .
Colon ies upon CYA (col our pl at e 143 , B and b) sp re ad ing broadly , a tta in ing
a di ameter a l' 50 mm in 14 days at room temperature (20- 25 C ), co ns isting 01'
a close -textu re d ba sal felt , deepl y velve ty a nd stron gly sulcated by numerous
radial furrows, grad ually raised tow ard s th e colony ce n tre, with e n tire m argin
white ; heavily spor ing th roughout in blue- gr een shades (Sguy, PI. XXX,
443) ; conspicuously zonate; odo ur faint, indefinite ; ex u date abundantly
produced , as crys tal clear , p al e a m bc r-colou rc d drops ; a m be r-colou red
pi gment d iffu sing into th e surro u nd ing aga r ; colony reverse (co lour plat e 143,
b) in sa ndy to cinnamon shades, a nd strongly pulling a n d splitting th e
underlayin g agar ; penicilli esse n tially as described a boye .
C olon ies upon MEA (colour plate 143 , C a nd c) growing restrictcdl y,
att aining a di amet er 01' ab ou t 25 mm in 14 d ays a t ro om temperature
(20- 25C), con sisting 01' a dense velvety felt , slightly radiat ely furro wed in th e
ce ntre, alm os t flat , with irregularly di ssected ye llow m a rgin largel y su b me rged
ex te nd ing a bo u t 4 mm beyond th e ae
r ia l growth, co ns isting 01' stolon-like
/
hyphae co m para tively stout ; h ea vily spo ring throu ghout in gray-g reen shad es
(S guy, PI. XXIX , 428); exu date a n d odo u r lacking ; no coloured pi gment
diffusing into the surrounding ag a r; colony rev erse (co lou r piat e 143, e) in

IJI'lI ll ge sh ad cs: pcni cilli cssc n tia lly as dcscribcd ab oye .

' I'he s pc cies dcscripti on is cc n ic rcd Up OIl th c t yp e srrn in AT C C 10,111 - C IlS

= C M I :~9 ,B2 4 = NRIU. B59 - )F D 5, 1l :>1l - Q M 7, 653 , isola tc d by C h.
'1'111111 1 Irom mttin g mu shroorns in th c su n u ne r 01' 1904 a t S to rrs, C onnccti cut ,
JSA , a nd upon strain C BS 227 .2 B = I.JFM 71 24 , isol ated by K . Z al esk i (a s P.
hialounezense fr orn forest so il under co n ifers in Poland , 1926 .
~;J ( ;. : l l

I-H . Penicillium paxilli Bainier: fig. 144, ca la u r plat e 144 , and SEM plate
::!:i, (tap : left) .

W-!J..LJ.J..W

10J,lm

O O 00
B O

OQO

DO O

'i".""
:';;'~ t

. _,~.

-","

.,. ~ \~\,

'/I/~

,~,~,,_c;"". '. ;~i . e . l. .

~~.;~N
. I <:'5/'~
,'_

"1"

100J,lm

~J',;" . ii; " "

""""i\',

fR.

Fig. 144. Camera lucida

drawings of Penicillium
paxilli Bainier , CBS 360.48.
A, Detailed drawings of the
penicilli. B, Mature conidia.
C, Habit sketches.

Penicillium paxilli Bainier - Bul. Soc oMycol. France. 23: 95-96. 1907. See also Ch.
Thom, 'The Penicillia' , pp. 294-296. 1930; also Raper and Thorn, 'A Manual of the
Penicillia.' pp. 414-416. 1949.

Colo n ies upon C A (co lo ur plate 144 , A a n d a) spread ing rather b ro a dl y,

attain ing 'a di amet er 01' abou t 50 mm in 14 d a ys at room temperature
(20 -25 C ), co ns isting 01' a close-textured b as al felt , deeply velve ty to lanat e ,

co ns p icuo us ly zo na tc, almos t planc 01' show in g lew sha llow ra d ial u rrow s:
colo ny su rlacc ovc rla id by a loosely WOVCII nctwork 01' whit e loccosc ac ria l
h yphae , either in localized sec to rs 0 1' in marginal arcas, rorn whi ch scco ndury
conidial structu res are dcvelop ed ; flat grow ing m argin white su rro u nclcd by /1
narrow frin ge 01' sub me rge d hyphae exte nd ing about 1 mm beyond th e aeriul
grow th; heavily spo r ing throu gh ou t , at first in avella neo us sha des (Sguy , 1'1.
XIII , 190) during th e first 2 week s, sha d ing th e blue- gr een (Sguy , PI. XXI X,
425) with th e late ripe ning 01' con id ia; exudate a nd od our lacking; pal e a mb crcoloured pi gm ent slowly di ffusing into th e surrou nd ing agar; colony reverso in
dull yellow to cinnam on sha des (colour plat e 144 , a); peni cilli typically
com pact (fig. 144 , A), consisting usually 01' a sing le termi nal ve rticil 01' 2- 11
metulae from which arise di vergent chains and loos e colu m n s 01' conidia (Ii g.
144 , C) ; con id iopho re s arising primarily from th e substratum in a den se anrl
eve n sta nd, occasionally a s short branches from aeri al hyphae , va riable in
len gth , com mo nly 200 -300 p.m by 2. 5 - 4 .0 p.m , so me times longer u p lo GOO
tuu ; with walls sm ooth 01' nearl y so; metulae 12-1 5 p.m by 3 .0- 4 .5 uit, wi th
apices enla rged to a vesicle up to 4- 6 p.m in d iamet er; phialides bottle sha pcd ,
in wh orl s 01' 3-9 eleme nts in th e verticil , mos tly 8-11 p.m by 3 .0-3. 5 .ull ,
tap ering abr u ptly to a sho rt narrowed neck ; conid ia mostl y ellipticnl,
com mo nly 3 .0 - 3 .5 p.m by 2.0 -3 .0 p.m in d iame te r, with walls smoo th (J I'
neari y so wh en obse rve d under th e light mi croscop e (fig. 144, B) but delical ely
rou ghened wh en obse rved under th e SEM (pIate 25, top : left ) ; con id ial chains
loosely parallel 0 1' divergent colu m ns (fig . 144 , C).
Colonies upon C YA (colour plat e 144 , B a nd b) spreadi ng broadly , atta ining
a di ameter 01' 50 mm in 14 days at roo m temperature (20-25 C), cons isting 01'
a sim ila riy textured ba sal felt as aboye, but m arked by numerous and dcep
radial fu rro ws, with ce ntre some wha t raised and irregul ari y fold ed , wi th a thin
white margin surro u nde d by a n arrow frin ge 01' su b me rge d hyphae extend ing
abo ut 1 mm beyond th e ae rial gro wth; heavily spo ring th rough out at first in
ave lla neo us shades (Sguy, PI. XIII , 190), th en becoming blue-gr een colou red
with the rip en in g 01' con id ia (Sguy, PI. XXXI, 463 ) whi ch sta rts in margin al
areas; odo u r lacking 0 1' faint ; ex udate lacking ; pale amber-pigment diffusin g
into th e su rround ing aga r; colony reverse (colou r plat e 144 , b) in orange to
cinnamo n shades ; peni cilli esse nt ially as descr ibed aboye .
Colonies upon M EA (colour plate 144 , C and c) spreadi ng broadl y,
atta in ing a di ameter 01' 50 mm in 14 day s at room temperature (20 - 25C) ,
consisting 01' a looser textured ba sal felt , conspicuously za nate , typi cally
velvety , with ve ry thin m argin su rrou nde d by a eringe 01' su bme rged hyphae
exte nd ing abo ut 2 m m beyond th e ae rial gro wth; colony su rface plane , not
furro wed ; heavily spo ring througho u t in brownish gree n shades (Sguy, PI.
XXI , 303) an d rem aining persistently so ; ex udate and odo ur lacking; no
coloured pigment di ffusin g in to th e su rrou nd ing agar; colon y reverse (colour

plute 144 , e) dcvclopin g ycllowish cinnamon shudcs with a grec nish cast
showing throu gh ; peni cilli essc n tia lly as descri bcd aboye .
'I'he species descri ption is cen tercd u pon thc type stra in c ss 360 .48 = C M I
'10 ,226 = AT C C 10,480 = NRR L 2,008 = QM 725 = IJ FM 7122 , isolat ed by Prof.
W. C. Hutchinson , Bar ro Colora do Island, Panam a , from optical in struments,
I !l17. Stra ins of thi s species are frequently isolat ed from deteri or ating mili ta ry
cq uipment , from fleshy fung i, nuts, etc .

14 5. Penicillium volgaense Baljakova el Milko: fig. 145, colour plat e 145,

an d SEM plat e 25, (top : ri ght).

oo a

000

0
0O O

OO sO

11

0
0 O
w..u.l=J
10

urn

9>!c r ~

_~

L...-.-...-J

100 prn

. ,Y

.l?.

Fig. 145. Camera lucida

drawings of Penicillium
volgaense Beljakova et
Milko, CSS 626.72. A,
Detailed drawings of the
penicilli. B, Mature conidia .
e, Habit sketches.

Penicillium volgaense Beljakova et Milko - Mikologia i Fitopathologia. 6: 146-149.

1972.
Beljakova and Milko's original diagnosis follows, since this adequately presents
the species in question :
'Colonies upon Czapek's agar attaining a diameter of 22-30 mm in 10-12 days,
velvety, radiately furrowed, with central tufts consisting of aerial interlacing

hyph ae, heavlly sporlng, at first blue-gray wlth a greenlsh cast , th en (after 45 days)
shading to olive gray to tobacco, and usually (after 3 months) becoming violet-gray
in colour; with even white margin, 1-1.5 mm wide; exudate Iimited, colourless;
colony reverse in pale lemon yellow shades. Conidiophores erect, not branched,
occasionally with a side branch 260-400 fLm by 4-4.6 fLm, with apices inflated up to
5 (7.S) fLm in diameter, delicately roughened, arising either from the substratum or
as branches from aerial hyphae or ropes of hyphae (particularly in the colony
centre). Penicilli typically cornpact, usually symmetrical, two- to three-stage
branched , and asymmetrical with a side branch under the level of the metulae .
Branches when present 9-11.7 fLm by 2.5-3.3 fLm, with apices enlarged in a vesicle
up to 3.6 fLm in diameter, usually numbering 4 in the verticil. Metulae 7.S-10.0 fLm
by 2.6-4.0 fLm, slightly enlarged at the apex, numbering up to 5 elements per
verticil. Phialides 7.S-9.0 fLm by 2.0 fLm, numbering 6-S per cluster. Conidia rough
walled, at first elliptical, 2.6-3.3 fLm by 2.0-2.6 fLm, then becoming globose,
2.6-3.6 fLm in diameter forming more or less divergent chains, blue-gray coloured in
mass.
Colonies upon malt extract agar appearing velvety to tanate, heavily sporing, at
first bluish gray, with margin white up to 2 mm wide, then becoming (after 45 days)
dark gray, with the colony reverse reddish coloured. Conidiophores arising either
from the substratum and from aerial hyphae or from ropes of hyphae . Otherwise as
described aboye. Odour faint, suggestive of dried fleshy fungi. Maximum
temperature of growth about 30C, minimum SOC, optimum 20-26C.'
(Translated from the Russian by Prof. L. Vilas).

O ur no tes follow:
Co lonies u pon CA (colour plat e 145, A and a) growing rather rest rictedly ,
attaining a d ia meter of 30 mm in 14 days at roo m tem perature (20-25 C) ,
consis ting of a close -tex tured b asal felt ove rla id by a dense network of white
floccose ae ria l hyphae len din g a deeply lanat e appearance to the colony
surface; slight ly raised in th e ce nt re , not fu rro wed, with a thin eve n m argin
surrou nde d by a narrow fringe of su bm er ged hyphae exte nd ing a bout 2 mm
beyo nd th e aerial grow th; medium sporing throughout in gray-gre en shade s
(Sguy, PI. XXVII , 404), but with conid ial areas almost com pletely obs cu red
by th e floccose ove rg row th of white ae rial hyphae ; exudate and odou r lacking;
pale yellow -coloured pigment di ffusin g into th e su rround ing agar; colony
reverse (co lour pIate 145, a) in pale yellow shade s; peni cilli (fig . 145, A) as
descr ibed aboye, suggestivc of th e Penicillium paxilli typ e , usu all y consisting of
small ve rticils of ram i, ranging from 20 to 25 I-tm in length by 2.0 -2.5 I-tm in
di am eter, be a ring wh orl s of 2- 3 metulae , mostly 9- 15 I-tm by 2 .5 -5 .0 tut,
ea ch su pp orting cro wded clus ters of 2- 8 phial ides u suall y 8 -1 OI-tm by 2.5 - 3 .8
tut , tape r ing to a short narrowed neck ; all ele me nts in th e penicillu s are m or e
or less closely appressed ; co nid iopho res smooth walled , com mo nly short , up to

I!jO Itlll in lcn grh hy 2 .:i -:~ . 0 Itlll in di nmctc r , ari sing' cithe r rom the
1I11i )slra tlllll 01' as short branch es Irom ae ria l hyph ac 01' from rop es 01' hyph ac ,
pur ticulurly in th e colony ce nt re; co nid ia cllipiical :1 .0 - 4 .5 utt: by 2.2-2. 8 tm
11) subg lobose , a ppear ing smoo th walled or nearly so wh en obse rve d under th e
ligll( micro scop e (Ii g. 145, B) but deli cately rou gh ened wh en obse rve d under
t lu: SE M (plate 25, top : ri ght) ; conid ial chains forming loosely paraUeI 01'
dcflnir ely ta ng led (fig . 145, C ).
Colonies upon C YA (colour pla te 145 , B a nd b), a ttain ing a diameter 01' 65
111111 in 14 days at room temperature (20- 25C), cons isting 01' a c1ose-textured
hasa l clt , a ppearing deepl y velvety to subla nate 0 1' funi cul ose , m arked by
1III III CroUS and deep radi al fur row s, slightly ra ised in the ce ntre, with thin white
Illa rgin surrou nded by a wide frin ge 01' su bmerge d hyphae exte nd ing about 4
111111 bcyond the ae rial growth ; heavily sporing throughout in brownish gray
sltadcs (Sguy, PI. X LVII , 694), with conid ial a reas loosely ove rla id by a
ru-twork 01' floccose and fu n iculose veget ative hyphae ; odo ur and exudate
la('kin g; pale yeUow-colou red pigment d iffusing in to the surrou nd ing agar;
ro lony reverse in cin namon shades (colour plat e 145, b); peni ciUi esse ntia Uy as
. k-sc rib ed abo ye.
C olonies u pon M EA (colour pia te 145, C and c) growing m or e res trictedly
Ihan up on C YA , atta ining a di amet er 01' 45 mm in 14 days at roo m
11'llIperatu re (20-25 C), co ns isting 01' a den se velvety felt , com pa ratively thin
.uid flat , not furrowed, with th in eve n m argin mostly su bme rge d , exte ndi ng
nbout 4 mm beyond the ae rial growth, colony su rface loosely ove rla id by a
uct work 01' ae rial ve geta tive hyph ae and rop es 01' h yphae ; heavily spo ring
throug ho ut in a rtemisia gree n shades (Sg uy, PI. XXIX , 429); exudate a nd
odo ur lacking ; no coloured pigment di ffusin g into th e surrou nd ing agar;
ro lony reverse (colour plat e 145 , c) developing yeUowish cin namo n sha des ;
pcni cilli esse nt ially as descri bed aboye.
T he species description is ce ntered upon the typ e strain CBS 626. 72 = VKM
11-1 ,127 = C M I 167 ,384 = IJ F M 7114 , isolat ed by L.A . Belj ak ov a and A .A .
Milko in 1966 fro m soil near Pless, U SSR. It is kn own only from th e type
st ra in .
Penicillium volgaense appears to be most sa tisfac torily ass ignable to th e P. breoirompactum series near P. paxilli Bainier on accou nt 01' the resemblance 01' its
peni ciUi with those 01' th e la tter spec ies. The presen ce 01' terminal ve rticils 01'
metulae in both form s is stro ng ly su ggestive 01' th e Penicillium citrinum series,
bu t they d iffer from it by its larger peni ciUi with aU elements m or e close ly
app resse d .

146 . Penicillium skrj abinii Sh moti na el Golov leva : fi g. 146 , colour plate
146 , a nd SE M plat e 25, (centre : Ieft ).

' H IlI

(J q:j)Q)
@>

0: (~~\,

0:

\:''\:"~ll ' "~:

"

(b(J
e

"

"

@
(J ,s O

O,

a
~

Qj

"
O

e
L--J

100~m

f./{.

Fig. 146. Camera lucida

drawings of Penicillium
skrjabinii Shmotina el
Golovleva, CSS 439.75. A,
Detailed drawings of the
penicilli. S, Mature conidia .
e, Habit sketches .

Penicilllurn skrjabinii Shmotina et Golovleva - Mikologia i Fitopatologia. 8: 530-532,

1974.
Shmotina and Golovleva 's original diagnosis follows, as this adequately presents
the species in question when recently isolated :
' Colonies upon Czapek's agar attaining a diameter of 38-46 mm in 12-14 days,
velvety, radiately furrowed, surface irregular, w ith centre folded, pale pink
coloured , w ith margin granular surrounded by a fringe of submerged hyphae
extending 1-2 mm beyond the aerial growth, somewhat zonate. in blue-green
shades , green , dull green , or chrysolite green colours; colony reverse at first cream
coloured, then shading to terracotta in age, with marginal and submarginal areas in
lemon yellow , sulphur yellow to whitish tints. Exudate colourless or cream
coloured , limited in amount or lacking . Odour pleasant (suggestive of the cheese
mould's aroma). Conid iophores erect, not ramified, with apices enlarged up to 4-5
Ilm in diameter, conspicuously warty, arising from the substratum or as branches
from aerial hyphae ranging from 25 to 500 Ilm, or up to 600 Ilm in length by 2.1 Ilm in
diameter. Penicilli cornpact, mostly biverticillate and asymmetric, occasionally

Imple lappearing monoverti eillate) ; bran ehes wh on present , in number of one per
punlcillus, elosely appressed against th e main axis or divergent, eonspieuously
w nrty, 26.4-32 .tm or up to 40 Ilm in length by 2.4-3 .tm up to 3.2 .tm in diameter;
rnot ulae parallel or somewhat divergent, with apiees inflated up to 4 .tm in diameter,
onspieuously warty, (3-4-6-8) 7.7 .tm- 16.1 .tm up to 20.3 .tm in length by
.1- 2.8, up to 3.0 .tm in diameter; phialides in erowded vert ieils, 6-8 elements in
lile whorl, bottle shaped, tapering to a narrow neek up to 2 .tm long , about 7.7 -10.5
IHl1 up to 19 .tm by 2.3-3.0 .tm, up to 3.5 Jlm; eonidia elliptieal, globose or
subqlobose, rough walled, 1.8-2.4 Jlm by 3.5-5.0 .tm in diameter, in ehains of
50- 100 Jlm long .
Colonies upon malt extraet agar attaining a diameter of 47-54 mm in 10 days,
wa nular, w ith centre velvety, slightly raised, irregularly folded in a radial pattern, in
loccose seetors narrowing towards the eolon y margino The eolony surfaee
coloured in dark green to lead gray shades, with seetors in lead gray t ints and
colony cent re and margin white ; colony reverse developing tobaceo shades,
radiat ely furrowed in the centre . Exudate Iimited in amount, as minute droplets
enmeshed in the mycelial felt. Odour pleasant , 'mouldv' .'
(Translated from the Russian by Prof. L. Vilas ).

Our notes follow :

Colon ies upon C A (colour plate 146 , A and a) attaining a d iame te r of 45 mm
in 14 days at room temperature (20- 25C), consisting of a loose-textured basal
k it, velve ty, conspicu ou sly zo nate, plane , not furrowed , overl aid by a loosely
wove n nctwork of deep ly floccose aerial wh ite hyphae, partially obsc u ring th e
otherwise hea vily spo ring ae ra s, len d ing a deep ly lanate appearance to the
colony su rface; very thin m argin surrounded by a wide frin ge of su b me rged
hyph ae exte nd ing abo ut 5 mm or m or e beyond th e aeri al gro wth; heavily
spo ring throughou t in well m arked zones in grayish gree n to brow nish shades
(Sguy, Pis XXVII', 404, and XXIX, 427); exudate and odo ur lacking ; pale
yellow-co loured pigm ent d iffusin g into th e surround ing agar; colony rev er se
(colour plat e 146 , a) colou rless to very pale cream shade s; pen icilli essentially
as described aboye, in ou r obse rvation only conid iopho re s were con spicuo usly
wa rty , particu la rly at the base, branches a nd m etulae were found always
smooth walled ; conidiop hores usu ally long wh en a rising from the su bstratum,
up to 700 tun in length by 3.0-4. 0 J.m in di amet er ; rami occas ionally obs erved ;
peni cilli com m only cons isting of verticils of 2- 3 more or less appressed
met ulae at th e tips of th e stipes about 14 - 20 J.m by 3 - 4 tui , with apices
enla rged in a vesicle up to 3 -4 utt , some time s up to 5 J.m in diameter;
phi alides in crowde d cluste rs of 2- 8 elem ents in th e vert icil, bottl e shaped,
10-1 5 J.m by 3-4.5 J.m , cons picuous ly tap eri ng to a lon g narrowed neck ;
conidia u sually elliptical, 3.2 - 4 .5 J.m by 2.5- 3.0 tui, cons pic uo usly
rou ghe ne d, wit h rou ghness arranged in a spirally pattern (fig. 146, B, and

SEM plat c 25, centre : lc ft); co nid iul chnins lormin g loosely para lid lo
divcrgent (Iig. 146, C ).
Colon ies upon C YA (colour plate 146 , 13 and b), aua in ing a di am ct cr of 40
mm in 14 days at room temperature (20-25 C), co n sistin g o!" a closc- rextu red
basal felt, d eep ly velv ety , somewhat zanate, strongly and dccply radiatc ly
furrowed, with centre irregu la rly fold ed and sligh tly rai sed, with colon y surfacc
overlaid by a thin , closely wov en n etwork of fine white aerial hyp hae influcn cin g gr eatly the colony appearance; margin entire su rro u nde d by a very
narrow fringe of submerged hyphae extending less than 4 mm beyond th e
aerial growth; sporing we ll but gr adually , starting at th e margin and thcn
proceeding towards th e centre in deep blue-green shades (Sguy, PI. XXXI,
461), with the ripening of conidia, elsewhere developing cin n a m on shades;
odour faint, ' m ouldy'; exudate abundantl y produced , as pale amber-coloured
drops ; a m be r-coloured pi gment diffusing into th e surrounding agar; colony
rev erse (colour plate 146 , b) developing reddish brown shades; penicilli essentiall y as des cribed ab oye .
Colonies upon MEA (co lour pl at e 146, C a n d c) attainin g a diamet er of 40
mm in 14 days a t room temperature (20 -25 C) , consisti ng of a loos e-textured
basal felt , deeply velv ety to sublanate, somewhat tufted by means of whitc,
floccos e, interlacing, ae r ial hyphae, with a very th in m argin surrounded by a
fringe of submerged hyphae exte n d in g ab ou t 3 mm beyond th e aeri al growth;
he av ily sporing throughout in deep gray-green shades (Sguy , PI. XXI, 308,
309); odour faint , aromatic; ex u date lackin g ; no coloured pigment diffusing
into th e surrounding agar; colon y rev erse (co lour plate 146 , c) dev elop in g
yellowish oran ge sh ad es with a green ca st showin g through ; penicilli as
described aboye but with longer stipes a rising primarily from the substratum.
The spe cies description is ce n te re d upon th e typ e strain CBS 439 .75 = VKM
F-1940 = IJFM 7123 , isolated in 1971 by L.A . Golovleva from soil , near the
village of Mujink, in the vicinity of Blagoveshczensk, in th e Amur region,
USSR.
Penicillium skrjabinii is a distincti ve sp ecies which appears mo st clo sely related
to the spe cies included within the P. brevi-compactum seri es , particu larly to P.
paxilli Bainier and P. stoloniferum Thom, but it can easily be differentiated from
both of th em by its conspicuous ly warty stipes and by its cu ltu ral ch a racteristics .

147 . Penicillium arenicola ChaIab uda: fig. 147, coIour pIate 14 7, and
SEM pIate 25, (centre: right) .
Penicil!ium arenicola Chalabuda - Nov. Syst . Crypt . Inst. Bot. Komarov ii, Acad.
Sci. USSR . 6: 162. 1950. See also M.M. Kulik, 'A Compilaton of Descriptions of
New Penicil!ium Species'. Agric. Handbook No. 351. Agre. Res. Ser. U.S. Dep.
Agr ie. p. 16. 1968.

0. 00

O
000

O BO~OC)

oOa O O
QO G

1111
L...-....J

100jJm

CK .

Fig. 147. Camera lucida

drawings of Penicillium
arenicola Chalabuda, CSS
220.66. A , Detailed drawings
of the penicilli. S, Mature
conid ia. C, Habit sketches .

Coloni es upon CA (colour plat e 147, A and a) growing restrictedly ,

altaining a diameter of abou t 15 mm in 14 days at room temperature
(20- 25C ), co nsisting of a tou gh, close-textured basal felt, with surface finel y
Iuni cu lose , ap pare nt ly velvet y , with thin , even m argin , white and en tire ,
surrou nde d by a narrow frin ge of submerged hyph ae extend ing lcss than 1 mm
beyond the ae rial gro wth, very slightly furrowed to flat; very poorly sporing,
devel oping pale och re-yellow shades; exudate and odo ur lacking; no colou red
pigm ent diffusing int o the surrounding agar; colony rev erse (colour piate 147 ,
a) in dark brown shades ; penicilli u su ally typ ically asym me trical (fig. 147 , A)
with a11 elements closcly appressed, con sistin g of verticils of ram i eac h
terminating in clus ter s of metulae at th e tip of conid iophores whi ch are
cons picuous ly septate and smooth walled , some times of branche s a nd m etulae
ar ising at different levels on th e conidiophores; con id iophores arising primarily
from th e substratum , abou t 50- 200 /lm by 2.5 - 5. 0 tut: with apices in flat ed up
to 4- 6 /lm in di ameter ; ra m i 10- 15 /lm by 3.5 -5. 0 psu ; m etulae m ore or less
cylind rical, 7-1 7 /lm by 3- 5 /lm ; phialidcs in crowde d clus ters of 2- 6 elem ents
in th c ve rticil, bottle sha pe d, 8 - 13 /lm by 2.5 - 4. 0 /l m , gr a?ua11y tap eri ng to a
short narrowed neck ; conid ia pyriform to elliptica l 4 -6 /lm by 2.5 -3.5 tui,

-r

d cli cat cl y rou gh cnccl (fig . 14-7 , B, a nd S E M pl at c 2;>, cc n t re : ri ght ); co n id ia l

chains sho r t, loosely parall el (Iig. 147 , C) .
C olon ies upon C YA (co lour plate 147 , H a nd b ) a tt a in ing a di am ercr of :i5
mm in 14 days at ro om temperature (20 - 25 C ), co nsisting of a fun icul osc
b asal felt a ppearing fin ely velvety, stro ngly a nd d eepl y radiat ely furrow cd ,
with ce n tre rais ed in a crate rifor rn pattern , fin ely zanate , with thin a nd evc n
marg in e n tire; spo ring ab u n dantly in ye llow ochre shades, but co n id ia l a rcas
some what a bsc u re d by a ve ry thin netw ork a l' closely wov en white ae r ial
h yphae ; ex u date lim ited in a mou nt , as minute colou rless d ropl et s e n meshed in
th e m yceli al felt ; odou r lacking ; da rk a m be r-colou re d pigment diffusing in to
the su rrou nding agar; co lony rever se (colour plat e 147 , b) de velopin g deep
reddish brown sh ades ; penicilli esse n tially as descri bed a boye .
C olonies upon MEA (calour pia te 147 , C a nd c) sp readin g b ro adly,
a ttain ing a di amet er al' 50 mm in 14 da ys a t room te m pe ra ture (20 -25C),
co ns isting al' a fin e funiculose felt , a ppear ing velvety, flat , not fu rrowed , with
thin eve n margin surro u nde d by a n arrow fr inge a l' sub merge d hyphae
ex te nd ing abou t 1 mm beyond th e ae ria l grow th; heavily sparing throughout in
ye llow ochre sh ades (Sguy, PI. X V , 225); ex u date a b u ndan tly produced , as
large crys tal c1ear ye llow d rops ; odo ur lacking; no colou re d pi gment d iffusing
into th e su r ro u nd ing agar; colo ny reverse d evel oping re dd ish ochre shades
(colou r pia te 147 , e); pen icilli as descr ib ed aboye bu t with co n idiopho res
com monly longer and a r ising primarily fro rn th e su bstratu m .
The sp ecie s d escription is ce n te re d upon th e type stra in CBS 220 .66 = VKM
F- 1,035 = CMI 117, 658 = ATcc 18,330 = IJFM712 5 ,isolat ed in1949byT .V.
C hala bu da from soil under co n ifers in the e nv irons of Ki ev , U. S . S .R.
Penicillium arenicola C halabu da is a qui te di stinct ive species wh ich a ppears to
be most satisfactorily assignab le to th e P. breoicompactum series than a nywhe re
else, but it c1early differ s frorn the othe r for rns inc lu de d in th e series, b oth in its
cu ltural charac te ristics and in th e det ail s of m orphology , particularly th e lack
of green a nd blue shade s in its colon ies.

148 . Penicillium brunneo-stoloniferum Abe ex R amre z : fi g. 148 , col our

plat e 148 , and SEM plat e 25, (bottom : left ).
Penicil/ium brunneo-stoloniferum Abe - J. Gen. Appl. Microbiol. Tokyo. 2:
104-105. 1956. See also M. M. Kulik, 'A Compilation of Descriptions of New
Penicil/ium Species' . Agric. Handbook No. 351. Agric. Res. Ser. U.S. Dep.
Agric. pp . 22-23. 1968.

Penicillium brunneo-stoloniferum was isolat ed from J apanese soil by S . Abe in

194 9 , wh o prop osed th e n ame P. brunneo-stoloniferum and described th e species
without providin g a Latin diagn osis. F u rthe r more, th is species wa s

(J)

68

le:

~ ~Q)

C);.,@
:~
0 11

l~m

'{' ..l

Fig. 148. Camera lucida

drawings of Penicillium
brunneo-stoloniferum Abe ex
Ramrez, CBS 317.59. A,
Detailed drawings of the
penicilli. B, Mature conid ia.
e, Habit sketches.

syno ny rnized with P. ochraceum b y Smith (196 3), but followin g the opinion of
Sa mso n et al. (1976), at present P. brunneo-stoloniferum ca n best be regarded as a
separa te spec ies. In orde r to va lidate the spec ies a L atin di a gn osi s of it is given
below; th e n ame of the ori gin al a u thor is m aintained under R ecom m end ation
46 C of th e International Code of Bot anica l N omenclature (Stafleu (E d .),
1972 .).
Latin dia gn osis of Penicillium brunn eo-stoloniferum Ab e ex R amrez :
Coloniae in agaro Czapekii, post duas hebdomadas ad 20-25C, 45 mm in
diametro attingunt, velutinae, planae , denso te xto basali coacto mycelio , parum
prominentes ad centrum , cum aequabili margine e submersis hyphis (3 mm)
formatur; fortiter sporulantes fusco colorantur; exsudatio adest , in magnis guttulis
fl avis; odo r abest; reverso sine colore vel pallido flavo; cum pigmento flavo
diffluente in medio; penici lli pleraque biverticillati asymmetrici, valde compacti les,
brev is; con idiophora septata, laevia , e substrato vel e mycelio basali exoriuntur,
plerumque usque 120-200 .tm in longitudine per 4.0-5.0 .tm in diametro, cum

' l' j ' l'

apicibus tenus 4.5-5.5 Jt m inflatibus; rami brevia, 17-20 Jtm per 3.5-4.0 Jtm;
metulae 3-6 per verticillum, plerumque 20-30 Jtm per 5.0-5.5 Jtm; phialides
lagaeniformae, 13-24 Jtm per 4.0-5.0 Jtm, 4-7 per verticillum, cum exiguo
acuminato collulo; conidia elliptica vel subglobosa, plerumque 4-6 Jtm per 3.5-4.0
Jtm in diametro, fortiter echinulata; in catenis laxis parallelis vel diversis formantur.
Neque ascomata neque sclerotia visa sunt.
Coloniae in agaro Czapekii cum extracto levedinis, post duas hebdomadas ad
20-25C, 40 mm in diametero attingunt, velutinae, cum parum radiatis sulcis,
denso texto basali coacto mycelio, cum margine continuo; fortiter sporulantes,
fusco colorantur; exsudatio adest, magnis succineis guttulis; odor abest; cum
pigmento succineo in medio diffluente; reverso rubro cinnamomeo. Neque
ascomata neque sclerotia visa sunt.
Coloniae in agaro hordeaceo, post duas hebdomadas ad 20-25C , 30 mm in
diametro attingunt; cum margine inaequaliter, velutinae, planae; fortiter
sporulantes, fusco colorantur; odor et exsudatio absunt; sine pigmento diffluente in
medio; reverso ochraceo cinnamomeo; neque ascomata neque sclerotia visa sunt.
Typus cultura e solo Japonico a S. Abe et Y . Nakano in anno 1949 isolata est,
FAT 1132 = CBS 317.59.

Colonies upon CA (colour plat e 148 , A and a) attaining a di ameter of 45 mm

in 14 days at room temperature (20 -25 C) , almost fla t , no t furrowed, velvet y,
very slightly rai sed towards the centre, with eve n margin surro unde d by a
fringe of submerged hyphae extending about 3 mm beyond the aerial gro wt h;
heavily spori ng througho ut in brown shades (Sguy, PI. IX , 132); exudate
abu ndantly produced as large yellow drops ; odour lacking : pale yellow coloured pigm ent di ffusing into th e su rrounding agar ; colony rev ers e (colour
pla te 148 , a) developing pale ye llowish shades ; pe nicilli typi cally asymme trical,
consisting (fig . 148 , A) of termina l verticils of me tulae or lat eral ram i each
bearing a verticil of metulae, wit h all elements closely a ppress ed sugges tive of
the Penicill ium paxilli Bainier coni dial str uctures in the P. brevi-compactum series;
with conid iophores septa te a nd smooth walled , arisin g primarily from the su bstratum or from the ba sal felt , rangin g from 120 to 200 flm in length by 4-5 flm
in di ameter, with api ces som ewhat enlarged u p to 4.5 -5 .5 fl m in diamete r;
rami whe n present short, m ostly 17-20 fl m by 3.5-4 .0 flm ; m et ulae
cylindrical, u su ally 20 - 30 fl m by 5.0-5 .5 fl m ; phialides bottl e shaped, about
13- 24 fl m by 4 .0 -5 .0 fl m , n umberin g 4- 7 elements in th e verticil, tap ering to
a conspicu ous ly long narrowed neck ; conid ia u su ally elliptical to su bg lobose,
4- 6 flm by 3.5-4.0 flm in di ameter , conspicu ous ly echinulate (fig . 148 , B, a nd
SE M pla te 25, bottom : left ); con id ia tending to form loosely parallel or
di vergent chains (fig. 148 , C).
Colon ies upon CYA (colour plat e 148 , B and b) a ttaini ng a diameter of
40 m m in 14 days at roo m temperature (20 -25 C) , sim ila rly text ured as

lescribcd above but dccper velv cty und with Iew deep radial urrow s, with
e vc n mur gin c n tirc ; heavily spo r ing throu ghout in brown shadcs (S guy, PI.
X LV 11, 696 ); cxud a te abundantly produced, as yellow-coloured drops ; odo u r
luckin g; amber-colourcd pigment difu sin g into th e su r ro u nd ing a gar; colony
rc vc rsc dev eloping reddish ochrc sh ades (colour plate 148 , b) ; penicil1i
l'ssentially as described aboye .
C olon ies upon MEA (co lou r plat e 148, C a nd c) grow ing restrict edl y ,
uu ainin g a di amct er 01' a bo u t 20 mm in 14 days a t room tempera ture
t ~ () - 25 C) , co ns isting 01' a d en se velvet y felt , almost flat , not fur rawed , with a
slight clev ation in th e colony ce n tre in a cra te r ifor m pattern , with irregul arly
.Iissccted m argin ; hea vily spo ring throu gh out in dark brown shades (Sgu y , PI.
X LV II, 701) ; ex u date a nd odo u r lacking; no coloured pi gment diffu sin g into
lile su r ro u nd ing agar; colony reverse (co lou r plate 148, c) dev eloping ochre
shades ; penicil1i esse ntially as descr ibed a bo ye.
Thc sp ecies d escription is cen te re d upon th e typ e strain C BS 3 17.59 = C M I
hH ,217 = ATCC 18 ,311 = IFO 5 ,72 7 = FAT 1,132 = QM 7, 306 = IJFM 7136 ,
.solat ed in 194 9 by S. Abe a n d Y . N a ka no frorn Japanese soil.
Penicillium brunneo-stoloniferum Abe is a di stinctive spec ies whi ch a ppears to be
III ClSt sa tisfac to r ily ass ignable to th e P. breoi-compactum se ries th an a ny whe re
lsc , but it clearly differs fro m th e othe r species included in it , b oth in its
ultu ral characte r istic s a nd in th e d etails 01' m orphology .

Species of dubious position

Penicillium amagasakiense Kusai - Ann . Rep . Sci. Works Fac . Sci. Osaka Univ. 8:
43. 1960. See also M.M. Kulik, 'A Compilation of Descriptions of New Penicillium
Species'. Agric. Handbook No. 351. Agric . Res. Ser. U.S. Dep . Agric. p. 16.
1968.

Penicillium amagasakiense Kusai is a species d escribed in 1960 that is believed

lo be closel y relat ed to spec ies included in the Asymmct rica-velutina su bsec tion. H ow eve r , it is presented h ere as a spe cies 01' dubious position a n d h as
not been included in th e keys to species since th e au thor 01' th e present stu dy
was not able to exam ine a n d stu dy the typ e strain 01' th e spec ies in question,
becau se al1 attem p ts to obtain it fram International C u lture Col1ectio ns fail ed .
Ku sai ' s or iginal decription fol1ow s:
'Colonies of P. amagasakiense on Czapek's agar, steep agar, and Sakaguchi and
Wang agar grow rapidly at 25C. (Average colony diameter after 12 days was 52
mm on Czapek's agar, 70 mm on steep agar, and 60 mm on Sakaguchi and Wang
aqar). This fungus grows more rapidly than P. notatum and P. chrysogenum and
furthermore it can grow in a medium with nitrite as the sole source of nitrogen . (lt is
a producer of notatin, also called glucose oxidase.)

Colonle s grown on agar have a velvety textura and are white. The surface of the
colony is smooth and the underside is cherry yellow. The colony exudes a cherry
yellow substance int o the culture medium during cultivation.
It is difficult to induce conidia formation by P. amagasakiense on the above three
media . To observe the production of conidia , potato agar was used. The penicillium
was then typically biverticillate and asymmetric. Conidiophores of 80-200 ,tm by
3-4.5 ,tm arising from the substratum. No branches were observed. Metulae usually
in groups of 4-8, 9-15 ,tm by 2.5-4.0 ,tm, with the apices not enlarged.
To classify this strain, some microbial properties of P. amagasakiense were
observed. The results indicated that the growth of this strain is very rapid on Czapek
or steep agars and that it can assimilate nitrite . The above properties, the textura
and colour of the surface and the reverse of the colonies, and the exudate were
fairly similar to those of P. !i!acinum Thom . There are, however, sorne differences
between .the two Penicillia: Penici!lium amagasakiense rarely forrns conidia and P.
!i!acinum cannot produce glucose oxidase.' (Without a Latin diagnosis).

Penici//ium aspergi//oides Rudakov - J . Bot. U.S.S.R. 44: 867.1959. See also M.M.
Kulik, 'A Compilation of Descriptions of New Penici!lium Species'. Agric.
Handbook No. 351. Agric . Res. Ser. U.S. Dep. Agric. pp. 17-18. 1968.
Penicillium aspergilloides Rudak ov is a spec ies described in 1959 that is
believed to be closely relat ed to the species included in th e Asym metri ca - velutin a su b-sectio n. H owever, it is presented here as a no ther form of
dubiou s positi on and ha s not been included in th e keys to spe cies in th e present
study for th e same reasons not ed abo ye .
Rudak ov ' s original di agn osis follows :
'On solid media it forms circular colonies 2-3 mm deep. Colour of the mycelium
white. The growing margin is surrounded by a fringe in a radial pattern 1 mm long .
After 3-4 days a pale orange colour appears in the colony centre . This colour
gradually becoming darker and extending to all hyphae which grow wider upon the
substrate to 40-50 mm in diameter.
In aging the aerial hyphae are lost, substituted by a layer of conidial structures.
On further growth, colony colours become duller and duller, changing from orange
to dirty crimson . Sometimes short tufts of gray aerial hyphae appear in the colony
centre. Yellowish milky-coloured pigment diffusing into the surrounding agar.
Conidiophores of young colonies are rather short. When older, the conidiophores
are coarser and longer. In old cultures , the conid iophores become slightly curved,
sabre shaped, 1.5 mm long by 5-6 ,tm in diameter.
The colour of the conidiophores is pale brown with an orange casto The walls are
thick and smooth . The penicilli consist of verticils of metulae, each bearing a whorl
of phialides, or occasionally simple penicilli (appearing monoverticillate) consisting
of small c1usters of phialides at the apex of the conidiophore. Usually variously

rmn llud, w ith rami arranged in seve ral stagos bolow the level of the metulae, but
Inootly ono -sided branched . Rami are usu ally arranged together in the pattern of a
Illr oo-pronged penicillus, abou t 30 J!m in longth , bearing at their apices small
vurtlclle of generally 2 metulae per cluster, about 9 J!m in length by 3.8 p,m in
lmnutor, usually cylindrical.
Phlolides are bottle shaped somewhat swollen in the middle, tapering to a
narrowed neck, commonly 7.5 by 2.3 p,m, in small c1usters of 3 elements per verticil.
nidia ellipsoidal, colourless, with smooth walls , 1.9-2.4 J!m by 2.2-2 .8 J!m ,
fonnlng long cha ins , loosely parallel or divergent .' (Without a Latin diagnosis).
ITrenslated from the Russian by Prof. L. Vilas) .
'mic:illium pidoplichkanum Chalabuda - Nov. Syst . Ins . Bot. Komaravii, Acad. Sci .
U.S. S .R. 6: 161. 1950. See also M.M. Kulik , ' A Compilation of Descriptions of
Now Penicillium Species' . Ag ric. Handboo k No. 351. Agric. Res. Ser . U. S. Dep .
Agrc . pp. 65-66.1968.
l 'rnictllium pidop lichkan um Chalabuda is a no the r species of dubious position
11t ;1I is believed to be closcl y rel ated to th e spec ies included in th e A symme-

trie'a - ve lu tin a su b -sec tion . For identical rcasons as statc d aboye , it is

I'l'l'scllt ed he re a nd h a s not been included in the keys to specics .
( 'halabud as o riginal di a gnosis follows:
' Colonies upon Czapek's agar growing restrictedly, velvety, at f irst white , shad ing
lo brow n after a long period ; co lony reverse at f irst colourless, becom ing later milky
yullow. Colonies upon sugar beet root extract often irregularly shaped, less circular,
vulvety , pubescent, tough basal f elt. At first white, becoming later yellow and
finally brown, not zonate , colony reverse dark brown. Hyphae pale yellow coloured ,
o l ten conspicuously septate, 15.4-345 J!m by 2-3.5 J!m . Heavily sporing, with
ponicilli typically biverticillate asymmetrical, arising from the basal felt in a close and
uven stand, consistng of verticils of 4-7 metulae, abou t 8.5-12 J!m. Phialides
bot t le-shaped, 8-14 by 2.8-3.5 p,m . Conidia limoniform, delicately roughened,
3.5- 4.5 by 2.8-3.5 J!m in diameter, pale yellow in rnass.'
(Translated f rom the Russian by Praf. Dr . L. Vilas),

Penicillium amagasakiense Kusai and P. aspergilloides Rudakov were both

described without providing a Latin diagnosis.

419

haptcr 6

Asyrnrnetrica-lanata sub-section

IN THI S SUB -SECT ION , T HE C O LON IES T YPICALL Y CONS IST OF COTTONY, LA NAT E OR
deepl y floccose networks of fin e aerial veget ati ve hyphae ove rlaying th e basal
k it, with co nidiopho res gen erally arising as branch es from ae rial h yphae , less
lrcquen tly from the su bs tra tu m , with co nid ial struc tures com mo nly a p pe aring
at first in th e colon y cent re, th en progr essin g gradually toward s th e m argin al
areas . Pen icilli u su ally com parat ively large, irregula rly ram ified, with metulae
a nd phi ali des frequently arising at different nod es of th e con id iophore . The
conid iopho res are usu all y fairl y coa rse with wall s more or less rou ghened.
Conidia l cha ins forming tangled m asses ra ther th an well-de fine d colu m ns .
T he lin es of demarcati on between the Lanata su b-sec tion a nd th e other su bsect ions of th e Asymmetrica are ad mi ttedly arb itra ry. T he L anata su b -sec tion,
indee d, me rges in sen sibl y into each of the ot he r su b-sec tions of the
Asymme trica, and species are pl aced in one or th e other on th e judgement of
the obse rve r.
Fo rms that m ay be strongly lanate but produce typi call y di varicat e pe n icilli
are placed in th e Divaricata. Sp ecies wh ich m ay sho w more or less floccul en t
colonies but a p pearing velvet y are included in th e vclu tina . Form s
predominan tly lanat e , but sho wing tra iling or a sce nd ing ropes of h yphae are
ass igne d to th e Funicul osa . Species with con id iop ho res a rising primarily fro m
the su bs tratu m, collec ting in d efinite bundles or tufts, are placed in the
Fascicul at a.
In fact , th e Lan ata su b-sec tio n of th e Asymmetrica is a n artificia l gro u ping
01' forms wh ich is ret ained here primarily as a m att er 01' conve nience , a nd is not
pres u me d, th erefore , to represent a natu ral grou ping of spec ies th at should be
cxpec ted to ex h ib it unique bioche m ical or ph ysiological charac te ristics.
O f th e nine spec ies recogni zcd by R aper a nd Thom ( 1949) in thi s series,
only Penicillium camemberti T ho m, P. comm une Thom , and P. lanosum W estling
still remain a s separate species, th e other six, namely P. caseicolum Bainier, P.
lanoso-viride Thom , P. lanosocoeruleum Thom , P. biforme Thom , P. lanoso-griseum
T ho m, and P. aurantio-candidum Di erckx h ave been reduced to sim ple
synony m ies: P. caseicolum Bainier is ac tually regarded as syno ny mo us with P.
camemberti Thom (Samson et al. 1976), P. lanoso-viride Thom is regarded as
syno ny mo us with P. verrucosum Di er ckx va ro verrucosum (Samson et al. 197 6),

-r "- \1

and th e rernaining four spe cics are a c tu ully rcgu rded as syno ny rnous with l ',

oerrucosum Di erckx va l'. cyclopium (W cs tling) (Sa rnson et al. 197G).

H owever , three addi tional sp ecics have bee n includcd in thi s series in lhe
cu rre n t study, namely: Penicillium gnseum Bon orden, P. echinosporum Nehi ra (1,\'
R amrez, and P. giganteum R oy el Singh , without presuming, th eref orc, a c10HI'
relationship bet ween th em . They are a ll con sid ered together prirnarily aH /1
m atter of co nvenience . H ow ever, they sho w sorn e charac te ristics in co rnmon ,
particul arly the lanat e characte r of th e colon ies.

K ey to se r ies a n d spe cies

Pag
1. Colonies typically lanate, without evidence of synnemata or ropes of
hyphae or with such structures reduced and insconspicuous if present o
A. Colonies remaining persistently white with the ripening of conidia or
becoming slightly coloured in gray-green shades.
P. camemberti series . ... 420
1. AII elements of the conidiophore except the stipe smooth walled.
P. camembertiThom . . . . 421
2. Conidiophores conspicuously roughened.
P. griseum Bonorden ... . 423
B. Colonies developing some shade of green or brown.
P. commune series . ... 425
1. Conidiophore walls roughened .
a. Conidia smooth walled or nearly so.
P. commune Thom . ... 426
b. Conidia strongly echinulate.
P. giganteum Roy et Singh . .. . 433
2. Conidiophore walls smooth.
a. Conidia globose or nearly so, less than 4 Lm in diameter, delicately roughened.
P. /anosum Westling .. .. 428
b. Conidia strongly elliptical, conspicuously spinulose, up to 6Lm in
long axis.
P. echinosporum Nehira ex Ramrez .. . . 430

Penicillium camem berti se r ies

O U T ST ANDIN G F EAT U RES

Colo nies floccose , cottony, d eepl y lanate , at first white and at m aturity
remaining so or becoming pale gray-gree n dep ending upon th e species; colon y
rever se yellowish cream and agar uncoloured or nearl y so .
Conid iopho re s arising from th e substratum or from th e loose ae rial network

''1' :,( I

01' vcgct ntiv c hyph ae , va riabl e in lcn gth up

l o 600

11lIl

by 3 - 4 ,.un in di amct cr,

with walls smooth 0 1' consp icu ou sly ro ug hc ned.

Pcnicilli asymmctrical , irrcgularl y branch ed a n d va r iable in pattern , with
mctulae a nd phi alides usu all y borne a t differen t levels, beari ng co n idia in
I a ng led ch a in s.
Con idia at first elliptical , beco m ing subglobose at m aturi ty , smooth walled
(JI' nearly so, co m p ara tively large up to 5 p,m in d ia mete r.
Two species a re inclu de d in th is series: Penicillium camemberti Thom a nd P.
griseum Ronorden . The for mer spe cies reg u la rly occurs a nd is res po n sib le for
the pradu ctio n of soft cheeses of the Came m be rt ty pe, in clu d in g Camembert ,
Brie , a n d N e u fchate l. It s n atural h abitat is still not known . AIl strains reported
in th e litera ture h a ve been ob ta ine d fram chee se sa m ples 01' fro m the ri pen ing
roo ms in th e cheese in du stry . The la tt er species, on the othe r h and , ha s never
becn sa tisfac tor ily idc n tified a n d descr ibed althou gh thi s n ame ha s b een used
sev era l times by va r ious au thors . The strain exam ine d b y us was received from
ih e American T yp e C ult ure Collection under th e numb er ATCC 20, 068 as
Penicillium griseum Bonorden. It h as tentat ively been assigned to this ser ies on
accou nt of its cu ltural ch a racter istics a nd d et ail s of morphology .

149. Penicillium camemberti 1 'hom : fig. 149 , colou r plat e 149, and SEM
pla te 25 , (bo tt om : right) .
Penicillium camemberti Thom - Bull . Bur. Anim. Ind. U.S . Dep. Agrie . 82: 33.
1906; see also Thom, Bull. Bur. Anim. Ind. U.S. Dep. Agrie. 110: 50. 1910; also
Thom, 'The Penieillia', pp. 312-313.1930; also Raper and Thom, 'A Manual of the
Penicillia', pp. 426-427. 1949; see also Samson et al. Antonie van Leeuwenhoek.
43: 342-344. 1977.
Synonymies

Penicillium aromaticum 111 Sopp - Z. Bakt. Parasitenk. Inf . Hyg. Abt. 11. 4: 161.
1898.

Penicillium album Epstein - Areh. Hyg. 15: 360. 1902. (Non P. album Preuss Linnaea. 24: 135. 1851, 01' P. album Rivolta - Paras. Vgt. p. 452. 1873).
Penicillium epsteinii Lindau apud Rabenhorst - Deutsehl. Krypt. Flora, Pilze Bd.
8: 166. 1907.

Penicillium rogeri Wehmer apud Lafar - Teehn. Myeol. 2: 226. 1906.

Penicillium caseicolum Bainier - Bull. Soc, Myeol. Franee. 23: 94. 1907.
Penicillium camembertiThom varrogeriThom - Bull. Bur. Anim. Ind. U.S. Dep.
Agrie. 118: 52. 1910.

Colon ies upon C A (colour pIa te 149 , A a nd a) a ttain ing a di ameter ofa bo u t

1::

LwJ..l=J

10pm

O sO
0 0 0 0
0 0 0
0 0 0

00 O (l{ .

Fig. 149. Camera lucida

drawings of Penicil/ium
camemberti Thom, CBS
299.48. A, Detailed drawings
of the penicilli . B, Mature
conidia.

30 mm in 14 days at room temperature (20 - 25 C ), consisting of a dense basal

felt ove rla id by a deepl y floccose ae rial m ycelium usu ally up to 10 mm de ep , at
first white , remaining persistentI y so or shad ing to yellowish , pinkish or gr aygr een colou rs, th e latter shade com mo nly appearing rather lat e ; eve n margin
surro unde d by ~ frin ge of su b me rge d hyphae exte ndi ng ab out 1 mm beyond
th e ae rial gro wth; medium spo ring throu gh out ; odo ur faint, m ouldy ; exudate
lacking or as minute d roplets enmes he d in th e m ycelial felt ; penicilli asy m me tri cal (fig. 149, A) with conidio pho res up to 500 p.m lon g by 3.0 - 4 .0 p.min
di ameter , two- to thre e-stage branched; stipe of the conid iopho re u sually rough
wa lled, seldo m smoo th, but rami , m etulae a nd phialides are smooth walled ;
conid iophores arisin g from subme rged hyphae , occasionally from aerial
hyphae , usuall y septa te ; ra m i, usu ally two pe r vertici l, 15- 20 p.m by 3.5 - 4 .0
p.m; metulae in gro ups of 2 -3 elem ents in the ve rticil, 1O-1 5p.m by 4.0-4.5
utt , with apices slightIy enla rge d up to 4- 5 p.m in di ameter; phialides bottle
shape d , 9 - 15 utt: by 3 .5- 4. 5 p. m, tapering to a short neck , numbering 2-8 in
th e verticil; conidia co m paratively large , globose to su bgIobose , 4 -5 p.m in
diamet er, smo oth walled or nearl y so (fig. 149 , B, a nd SEM plate 25, bottom :
right ); coni d ial chains tangled . C olony reverse (colou r pIat e 149 , a) in pinkish
yellow sh ades; no coloured pigm ent diffu sin g into th e surro u nd ing agar.

lolo n ie upon C YA (co lour plat c 14-9, B ami h ) a tta in ing a di amct cr 01'
uho u t 40 mm in 14 d a ys a l roo m tcmporutu rc (20 -25 C), sim ilar in tcx tu re
nnd cu ltural charac tc ristics lo th osc grow n upon C A, hut m ore deepl y Iloccosc ;
hc u cr spo ring in vc ry palc grccn shadcs; odo u r ca rthy ; ex udate as minu te
. Iro plcts cn mcshe d in the mycclial felt ; co lo ny rcve rse radiatcl y wrinkled
(colou r plate 149 , b) , developin g yello wish crcam shades; no coloured pi gment
d ifTu sin g into th e su rro u nd ing agar; penicilli esse nt ially as d escrib ed aboye.
Colonies upon M EA (co lour pla te 149 , C a nd c) a ttain ing a diameter 01'
nbo u t 30 mm in 14 days a t room te m perat ure (2 0 -25 C), sim ilar in texture
.md cultu ral characte r ist ics to th e preceding , bu t spo ring m ore ab u n dan tIy in
pastel gree n shades (Sguy, PI. XXVII, 400 ); odour an d ex u date la cking; no
roloured pi gment diffu sing in to th e su rrou nding agar; colony reverse (co lou r
pla te 149 , c) in yellow ish cream shades; grow ing m argin su r ro u nded b y a
frin ge 01' su b merged hyphae extending abo u t 2 mm beyond the ae r ia l gro wt h ;
pcn icilli esse ntia lly as dcscribed a boye but with the stipes 01' th e co nid iopho res
a pp caring m ore co nsisten tIy roughe ne d than u pon CA a n d C YA .
T he species description is ce n te re d upon th e typ e st rain ATCC 10 ,4 23 = NRR L
1177 = C BS 299 .48 = IJ FM 7217 , isolated by C h . T ho m in Storrs , Con necticu t,
USA , in 1904 fro m Frenc h Camembert cheese . T he strain h as re maine d in
con tinuou s laboratory cu ltivatio n since th at time wi th out a ppare nt chan ges in
cultural ap pearanc e 01' dct ail s 01' m orphology .

150. Penicillium griseum Bonorden : fig . 150 , colour pia te 150 , a n d SEM
p ia te 26 , (top : left) .

Penicillium griseum Bonorden - Abh. Geb. Mykol. p. 92. 1864. See also Thom,
'The Penicillia', pp. 563-564. 1930.
Colo n ies upon CA (col our pIa te 150 , A a n d a ), growing r apidly , attaini ng a
diame te r 01' 55 mm in 14 d ays a t roo m te m pe rature (20-25 C) , co ns isting 01' a
cIose-text u re d b asal felt overlaid by a deeply floccose aerial m ycelium infl uencin g greatIy th e colony a p pearanc e , with ce n tre irregul arly r ai sed b y tufts 01'
whi te vegetative hyphae ; thin eve n m argin surrounded by a narrow fr inge 01'
subme rge d h yphae cx te n d in g a bo u t 5 mm beyond th e ae r ial gro wth; hea vily
spo r ing throu ghou t in pal e gray-g re en shade s, but with co n id ia l areas m ostIy
obs cured by the wh ite ove rg ro wth 01' floccose vegetativ e hyphae ; colony surface
dcvcloping pale green shades with the rip ening 01' co n id ia (Sguy, Pi s. XXI ,
3 10, a nd XX , 299) ; odo u r fa int, ' m ou ld y ' ; ex u date limited in a mou n t as
m in u te d roplet s enmeshe d in the mycelia l felt; n o coloured pi gment d iffus ing
into the su rro u nding agar; colo ny reverse (colour pla te 150 , a) develop ing
ye llow-orange shades ; penicilli (fig . 150 , A) ty p ica lly asym metrical , co n sisting
01' two- 01' three-st a ge branch ed stru ctures, with all c1emen ts 01' th e con id io-

--:

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'

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:.:;

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ffi

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.:':".

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o:::

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."

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:~:'

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~~:.

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!Y

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1rl1r
100jJm

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Fig. 150. Camera lucida
drawings of Penicillium
griseum Bonorden, ATCC
20,068. A, Detailed drawings
of the penicilli. B, Mature
conidia. C, Habit sketches.

phore except phialides , conspicuo usly rou ghen ed ; conidiophore s arising from
sub merged hyphae and from aerial hyphae , usually 100 -150 tm by 2.5- 3.5
tui , septate , with api ces enla rged in a vesicle up to 5- 6 tm in diamet er; rami
when prese nt upto 20 tm by 3 .0 ustv ; metulae in groups of2- 3 elements in th e
verticil , of unequal length , com monly 11-18 tm by 2.5 - 3 .5 tm; phi alides
bottl e shape d, in clu sters of 2-6 elements in th e ve rticil, 9-1 2 tm by 3- 4 tut,
tap ering to a short neck ; con id ia globose to subg lobose , sm ooth walled or
nearly so (fig . 150, B, and SEM plat e 26, top : left) , 3 .5-5 .0 tm in diamet er;
conid ial chains forming loo sely parallel short colu m ns (fig . 150 , C ).
Col oni es up on C YA (colour plate 150 , B and b) attaining a diameter of
about 50 mm in 14 days a t room temperature (20 -25 C) , con sisting ofa close
textured b asal felt , strongly sulca ted by numerous radial furrows and slightly
raised in th e colony centre , a ppearing deepl y velvety to subla nate, with th e
colony surface ove rlaid by a den se netw ork of dirty white-coloured aerial
vegetative hyphae influen cin g greatl y th e colony appeara nce; thin membra
nous m ar gin , su rrou nde d by a fringe of submerged hyphae exte nd ing about 1
mm beyond th e aer ial gr owth ; medium sporing in the same shad es as described

aboye; with co uid ia l ureas obs cured by thc dense whit e ovcrgrowth ; exudale
nnd odour lacking; no colou red pigm cnt dillusin g into the surrounding agar;
colony rcv crs c (eolour plat e 150, b) in yellow-orange shades; penieilli as
described above .
Colonies upon MEA (colour platc 150, C and c) attaining a diameter of 50
111m in 14 days at room temperature (20- 25C) , consisting of a looser textured
basal felt, not furrowed, but raised in the centre, with thin margin surrounded
by a wide fringe of submerged hyphae extending about 4 mm beyond the aerial
growth; colony surface partially overlaid by a white overgrowth of densely
woven aerial hyphae ; sporing sparingly in marginal areas and heavily in the
centre in blue-green shades (Sguy, PI. XXIX, 429) ; exudate and odour
lacking; no coloured pigment diffusing into the surrounding agar ; colony
reverse (colour plate 150, c) developing pale orange shades in the centre and
pale cre am to colourless elsewhere with a green cast showing through ; penicilli
essentially as des cribed aboye .
The species description is centered upon the type strain ATCC 20 ,068 = IJFM
7206, isolated from soil. The type strain , probably lost, was reported by
Bonorden as isolated from rotten leaves in 1863, in Westphalia , G ermany .
Furthermore, it was so poorly des cribed by the author as to make th e identifi
cation impossible.

Penicillium commune series

UTSTANDING FEATURES

Colonies growing moderately to rather restrictedly, typically loase textured,

lanate, comparatively deep, commonly up to 1- 2 mm, usually medium to
heavily sporing with conidial areas in dull bluish green, yellowish green 01'
brownish shades ; with conidiophores comparatively coarse, either conspi
cuously roughened 01' smooth walled ; colony reverse usually devcloping
orange, yellowish 01' drab shades.
Conidiophores arising from the substratum 01' from the basal felt , compara
tively coarse in most species, often long, up to 500 /lm 01' more, som etimes
short, depending upon the species.
Penicilli com pa ratively large, often irregularly branched , with metulae and
phialides usually borne at different levels in the penicillus .
Conidia variable in form and size, ranging from globose, subglobose to
elliptical, either smooth walled 01' conspicuously echinulate, with conidial
chains usually tangled 01' loosely parallel .
The series as understood by Raper and Thom (1949) has been broadened
enou gh to accomodate two additional spe cies, namely, Penicillium echinosporum
Nehira and P. giganleum Roy el Singh, together with P. commune Thom and P.
lanosum Westling, originally included in the seri es,

' I':':ll

Penicillium communc ser ies is probabl y artificial in cha ra ctc r. H o wcvcr , it is

rctaincd herc prim nril y as a ma ttc r of co nvc n ic ncc ,
151. Penicilliu m commune Thom : fi g. 151 , colo ur plat c 151 , and S EM

plate 26, (to p: left).

o
'. 1

"

', } lO}lm
.'

-"..,

~r"" 1\1 ~f
(Y!i$!;'~~ e
\ :;;r...
1

. ~

'-------'

100 jrrn

..N8ogo0

ir

)U i~ 86~c?
oogoo
00 0
O
R .

Fig. 151. Camera lucida

drawings of Penicil/ium
commune Thorn , CSS
311.48. A , Detailed drawings
of the penicilli . S, Mature
conidia. C, Habit sketches .

Penicillium commune Thom - Bull, Bur. Anim. Ind. U.S, Dep. Agrie. 118: 56-57.
1910; see also Ch. Thom, 'The Penieillia' , pp . 324-325. 1930; also Raper and
Thom, ' A Manual of the Penicillia ', pp. 439-440. 1949.

Synonym
Penicillium fusco -glaucum Biourge - Monographie, La Cellule. 33: 128-130. 1923.
See also Thom, 'The Penicillia', pp. 325-326. 1930.

Colonies upon CA (colour piate 151 , A and a) a ttaini ng a diameter of 35 m m

in 14 days at room temperat ure (20- 25 C), cons isti ng of a tough , close-

tcxturcd basal Ielt , decply llocco sc, ah out ;OO to 700 1L1lI d ecp, zon at c, su lca icd
by few sh allow radial lurrows: with whit c cvc n m argin surroundcd b y a fringc
01' sub mc rged hyph ac extc nd ing a bo u t :~ mm bcyond th e aerial growth ; colony
s u rface gradually raised towards the cen tre; spo r in g abundantIy in zones in
diffcrcnt shades of blue-green (Sguy , PI. XXXI , 465, 463, 462), depending
on th e degree of maturity of conidia ; exu d a te colourless, as minute droplets
cnrneshe d in the mycclial felt ; odour ' m ou ld y' ; amber-coloured pigment diffusing into the surrounding agar ; colony reverse (colour pIate 151, a) developing
reddish sienna brown shades in the centre and subcentral areas and pale
cinn a m o n in the margin ; penicilli (fig . 151, A) typically asymmetrical, with
clcm ents of the penicillus arising at different levels of the conidiophore in an
irregular pattern; conidiophores varying from very short up to 500 lLm in
length by 5 .0 lLm in diameter in marginal areas of older colonies , with walls
con sp icu ou sly roughened to coarsely granular in age; rami variable , up to
15-20 lLID by 2.5-4 .0 lLm ; metulae in sm all groups of two elerne n ts in th e
verticil , usually 15-20 lLm by 3 .0-3 .51Lm; phialides few in the verticil, 2-4 per
cluster, 12-17 lLm by 3-4 tui , tapering to a conspicuous narrowed neck;
con idia elliptical to subglobose, usually 4- 5 lLm, occasionally up to 7 lLm in
long axis , with walls smooth 01' nearly so (fig . 151 , and SEM plate 26 , top :
right) ; con idia l chains tending to diverge (fig. 151 , C) .
Colonies upon CYA (colour pl ate 151, B and b) spreading broadly , attaining
a di amet er of 50 mm in 14 days at room temperature (20-25 C), consisting of
a close-textu re d basal felt, strongly sul cated by numerous a nd d eep radial
furrows, with centre slightIy raised , with the colony surface overlaid by a
floccos e overgrowth of white vegetative hyphae influencing greatIy th e colo ny
aspect ; con sp icu ou sly zonate; with thin even margin surrounded by a fringe of
submerged hyphae exte nd in g a b ou t 1 mm beyond th e aerial growth ; heavily
spo rin g throughout in various shades 01' green , brown-green and blue- green
(S guy , PIs . XXXI, 462, XXIX, 427 , XXX , 448) depending on the degree 01'
ripeness 01' conidia; odour strong, ' m ou ld y '; exu d a te abundantIy produced, as
cry stal cIear yellow amber drops; amber-coloured pigment diffusing into the
su r rou nd in g agar ; colony reverse developing deep reddish sienna brown
shades with margin cinnamon (colour pl ate 151, b) ; penicilli esse n tia lly as
described ab ove .
Colonies upon MEA (colour pl ate 151, C a nd c) attaining a di amet er 01' 30
mm in 14 days at room temperature (20-25 C) , consisting of looser textured
basal felt , plane, not furrowed , deeply velvety, with few floccose tufts 01' bluegray aerial hyphae in localized zones 01' the colony surface, with thin even
m argin surrounded by a fringe 01' submerged hyphae exte nd in g about 3 mm
beyond th e aerial growth; heavily sporing throughout in blue-green shades
(Sguy , PI. XXXI , 461) ; odour spicy; exud a te lacking; no coloured pigment
d iffusing into the surrounding agar; colony rev erse (colour pIate 151, c) in

yellowish cin n um o n shades w ith a greell casi sho w ing through; peuici lli cssen tialIy as descri bed aboye .
T he spec ies descri ptio n is ce ntcrcd lIpon th e ty pe strai n c ss 3 11.48 = C MI
39,8 12 = ATCC 10,428 = NRRL 89 0 = I FO 5,763 = Q M 1,269 = IJ FM 386 7,
whic h was isol a ted by C h . T hom a t Storrs, Con necticut, USA , from cheese in
1904. The cu lture h as been u n der continuous laborat ory cu lt iva t ion withou t
significan t change in cu ltural or mo rphological characteristics.

152 . Penicillium lanosum W estling: fig. 152 , colour plat e 152 , and SEM

plate 26 , (ce ntre : left).

t.cw.l.Lw.J

10

um

O
O8 O 0

00
0

O0 O 0
0
0 .0 0
O OO

" ~'l

\ \-'>~'. ~l.
~~,~'

Fig. 152. Camera lucida

drawings of Peni~illium
lanosum Westling, CBS
106.11. A, Detailed draw ings
of the penicill i. B, Mature
conidia . C, Habit sketches.

Penicillium /anosum Westling - Arkiv Bot. 11: 55, 97-99. 1911. See also Ch. Thom ,
'The Penicillia', pp. 317-318. 1930; also Raper and Thorn, 'A Manual of the
Penicillia', pp . 431 -434. 1949.

Colon ies upon CA (colour pi ate 152 , A and a) attaining a diamete r of 40 mm

in 14 days at room te m perature (20 - 25 C), co ns isting of a tough, closetextured basal felt ove rlaid by a floccose overgrowth, len ding a la n a te

ippea ra ncc lo th e colo ny surface whi ch is a l111 osI plano, showing a slighl
clcvation in th c cen tre , with ew shallow urrow s; cvc n margin surroundcd by a
trin gc 01' submcrgcd hyphac cxte nd ing alxiut ~ 111m bcyond the aerial growlh ;
slighlly sporing in marginal arcas in dark gr a y-green shades (Sguy , PI.
X X IX, 433) , c1scwhere , rcmaining persistently white owing to th e f1 occose
overgrowth; odour fragant, indefinite; exudate limited in am ou n t , as crystal
clca r minute colou rlcss droplets en me shed in th e mycelial felt; no colou red
pigment diffusing into th e surro u nd ing agar ; colo ny rev ers e (colour pl at e 152 ,
a ) in pale yellow to cream sh ades ; penicilli com parativcly large (fig. 152 , A) ,
usy mme trical, irregularl y ramified and tending to beco m e div ergent , with
rouidiophores arising co m m only a s short branch es from ae rial hyphae 100 to
~ O O /lm in length, less frequently from the substratum , ranging th en from 250
lo 600 /lm by 2.8-3 .3 tui, with walls smooth 01' nearly so; rami variabl e ,
usu all y 25 - 35 /lm by 2.8 -3.0 tui, arising in small clusters 01' two in th e
vcrticil, often arising low on th e co nid iophorc ; m etulae in numbers 01' 2- 3
clcme n ts in th e verticil , abou t 9-14 /lm by 2.5 - 3.5 usn, with apices slightly
cnla rged up to 4 .0-4.5 /lm in diamet er, commonly borne at different Ievels 01'
t he penicillus; phialides in clusters 01' 2-10 elements in th e verticil, a bo u t 9-11
Ilm by 3.0 -4.0 tuu ; tap ering to a conspicuous narrow ed and short neck ;
conid ia globose to sub globose , com parative ly small, usually 2 .8- 3.0 /lm in
diameter, with walls delicately gra nular (fig. 152, B, and SEM plate 26,
centre : left); coni d ial chains loosely parallel to tangled (fig. 152 , C) .
Colonies upon CYA (colour plat e 152 , B and b) attaining a di amet er 01' 60
mm in 14 days at room temperature (20-25 C) , similarly textured as aboye but
with colony surface strongly and deeply sulcat ed by numerous raidal furrow s,
and centre irregularly fold ed and raised ; thin growing m argin f1at, su rrou nde d
by a fringe 01' submerged hyphae exte nd ing a bou t 5 mm bcyond the aeri al
gro wth ; colon y surface overlaid by a dense network 01' white f1 occose aeri al
hyphae influencing greatly the colony appearance ; heavil y sporing margin al
a rea s in dark gray-green sh ades (Sguy, PI. XXIX, 44 3) , c1sewhere th e
co nid ia l are as are entire ly ob scurcd by th e d ense overgrowth ; od our fragrant ,
suggestive 01' soap perfumes; ex ud ate limited in amount , as minute colou rless
droplets en me shed in th e mycelial felt ; no eoloured pigment diffusing into th e
surrou nd ing aga r ; eolony rev erse (eolour plate 152 , b) developing pale orange
to er eam shades; penieilli essen tially as deseribed aboye .
Colonies upon MEA (eolour plate 152, C a nd e) spreading broadl y ,
att a ining a diamet er 01' about 60 mm in 14 da ys at room temperature
(20 - 25 C), eonsisting 01' a loose-textured basal felt , loosely floceose and zanat e
in m argin al a reas, more den sel y lanate in central and subeentral areas , not
furrowed , flat, with cen tre sligh tly umbonate , with very thin margin largely
su b me rge d, extending abou t 5 01' more mm beyond th e aerial growth; heavily
sporing throughout in dccp blue-gray shadcs (S guy, PI. XXXV , 519);

cxudate a nd odou r lnckin g ; no COIOllJ'l:d pigm cnt difTusin g int o lh c SUIToun"

ding agaJ'; co lo n y rev e rso (co lour plat o 152 , e) d e vclop in g ye llow ish ti nls with 11
gree n cas t show ing th rou gh ; p e n icilli cssc n t iall y as d cscrib ed aboye bu t wi th
conid iophores arising mostly Irom th e subs trat u m, lendin g to the colo ny a
dee ply velve ty a p pearance .
The spe cies descriptio n is cen tere d upon th e type stra in CBS 106 .11 = C M I
40 ,224 = ATCC 10,458 = NR RL 2, 009 = [ Fa 6 ,099 = [J FM 7133 , isolat ed by R.
Westling in 1910 from Lecanora sp . , a nd upon other strains isola ted b y DI'.
A .T . M artnez Irom a n a ndo sol under a Fagus silvatica L. fores t , in the spa nish
prov ince 0 1' Navar ra , 1977 .
T he ten dency 01' these strains to produce penicil1i more 0 1' less di varicat c is
suggestive 0 1' a certain relatio ns hi p to th e Asymmetrica -divari cata .

153 . Penicillium echinosporum Ne h ira ex R amre z : fig. 153, co lo u r pl a t..

153, a n d SE M pi ate 26 , (ce ntre: ri ght) .

eQ~q
.G;>oa
.be ' o ;, "
fJ

C)

<&

() Q)

rfY
e

211
L.....-.I

100J.lm

fR.

Fig. 153. Camera lucida

drawings of Penicillium
echinosporum Nehira ex
Ramrez, CBS 344.51. A ,
Detailed drawings of the
penicilli. B, Mature conidia .
C, Habit sketches.

nicillium echinosporum Nehira - J. Ferment . Toc hnol. 11: 861. 1933. See also
M.M. Kulik , 'A Compilation of Descrip tion s of New Penicillium Species ', Agric.
Handbook No. 351. Agric. Res. Ser. U.S. Dep. Agric. p . 36. 1968.

Prnicillium echinosp orum Nehi ra was isolat cd l'r ornJ a pan esc soil in 1932 by T .
Ne h ira , who prop osed (he na me P. echinosporum and described th e species
wiihou t p ro vidin g a La tin di agn osis. In ord er to validat e th e spe cies , a Lati n
d iagn osis 01' it is given below ; th e name 01' th e original a u thor is m a int ained
1I11der R ecornmendation 46 C 01' the Interna tion al Code of Bot anical No m enrl nt u r (Sta f1e u (Ed .) , 1972) .
Larin d iagnosis of Penicillium echinosporum N ehi ra ex R am rez:
Coloniae in agaro Czapekii restrictae crescunt, post duas hebdomadas ad
O-25C, 15 mm in diametro attingunt, denso texto basali coacto mycelio,
velutinae, inaequaliter prominents, radiatae plicatae; cum undulato margine; fortiter
sporulantes coeruleo viridibus colorantur, cum hyphis aeris flavis tectae sunt; odor
abest ; exsudatio adest, exiguus, in mycelio submerso; cum flavo pigmento in medio
diffluentis; reverso rubro fusco ; neque ascomata neque sclerotia visa sunt; penicilli
bivert icillati asymmetrici, cum conidiophoris glabris, brevis , plerumque tenus 50 ~m
in longitudine per 2.0-2.5 ~m in diametro, plurimum e mycelio basali vel e hyphis
reptabundis exoriuntur; rami 15-25 ~m per 2.0 - 3.0 ~ m ; metulae 10-25 ~m per
1.7-2.0 ~ m ; phialides 2-6 in numero per verticillum, 11-17 ~m per 1.8-2.5 ~m,
lanceolatae, cum langa et acum inato collulo; conidia elliptica, 4-6 ~ m per 2.5-4.0
ILm in diametro, echinata, in brevis catenis diversis formantur.
Coloniae in agaro Czapekii cum extracto levedinis, post duas hebdomadas ad
20-25C, 25 mm in diametro attingunt, denso texto basali coacto mycelio,
velutinae, radiatae plicatae , prominentes in centro, cum aequaliter margine; parum
sporulantes, pallido coeruleo colorantur; exsudatio et odor absunt; cum flavo
pigmento in medio diffluentls: reverso aurantio; neque ascomata neque sclerotia
visa sunt.
Coloniae in agaro hordeaceo, post duas hebdomadas ad 20-25C , 25 mm in
diametro attingunt, laxo texto basali coacto mycelio, lanatae, concavae in centro,
parum prominentes in margine ; e reticulo floccosis albis hyphis aeris tectae sunt;
parum sporulantes; odor et exsudatio absunt; sine pigmento diffluente in medio;
reverso aurantio. Neque ascomata neque scierotia visa sunt.
Typus cultura CSS 344.51 e solo Japonico a T. Nehira, anno 1932 isolatus esto
Nehira 's original diagnosis follows:
'The mycelium is yellow-green upon Czapek's agar and the margin is white and 2
mm wide . After 7 days, colonies attaining 10 mm in diameter. The white marg in disappears in age and the colony surface becomes dirty dark green coloured ; the
colony reverse is bright yellow, but sometimes developing brown colours with

'1'""
yellow-brown zonation in age. At 33 lJC, the colonias become very dark green In
colour with the reverse yellow to purple brown. No growth occurs at 37C .
When viewed under the microscope, some ropes of hyphae may be observed ,
2-3 .tm in diameter.
Penicilli biverticillate asymmetric , 21 to 33 .tm; metulae 8 to 9 .tm; phialides 10 to
12 .tm. Conidia elliptical, 2 to 3 .tm , with pointed ends; some conidia are globose , 2
to 2.5 .tm; with walls roughened and echinulate .'
(Translated from the Japanese by Miss Masae Yamamoto).

Our notes follow :

Colonies upon CA (colour plate 153 , A and a) growing restrictedly,
attaining a diameter of 15 mm in 14 da ys at room temperature (20-25 C),
consistin g of a tough , close-textured basal felt , finel y velvet y in appearancc ,
irregula rly raised towards the ce nt re , with few shallow radial fur rows, with
thi n undulating margin entire , a t first white, th en becoming blu e-green with
th e rip ening of con id ia, with veg et ative hyphae yellow -coloured; medi u m
sp oring in blu e-green shades (Sguy , XXXIII, 484), sta rting near the margin,
wit h non- sporulating areas remaining yellow; colony surface overlaid by a
dense network of aerial hyphae lending a sublanate appearance to th e colony;
odour lackin g; exudate lim ited in amou nt , as very minute colourless droplets
en mes hed in the mycelial felt ; yellow -col oured pigm ent diffusing into thc
surrounding a ga r; colony reverse (colour plat e 153 , a) in dark redd ish brown
shades; pen icilli (fig . 153, A) typically biverticillate asymmetrical , with slender
conidiophores arising from th e basal felt, very short , usuall y less than 50 .tm in
len gth by 2.0 -2 .5 11m in diameter , irregu larly arranged with rami abo u t 15-25
11m by 2.0- 3 .0 11m , no more than two elem ents in th e ve rticil; metulae variable
in len gth in th e verticil, ranging from 10 to 25 11m by 1. 7- 2.0 11m , n umberi ng
2- 3 elem ents in the verticil ; phialides typi cally la nceola te , sugge stive of the
Biverticillata -vsymmetri ca secti on, in sm all clusters of 2-6 ele m ents in the
verticil, 11- 17 11m by 1.8 -2 .5 11m, taper in g to a long narrowed neck ; con id ia
elliptical, 4 -6 11m by 2.5 -4 .0 11m in di amet er, cons picuously ech inulate (fig.
153, B, and SEM plate 26 , ce ntre : right) ; con id ial cha ins short , loosely
div ergent (fig . 153, C) .
Colonies u pon CYA (colour pIat e 153, B and b) attaining a di ameter of
about 25 mm in 14 da ys at ro om temperature (20 -25 C) , con sisting of a
sim ila rly textured basal felt as described aboye , but more deep ly floccose ow ing
to the de ep er network of white aerial hyph ae overlayin g th e basal felt ; deep ly
radiately furrowed in marginal and submargin al area s, with centre umbonate
and m argin grad ually rai sed tow ards th e centre; wit h undulating even m argin
entire; po orl y spor ing in very pale blue shades in margin al areas , elsewhere
remaining persist ently white ; exudate a nd od our lacking; yellow-coloure d
pigment diffusing int o th e surrounding ag ar; colony reverse (colour plat e 153,

h) dev elopin g o ran gc sh ad cs: pcni cilli t:sscntiall y as dcscribcd a bo ye .

Colonies upon M EA (colour plat e 15:1, C a m i c) a uain ing a di amct er 01'
:11 )()1I ( 25 mm in 14 days at roo m tcmpcratu re (20 - 25 C ), cons isting 01' a looser
tcxt 1I 1'ed basal felt overlaid by a deepl y floccose netw ork 01' white aeria l hyphae ,
lend ing a lanat e appearance to the colony su rfaee ; thin even m argin a b ruptly
ru iscd and ce nt re eoncave ; colon y surfaee not furro wed; poorly sporing, not
influcnci ng th e colon y appearanee , whi eh remains persistently white; no
m [ou re d pigmen t di ffusin g into th e su rro u nd ing agar; eolony reverse (eolou r
plat c 153 , e) d eveloping yellow-o ra nge shades; penieilli esse ntially as described
a boye .
T he spee ies deseriptio n is eentered up on th e typ e stra in C BS 314.51 = IJ FM
7 1:1 1, isolat ed b y T. N ehira in 1932 from J a panese soil. It is kn own only fro m
1he type stra in.
O u r obse rva tions, in gene ral, eonfor m with th e author's original d iagnos is
() I' the spec ies, with a few di ser ep aneies, part ieularly th e presen ee 01' ropes 01'
hyphae and the eo m paratively smalle r eon id ia rep orted b y Ne hira .
The eo rree t plaeemen t 01' Penicillium echinosporum Ne hira is still ope n to
qu cstion . The rather velvety as peet upon C A and C YA and th e d eepl y floeeose
"ppearanee 01' th e eolonies upon M EA , a re suggesti ve 01' some relation sh ip
with bo th th e Asymme triea - velu tina a nd th e Asymmetriea-Ianat a su bscct ions, but , on th e othe r hand , the typieall y laneeolat e phiali des are
sugge stive 01' th e Biverti cillat a-symmetri ea see tio n as understood by R aper and
Thorn ( 1949).

154 . Penicillium giganleum R oy el Sin gh : fig. 154, coIour pIat e 154, and
SE M plat e 26 , (bottom: Ieft) .
Penicil/ium giganteum Roy et Singh - Trans Brit. Mycol. Soco 51: 805-806. 1968.
Roy and Singh's original diagnosis follows:
'Colonies growing well on Czapek's agar, attaining a diameter of 11.5 mm in a
week at 25C, dark brown; reverse yellowish brown, slightly floccose and wrinkled;
hyphae branched, delicate, mostly 2-3 /Lm broad; conidiophores fairly long, arising
as lateral branches from the main hyphae, often simple but sometimes branched,
septate, 2-4 /Lm wide, smooth walled; penicilli irregular, varying from
monoverticillate with 1-4 phialides at the apex of a short conidiophore to partially
biverticillate or fairly regularly biverticillate, but not typical of the section
Biverticillata-symmetrica; metulae 2-6 in a verticil, 7-15 by 2.5-4 /Lm; phialides
divergent with swollen base, tapering abruptly to a long , slender tip , 6-20 by 1.5-3
.m . Conidia globose, in chains with conspicuous connectives, at first olive green
becoming dark brown at maturity, with prominent spines, generally 6-9 /Lm,
occasionally up to 12 /Lm in diameter, connectives 1.5-2.5 .m long, occasionally 4
/L m .

.\ \

c~'~
, : ., :.

~, I!."
, ..-

"

'-

.' i

~,
;~. .
.. .

l-.....-.J

!i.

100jJm

Fig. 154. Camera lucida

drawings af Penicilliu m
giganteum Ray et Singh ,
CSS 144.69. A , Detailed
drawings of the penicilli. S,
Mature canidia. C, Habit
sketches . D, Cailed hyp hae.

The fungus f irst appeared on Martin's rose-Bengal med ium as a small blackish
colony which was later stud ied on Czapek's medium w ith 3% and 20% sucrose,
and on 2 % malt agar. It grew satisfactorily on these med ia but the growth was
maximum on Czapek's medium w ith 20% sucrose.'

Our notes follow:

Colonies upon CA (colour plat e 154 , A a n d a) growing rather restrictedly,
a ttain in g a diameter of 25 mm in 14 days a t ro om temperature (20 -25 C) ,
co nsisting of a tough, close -textu red basal felt ov erlaid by a deep n etwork of
f1occose white aerial vegetative hyph ae influencin g greatly th e colo ny
appearance; colony surface de epl y velvety to la n a te , colou re d in very pale
yellowish shades, with eve n thin m argin surrounded by a fringe of submerged
hyphae ex te nd in g about 2 mm beyond th e aerial growth ; very poorly spor in g ,
not influen cing the colo ny a p pe a ra nce ; odour lacking ; exu d a te lim ited in
a mou nt, as minute colou rless droplets enmeshed in th e mycelial felt ; pale
yellowish-colo ured p igment d iffu sin g into th e surrou ndin g aga r; col on y reverse
(colour plat e 154 , a) deve loping pale ye llow to orange shade s ; penicilli (fig.
154 , A) va r iable in co m plexity , com mon ly co n sisting of ve ry short con id io-

phores , up to :)() Itlll in length by :~.() -4.() Itlll in diumct cr, usunlly smooth
wnllcd , but arising roin trailillg hyphac more 01' less co nspicuo usly granular,
Iwarillg al th cir tips small verti cils 01' m crulac, commonly 2-3 clements in thc
vcrricil, 10-15 P.nl by :~ .0 -4 .() tui, cac h bcaring a small cluster of 2-3
phinlidcs per whorl , bottle shaped , 8-13 tuu by 3.0 -4.5 tui, tapering abruptly
11' l co nspicuously long narrowed neck; occasiona1ly simple penici1li
(al'pearing monoverticillate) are also observed, and metulae arising at different
Ie 'vds 01' the conidiophore are not infrequently encountered; conidia compara1ivcly very large, probably exceeding in diameter that of a1l known Penicillium
xpcc ics , globose, strongly echinulate (fig . 154, B, and SEM plate 26, bottom:
11'1"), about 6-10 p.m 01' occasionally up to 12 ust: in diameter; conidial chains
short , loosely divergent, with conspicuous long connectives between the conidia
(Iig . 154, C); numerous coiled hyphae, not reported by previous authors have
Iicen observed in the present study (fig . 154, D), suggestive either of asco mata
i 1I i 1ials 01' nematodes traps .
C olonies upon CYA (colour plate 154, B and b) attaining a diameter of
:d)()ut 30 mm in 14 days at room temperature (2-25 C), consisting of a
lough , c1ose-textured basal felt, similarly textured as aboye but radiately
lurrowed , appearing deeply velvety to lanate, with thin even margin
surrounded by a fringe of submerged hyphae extending about 2 mm beyond
I he aerial growth; poorly sporing, with colony surface remaining persistently
white to yellowish coloured; odour and exudate lacking; pale yellow-coloured
pigment diffusing into the surrounding agar; colony reverse (colour plate 154,
h) developing orange shades; penicilli essentially as described aboye .
Colonies upon MEA (colour plate 154, C and c) attaining a diameter of 30
111m in 14 days at room temperature (20- 25C) , consisting of a loo ser textured
basal felt, deeply velvety, plane, not furrowed, with wide thin margin
surrounded by a fringe of submerged hyphae extending about 5 mm beyond
the aerial growth ; very heavily sporing in central and subcentral areas in deep
mouse gray shades, with the conidiophores arising in a dense stand from a
closely woven network of trailing vegetative hyphae, with walls conspicuously
granular; exudate limited in amount, as minute colourless droplets enmeshed
in the mycelial felt; odour lacking; reverse of colony (colour plate 154, c)
developing yellow to orange shades with a green cast showing through; no
coloured pigment diffusing into the surrounding agar.
The species description is centered upon the type strain CBS 144 .69 = CMI
132,774 = IJFM 7149, isolated in November 1967 by C .N . Singh from soil at
the Botanical garden of the Banaras Hindu University, Varanasi-5, India .
Other strains were isolated simultaneously from the rhizosphere of Brassica
campestris L., at the same locality .
Penicillium giganteum Roy et Singh is a quite distinctive species characterized
particularly by its giant, strongly echinulate conidia , hence the specific epithet .

436

It a ppears lo be IllO S( sa tisfac rorily ass ig na b lc lo th e P. comrnune seri es on

accou n t 01' its floccose colo n ies , but no close rel ati on ship has been found
between P. giganteum a nd th e other spec ies included in th e ser ies . Howev er,
th ey are co ns idered together as a m atter 01' con ve n ience .

haptcr 7

Asym m ctr ica-funiculosa sub-section

TIIE SP ECIES CONSIDE RED I N T H IS SUB -SECTION , AS UNDERSTOOD BY R A P ER AND

' I'horn ( 1949), p roduce colon ies ranging from definitely floccose to almost
vclvcty , bu t typi call y showing pa rt of the aer ial hyphae combin ed in to
.uu istomosin g ropes or fu ni clcs; with con id iophores partly ar ising from
sub rnerged hyphae , partly as terminal portions of ascen d ing ae r ial hyphae ,
.uul in part as branch es fro m th e ro pes of ae r ial hyphae , but n ever forming
rrcct fascicles or syn ne mata; with peni cil1i cons iste n tly a symmetrical , u sually
I wo- or three-stage branch ed .
O f th c two recognized ser ies (Rape r a nd T hom, 1949), n amely Penicill ium
trrrestre ser ies and P. pallidum series, only th e latter h as been retained in
lite present stu dy, since al1 sp ecies incl ude d by Raper a nd Thom (194 9) in th e
ormer have ac tual1y been re d uce d to sy no ny m ies wit h oth er for ms incl u de d in
t hc Asymmetrica-fascicul at a Subsection (Samson, Stolk, a nd H adlok , 1976) .
T he members of thc Penicillium pa llidum series do not a ppea r to b ear a clos e
relat ion ship to a ny recogn ized ser ies. They seem to represent a gr oup wel1
a pa r t from al1 othe rs.

Key to species
Page

1. Conidial areas variously coloured, not in green shades; penicilli often

comparatively narrow with cellular elements laterally appressed; walls of
the conidiophores, metulae (and often the phialides) closely and
conspicuously roughened.
P. pa/lidum series
A . Conidia white to cream coloured.
1. Conidial chains divergent, becoming tangled in age.
P. pa/lidum Smith
2. Conidial chains in well-defined columns.
P. putteri/lii Thom
B. Conidia in light to dull gray shades .
P. namyslowskiiZaleski
C. Conidia in Iight violet, lavender, or vinaceous shades.
P. lavendulum Raper et Fennell

. . . . 438

.. . . 438
.... 440
. . . . 442
. ... 444

Pen icillium pallidum seri es

UTSTANDING FEAT UR ES

Colonies growing rather restrictedly upon Czapek's agar bu! broadly

spread ing upon Czap ek-yeast extract agar; usuall y rather thin, with su rface
growth somewhat funi culose ; conidial ar eas variabl e in colour , white to crcam,
buff, light gray , 01' lav ender to vinaceous, nev er developing true gree n shadc s,
C onidiophores com pa ratively short, generall y a rising from ropes of acrial
hyphae; cons p icuou sly septa te , with wall s of con id iopho res, including rami ,
metulae , a nd even phialides usu ally cons picuo us ly roughened .
P eni cilli usually com paratively narrow, with all elements tending to be
closely appressed .
Phialides arran ged in com pac t clu sters, closely parallel , com para tivcly
narrow, seldom exceeding 3. 0 p,m in diamet er, with walls usually echinulate.
C onidia strongly elliptical to cylindrica l, smo oth wall ed , ofte n ad he ring in
long cha in s.
Irresp ecti ve of th e true re la tions hip th at m ay exist betw een th e spccies
included in this series, all of th em , however , possess ce rta in rnarked
cha racteristics in com mo n whi ch enable th em to be cons ide re d together m or e
sa tisfacto rily than with a ny othe r of th e R aper and Thom ' s ( 1949) recognizcd
senes.

155 . Penicillium pallidum Smith : fig . 155 , co Io ur pIate 15 5 , and SEM

pl ate 26, (bottom: right) .
Synonymy
Geosmithia putteri/li (Thorn) Pitt - Can. J . Bot. 57: 2021-2030. 1979.
Penicillium pallidum Smith - Trans . Brit. Mycol. Soco 18: 88-89. 1933. Also Rapar
and Thom, 'A Manual of the Penicillia', p. 461.1949.

Colon ies upon C A (colour plat e 155, A and a) growing restrictedly,

a tt aining a di ameter of 25 mm in 14 days at room tem pe rature (20- 25 OC ),
consisting of a thin, loose-t extured ba sal felt, with margin largely submerged,
exte nd ing abo u t 5 mm beyond th e ae rial gro wt h, plane ; m ycelial gro wth
giving rise to a spars e dev elopment of con id ial str uc tures and sho rt funi culose
tufts 01' rop e of aerial hyphae bearing con id iopho re s; colony surface a t first
white , becoming cre a m coloure d in age; odo u r a nd exudate lacking ; no
coloure d pigment diffu sin g into th e surro u nding agar; colony rcverse (colour
plat e 155, a) colou rless 01' in p ale cre am sha des; penicilli asym metr ic a nd
bi verticillat e (fig . 155 , A) , with co nid iopho res a rising from trailing hyphae 01'
from ropes of h yphae, many borne from foo t cells as obse rve d in th e genus

Pen icillium [mllidum se ries

OUTSTANDING FEATU RES

Colonies growing rather restrictedly upon Czapek ' s agar but broadly
spread ing upon Cz apek-ye ast extrac t agar; usually rather thin, with su rface
growth somewhat funi culose; con id ial a reas variable in colou r, wh ite lo crca m ,
bu ff, light gray, or lav ender to vinaceous, never d eveloping true gree n shadcs ,
C onidiophores co m parative ly short , ge ne rally arising from ropes of acri al
hyphae ; con spicuo usly sep ta te , with wall s of con id iop ho res, includin g rami ,
m etulae , an d even phialides usuall y cons picuo usly rou ghen ed .
Penicilli u sually com paratively narrow, with all elements tending to be
closely ap pressed .
Phialides ar ra nged in com pact clu sters , closely parallel , com para tivcly
narrow , seldom exceeding 3.0 p.m in diamet er, with walls usually echinulatc.
C onidia stro ngly ellipt ica l to cylindrical, smo oth wall ed , oft en adherin g in
long cha in s.
Irrespecti ve of th e true rel ation ship th at m ay ex ist between the spec ies
included in this series, all of th em, however , possess ce rtain m arked
chara cte r istics in co m mo n whi ch enable th em to be cons ide re d together more
satisfacto rily th an with any othe r of th e R ap er a nd T ho m ' s (1949) recognizcd
senes.

155 . Penicillium pallidum S m ith: fig. 155 , co lo ur plate 155 , and SEM
plat e 26, (bottom : right) .
Synonymy
Geosmithia putterilli (Thom) Pitt - Can. J . Bot. 57: 2021-2030. 1979.
Penicillium pallidum Smith - Trans. Brit . Mycol. Soco 18: 88-89. 1933. Also Rapar
and Thom, 'A Manual of the Penicillia', p. 461. 1949.
Colon ies upon C A (co lou r pl at e 155 , A and a) gro wing restrict edly,
attaining a di amet er of 25 mm in 14 days at room te m pe rature (20 -2 5 OC ),
co nsisting of a thin, loos e-textured ba sal felt , with margin largely sub m erged ,
extend ing a bo u t 5 mm beyond th e ae rial growth , plane ; m yceli al growth
giv ing ris e to a sp arse devel opment of coni d ial structures and short funiculose
tu fts or rop e of ae rial hyphae bearing con id iopho res ; colo ny su rface at first
white , becoming cream colou re d in age; odo u r and ex udate lacking ; no
colou re d pi gment diffu sing into th e su rro u nd ing agar ; colony reverse (co lou r
plat e 155, a) colou rless or in pale cream shades ; penicilli asym metric and
biverticillat e (fig . 155 , A), with co nid iopho res ar ising from tr ailing h yphae or
from rop es of h yph ae , m any borne from foo t cells as obse rved in th e gen us

-,
",'

/f
'

<:~t e Jf7':

;~

'l
L.-J

100J.lm

\.' '; '~i'~ :<: '

)/

" <'

f.K..

Fig. 155. Camera lucida

drawings of Penicillium

pallidum Smith, CSS 248.32.

A, Detailed drawings of the
penicilli. B, Mature conidia.
C. Habit sketches.

Aspergillus; conidiophore walls cons picuou sly roughen ed, with metulae , rami ,
a nd phi alides similarly marked; m etulae in groups of 2- 3 elements in th e
verticil, a bou t 9-14 /lm by 2. 5- 3.0 /lm ; phialides with walls con spicuo us ly
g-ra nular, narrow, taperin g grad ually to a narrowed short neck , 8-1 5 /lm by
:.! .5 - 2 .8 /l m; conidia cylind rica l, 3.5 - 4 /l m by 1.8-2.0/lm , sm ooth walled
(Iig . 155, B, and SEM plat e 26, bottom ; right) ; con id ial chain s tending to for m
loosel y parallel colu m ns (fig . 155, C) .
Colonies upon CYA (colour plat e 155, B and b) spreading broadly , attain ing
a diamet er of a bou t 45 mm in 14 d ays a t room temper ature (20-25 OC ),
consisting of a tough , close-textured basal felt, strongly radiately wrinkled , a nd
slightly raised in th e ce ntre ; a ppearing powdery to chalky, with a thin eve n
margin surrounded by a frin ge of su b me rged hyphae ex te nd ing abo u t 2 mm
beyond th e aerial gro wth; heavil y sporing throughout in pal e cream sha des ;
odo u r and exud a te lackin g; pal e a mbe r-coloured pi gm en t diffu sin g into th e
su rrou nd ing agar ; colony revers e (colour plate 155, b) devel oping ora nge
shades; penicilli esse n tially as d escribed aboye , with co nid iop ho res usu all y
short, com monly two-stage branched , not exc eeding 175 tui: in len gth by
2.0- 3.0 /lm in di amet er , arising primarily from the basal felt a nd from ro pes of
ae rial hyphae.
C oloni es upon MEA (co lour plat e 155, C and c) attain ing a diamet er of
abo u t 45 mm in 14 days a t ro om temperature (20 -25 OC ) , con sisti ng of a

't'I'\J

looscr tex tu red ba sal fclt, co ns picuo usly funi cul ose all ovcr thc colo ny surfa cc ,
flat , not furro wed , slightly urnbon at c in the centre , with thin marg in
su rro u nde d by a fringe of subme rged hyphac cxtc nding abo ut 5 mm bcyond
the ae rial growth ; heavil y spor ing throughout in Naple yellow shades (Sguy ,
PI. XIII , 190), wit h conidiop ho res short, a rising mostly from ro pes 01' hyph ac ,
giving to th e colony surface a chalky appearance; odo u r and exudate lackin g;
pale yellow coloured pigment di ffus ing into th e su rrou nding agar; colony
reve rse (colour plat e 155, e) developing pale orange shades; pe n icilli esse ntially
as described .
T he species desc ription is centered up on ou r notes on Sm ith's type strain c as
260.33 = C M I 40,214 = ATCC 10,4 78 = NRRL 2,037 = I Fa 5, 758 = QM 1958,
isolated fro m cotto n ya ro, Eng land, 1931, a nd upon strain C BS 248 .32 = N R RL
2,156 = IJFM 7118, isolat ed in 1932 in H olland fro m an elm beetl e (Scolytus sp. ) ;
the latter strain prob abl y suffers a vitamin deficien cy , clearly shown by the
very res tr icted growth upon CA (colour plate 155, A) and the rapid growth
upon CYA (co lour piate 155, B) . H owever , th e typ e strain 01' Penicillium
pallidum, as noted in Smith's original diagn osis (1933) is characterized by
broadly spreading colonies upon CA, attaining a diamet er 01' 65- 70 m m in 14
days at room te m peratu re, showing numerous radial furrows, bei ng simi la r in
texture and cultural appearance to strain C BS 248.32 cultivated upon CYA
(colour pla te 155, B) .

156 . Penicillium putterilli T horn : fig. 156 , co lo u r pl ate 156 , a nd SEM

pl ate 27, (top : left) .
Penicillium putterilli Thom - The Penicillia', pp. 368-369. 1930. Also Raper and
Thorn , ' A Manual of the Penicillia', pp. 461-462. 1949.
Synonym
Geosmithia putterilli (Thorn) Pitt - Can. J. Bot. 57: 2021 -2030. 1979.

Co lonies upon CA (colou r plate 156, A and a) attaining a diameter ofabout

35 mm in 14 days at room temperat ure (20 -25 OC), con sisti ng 01' a toug h ,
close -textured basal felt, at first largely submerged a nd appearing wet, wit h
thin margin la rgely submerged extending about 3 mm beyond th e aerial
growth, later appearing velvety, with numerous ropes al' hyphae all ayer the
colony surface, which is plane a nd slightly raised towards the centre; me dium
sporing throughout in Naple yellow shades (Sguy , Pis. X II, 190, and XIV ,
200); odour a nd exudate lackin g; no coloured pigme nt diffus ing in to the
surrounding agar; colony reverse (colou r piate 156, a) at first colou rless,
develop ing pale yellow shades; peni cilli typically asymmetrical, one - 0 1' twostage branched, with all elements closely appressed and cons picuously warty

"o
o
0 0

i.w.WwJ

jo ,

10 prn

' o

o
00

trYJ}
~

100 pm

oiJO () ~> o
(J ~ O ~
O

~ C) ~o
O

~ .>

R..

c:;{:J

Fig. 156. Camera lucida

draw ings of Penicillium
putterilli Thom , CSS 233.38.
A , Detailed drawings of the
penicilli. S, Mature conid ia.
C, Habit sketches.

(from the stipe to the phialides) (fig . 156, A); conidiophores mostly arising
Irorn ropes of hyphae, commonly short, less than 100 11m by 2.5- 3.5 11m ;
rami , in small groups of one or two elements in the verticil, about 15-25 11m
by 3.0- 3.5 11m; metulae n umbering 2-3 elements in the verticil, 12-1 8 11m by
2 .5- 3 11m ; phialides in closely appressed who rls, numbering 5- 10 elements in
the verticil, cylind rical, conspicuously wa rty, 9.0-15 .0 11 m by 1.5- 3.0 11m ;
con id ia cylin d rical, 4.0-6.5 11 m by 1.5-2.5 11m , smooth wa lled (fig . 156, E,
and SEM plate 27, top : left); conid ial chains forming fairly well-d efin ed
colu m ns (fig. 156, C).
Colonies upon CYA (colour plate 156, B a nd b) spread ing broadly, attaining
a diamet er of about 60 mm in 14 days at room temperature (20-25 OC ),
consisting of a thi n, loose-textured basal fe1t, radiately furrowed and raised in
the centre in a crateriform pattern, wit h very thin margin surrounded by a
wid e frin ge of submerged hyphae extend ing about 4 mm beyond th e aerial
growth , with growing surface ov erlaid by numerous ropes of aerial hyphae
lending a powdery appearance to the colony ; cons picu ou sly zonate; heavily
sporing throughout in Naple yellow shades ; exudate and odour lacking ; no

colou red pi gm cnt difu sin g' inro th c su rro u nd ing aga r ; colo ny rev e rso stron gly
pulling th c agar away rom th e cu ltu re pl a te , a llow ing good sporu la tio n
underneath in a reas where th e a ir has becn a vaila b lc th rou gh th c a gar cracks
(colour plate 156, b) ; penicilli esse ntia lly as des cribed aboye.
Colo n ies upon M EA (col our plat e 156, C a nd c) growing r ather restrict edly ,
a ttain ing a di amet er 01' a bou t 30 mm in 14 da ys a t room te rnpe rature
(20 - 25 OC ) , co ns isting 01' a thin , loose-textured ba sal felt , pl ane , za nate , not
fu rrow ed , with a powdery a p pearance, with a ye ry thin m argin surro u nde d by
a frin ge 01' su b merged h yphae ex te nd ing a bou t 2 mm beyond the a erial
gro wth , with th e funiculose habit re d uced 0 1' la ckin g ; heavil y sp orin g
th ro ughou t in N aple ye llo w shades , wi th co n id io pho res m ostl y a r ising from the
su b st ratu m in a dense stand , wi th co ni d ia l chains producing fairly well-defined
colu m n s up to 200 /lm long ; odou r a n d ex u date lacking ; no co lo ured pi gment
di ffu sing into th e su r ro u nd ing agar; colo ny reverse (co lo u r pl at e 156, c)
develop ing ye llo w-o ra nge sh ades .
T he spec ies description is ce nte red upon strain C BS 233 .38 = C M )
40 ,212 = NR RL 2,024 = IJFM 7182 , isolated b y J .C. N eill in N ew Zeal and , from
Beilschmiedia tawa-discoloured timber. This cu ltu re a ppare ntly differs from th e
or iginal ty pe strain describ ed by Thom , in 1930 , from a cu lture se nt to him by
F. M . Putterill , Cape T own , So u th Afri ca , a nd quickl y lost from hi s collection,
only in producing co n id ia that a re m o re co ns iste nt ly wh ite , h ence a p proach in g
Penicillium pallidum in cu lt ural a p peara nce.

157. Penicillium namy slowskii Zaleski : fig. 157 , colour pIa te 157 , and
SEM plate 27, (top : right).
Penicil/ium namyslowskiiZaleski - Bull Acad. Polonaise Sci . Math. Nat . Sr. B. pp.

479-480. 1927. See Thom, 'The Penicillia', pp. 484-485. 1930; also Raper and
Thom, 'A Manual of the Penicillia', pp. 462-464. 1949.

Synonym
Geosmithia namyslowskii (Zaleski) Pitt - Can. J. Bot . 57: 2021-2030. 1979.

Colo n ies upon C A (colour pi a te 157, A) gro w ing ye ry restrict edly , a ttain ing
a di amet er 01' 15 mm in 14 da ys at room temperature (20- 25 OC ) , co ns ist ing 01'
a very thin b a sal fel t , wholl y su b merged, almost transp arent , producing
limited con id ia l structures , m ostl y near th e colony ce ntre, dirty white to
eolou rless; odour stro ng, ear thy; ex udate la cking ; co n id ia l str uc tu re s sparse ly
p ro d uce d, w ith co ni d iophore s ge nerally arisin g fro m su b merg ed h yphae , ereet,
com pa r a tively co arse, with walls co ns picuo u sly a n d coarsely ro u ghe ne d (from
th e stipes to th e phialides in clusiv e) .
C olon ies upon C YA (colour pl ate 157 , B a nd b ) growing lu xuriantl y a n d

1':'
, '

.'.. ~

"

.:

r..,"

,~Ilf:i

l'!
,

-:

"

.,"1 m"t~ '::

'.

:~. '.t -.

~o
o QO
00

..

::::.~
" ,
",

..

' ,'

'

: .:
.' : "
:'.:

::.:

G()O

OO2f
0

g~o 11[')
(

100~m

(J{.
Fig. 157. Camera lucida
drawings of Penicillium
namyslowski Zaleski, CBS
353.48. A, Detailed draw ings
of the penicilli. B, Mature
conidia. C, Habit sketches.

broadly sp read ing, attaini ng a diame ter of 65 mm in 14 days a t roo m

temperature (20 - 25 OC), sho wing a clear vita m in deficien cy ; consis ting of a
tou gh , close-textured basal felt, with flat thin m argin surrou nded by a fringe of
sub me rge d h yphae ext ending widely up to 10 mm beyond th e aeria l gr owth,
with su b marginal area radi at ely and deepl y furro wed , a nd with cen tral area
irr egul arl y folde d a nd sho wing a fu n iculose appearance , a ppcaring powdery in
thc m argin, wet in su b marg inal zo nes; heavily spo ring particul arl y in m argin al
areas in gray-gree n shades (Sgu y , PI. XXIX , 427 ); penicilli (fig. 157 , A)
typically one -stage branched , co mmonly showin g a terminal clu ster of
metul ae , but sometimes ramified , u su ally asym me tric in pat tern ; m etulae in
grou ps of 2-5 elements in the verticil, about 15-25 !-tm by 3.5 - 4. 0 tui: with
wall s con spicuous ly wa rty , and a pices some what enlarged up to 5 - 7 !-tm in
d iamet er , borne upon coarse con id iopho re s, usu all y short , ranging from 60 to
125!-tm by 3 .5 -4.5 !-tm , erect a nd with walls co ns picuo us ly warty , ge nerally
arising fro m submerged hyphae ; phialides in small clus te rs of 2-6 elements in
the verticil, closely a pp ressed, 12-1 4!-tm by 4.0 uit, with wa lls cons picuo usly
wa rty, tapering grad ually to a co ns picuously sho rt narrowed neck ; co nid ia at

11'1

first cylind rical, becomin g lat cr cllipt ica l , :~ .5 -5 . 0 by 2.B-:1. 5 J.tlll in d iamc tc r,
with waIls smooth (lig. 157, B, an d SE M plato 27, (0)): righ t); odo ur stron g,
sugges tive of the Beta uulgaris L. a ro ma; ex udare lim itcd in a mou n t , as minute
colou rles s dropl ets en mes hed in th e m yceli al felt ; no colou rcd pigm ent
d iffus ing into th e surro u nd ing agar; colon y rever se (co lour pl at e 157, b)
developing oc h rc shadcs.
Colon ies upon M EA (co lour pIate 157, C a nd c) spread ing broadly,
atta ining a di amet er of 60 mm in 14 days at ro om te m perature (20- 25 OC ),
cons isting of a ver y thin , loose-textured basal feIt , flat , not wrinkled except at
th e colony cen tre whi ch is irregul arl y folded and ra ised in a somewha t
cra te riform pattern , with very thin marg in largely su bmerge d, extc ndi ng
5 m m 01' m ore beyond the aeria l growth; heavil y spo ring th ro ugho u t in the
sa me gray-gree n shades as aboye (Sguy, PI. X X IX, 427); odour stron g , as
aboye; ex udate lackin g; no colou re d pi gm ent diffusin g into the surrou nd ing
agar ; colon y reverse (colo u r pl at e 157 , c) developing yeIlowis h to oc h raceous
shades with a p ale gree n cas t sho wing throu gh ; penicilli esse n tia Ily as described
aboye, with co nid ial chains forming fairly we Il-de line d colum ns (lig . 157 , C) .
The species desc rip tion is ce nte re d upon the strain CBS 353 .48 = N R RL
1,070 = CM I 40,033 = ATC C 11,127 = Q M 1,932 = IJ F M 7202, whic h is believed to
be Zales ki' s type strain, isolat ed fro m Polish soil u nder co nifcrs in 1926.
The a boye description is in fairly goo d agreeme n t with Zales ki's orig inal
di agn osis excep t for th e cons picuously roughe ne d coni d iophores (Zales ki
re po rted and iIlust rated aIl walls as smooth). In study ing th e a bo ye strain,
Thom ( 1930), and R aper and Thom ( 1949) rep orted con id iophore wall s as
ro ug h. The culture received by us from the Cen traalb u reau voo r Schimme lcultu re s, Baarn , T he N etherl ands, re ta ins th e characte r observed b y T ho m
and is believed to re present Zaleski' s original isolat e despite the re po r ted
differ en ces in th e .character of th e con idiopho re waIls.
158 . Penicillium lavendulum R aper el Fennell: fig. 158 , colour pl at e 158 ,
a nd SEM plat e 27, (ce n tre : left) .
Penicillium lavendulum Raper et Fennell - Mycologia. 40: 530-533. 1948. See also
Raper and Thom, 'A Manual of the Penicillia', pp. 464-465. 1949.

Synonymy
Geosmithia lavendula (Raper et Fennell) Pitt - Can. J. Bol. 57: 2021-2030. 1979.
C olonies upon C A (co lou r plate 158, A and a) attaining a di amet er ofabo ut
40 mm in 14 d ays at room temper ature (20 - 25 OC ), co ns isti ng of a loosetex tured b as al felt, plane 0 1' nearly so, comparatively thin , with irregul arl y
d issect ed m argin surrou nded by a fringe of su b merged hyphae extend ing abo u t

'r' r d

00

\:jo6

"

"
::.:

\j
'O sO (:)

68o

Fig. 158. Camera lucida

drawings of Penicillium
lavendulum Raper et Fennell,
CSS 344.48. A, Detailed
drawings of the penicilli. S,
Mature conidia .

5 mm beyond th e ae rial growth, with colony su rface ove rlaid by a loose

ne twork of floccul ent to cot to ny hyphae , showing sorne ro piness in m ar ginal
and submarginal area s; heavily spo ring throu gh ou t in violet shades near
laven der (Sg uy, PI. 1, 6, 11); odo ur a nd exu da te lacking; pu rpl e-coloured
pigme nt diffusing in to th e su rround ing agar; colony reverse (colou r plat e
158 , a) developing purple to deep lav en der shades (Sguy, PI. 1, 11) ; penicilli
typi cal1y asym me tr ical (fig. 158 , A) , va riable in size, with conid iophores
a rising from th e su bstratu m, but m or e frequ ently bo rne primarily as branch es
from aerial h yphae, com m only 100-1 50/lm in len gth by 3.0-4 .0 /lm in
di ameter , cons picuo us ly septa te , with walls eithe r closely echin ula te or sm ooth
walled; penicill i very irregul ar, one - or two -stage branched , with m etulae
com mo nly ar ising at different levels within the penicil1us; rami , va ry ing greatIy
in len gth , 15- 40/lm by 3.0- 4.0 tui , cons picuo us ly septa te ; m etulae about
8- 11 /lm by 3.5-4. 0 /lm, u su all y smooth wall ed ; phialides in closely
appressed clu sters, vari ab le in shape and size, m ostly cylind rical, 11-14 /lm by
2.5 - 4.0 /lm , abruptly tapered to an inconspi cu ou s neck , usual1y smooth
wa lled; con id ia stro ngly ellipt ica l to cylindr ical, 5.0-6 .5 pst: by 2.5- 3.5 /lm in

146

diameter , with wall s srnooth (Iig . 1:>8, B, a nd SEM plat e 27, ce ntre; lcft);
con id ial chains tending to adher e int o loose colu m ns,
C olonies upon C YA (colour plate 158, B a nd b) spread ing broadly , attainin
a di am eter of 55 mm in 14 days at room temperature (20 - 25 OC), co ns isting 01"
a close-t extured basal felt , strongly funi cul ose , com pa ra tively thin , radiatc1y
furrowed a nd somewhat rai sed in th e cen tre , with thin eve n m argin
surrou nde d by a frin ge of su b me rge d hyphae ex te nd ing abou t 3 mm beyond
the aerial growth; poorly sporing in dirty white shades, never showing
lav ender colou rs; odo ur lacking ; exudate limited in am ou nt , as colou rless
d ropl ets enme she d in th e my celi al felt ; pale yello w-coloured pigm ent diffusin g
int o th e su rro u nd ing agar; colony rever se (colou r plat e 158 , b) devel opi ng
reddish brown shades; peni cilli esse ntially as described aboye .
C olonies upon MEA (colour plat e 158 , C and e) att a ining a di amet er 01"
30 mm in 14 days at room temperature (20 -25 OC ), consisting of a loosetextured basal felt , conspicuo us ly funi cul ose with a cotto ny or velv cty
appeara nce , with a thin m argin surrounded by a frin ge of sub me rge d hyphae
exte nd ing abo u t 2 mm beyond th e ae rial growth, and centre slightly
umbon at e ; colony surface plan e , heavil y sporing throu gh out in cora l pink ,
sha des (Sgu y , PI. lII , 33); odo ur and exudate lacking; no coloured pigment
diffu sin g int o th e surro und ing a ga r; colony reverse (colour plate 158 , e)
developing ora nge to brown shades with a gree n cas t sho wing th rou gh ;
peni cilli esse ntially as dcscribed aboye but with m ost con idio pho res aris ing
fro m ro pes of h yph ae .
The- spec ies description is ce nte red up on th e type stra in CBS 344, 48 = C M I
40,5 70 = ATCC 10, 463 = NR R L 2, 146 = I F 7,729 = Q M 1929 = IJ FM 7113, isolat ed
in lul y 1947 as a laborat ory conta m ina nt by K .B, R aper a nd D.1. Fennell.
The correc t placement of thi s spec ies remain s somewha t ope n to qu estion
since it typi call y devel op s m or e irregularly and com plexly arrange d penicilli
th an othe r spec ies included within th e Penicillium pallidum series. T he spec ies is
appa rc ntly uncommon in n ature; Penicilliu m lavendulum has been rep orted twic e
as far as we know : th e type stra in and stra in CBS 582 .6 7, isolated from woo d of
Carya sp. by R . K enneth in 1967.
T he members of th e Penicillium pallidum series, characterized by th eir
appressc d roughe ne d peni cilli , th eir lon g cylind rica l, ro ughe ne d phi alides
with out a conspicuous ne ck and abse nce of green colours sho w a certain
resembl an ce a nd affin ity to P. cylindrosporum Sm ith a nd P. argillaceum Stolk,
Evans el Nil sson of th e su b -sec tion Asymmetrica -divari cat a .

haptcr 8

Asymmetrica-fasciculata sub-section

M EM BERS OF T HE F AS CI C U LATA A RE VERY ABUN DANT IN NATURE, E M B RAC ING

sorne of the m ost com monly isolated species of Penicillium. They are
cha rac te rized princip al1y by a co m plete or partial aggregation of th e
co nid iopho res into loos e or wel1-defin ed syn ne mata or fasc ic1es, lendin g a
gra nula r, tufted or ro ughe ne d a ppearance to th e colony surface . The penicil1i
a re usu al1y typi cal1y asym metrica l and two-sta ge branch ed .
They occur in soil, in air , as rots on store d fruit s and cereals, and upon a
great va riety of decaying organ ic su bstra ta . They are ofte n recovered from
ood product s, an imal feed a nd ph armaceutical products. T hey have become
the su bject of intensive bioch em ical research since isolates of sorne species h ave
been found to be potential m ycot oxin producers. H en ce , a co rrect identificatio n is important. U nfortunately , th e identification of sorne of the m ost
corn mo n fa scicul at e spec ies is ofte n problematic because of th e great variation
encou n te re d within th e species and th e presence of in tergr adin g stra ins.
R aper and Thom ( 1949) who treat ed th e Fasciculata as a sub -sec tion of th e
Asy m me trica section cons ide rably reduced the number of spec ies accepted by
Thom (1930) co ns ide ring m any of th em to be syno ny ms . M ore recently ,
Samson et al., (1 976) ca rried ou t an exhaus tive restudy of th is gro u p of fungi ,
cen te ring it primarily on th e m orphology of th e conidiogenous structures,
which led to a com plete revision and rearrangement of the Fascicul at a by
reducin g to sy no ny my 24 spec ies acce pted by R aper a nd T ho m ( 1949), both
within th e Fascicul at a and Lana ta su b -sections .

Acco rdi ng to th e mo st recent revision of th e gro u p (S amson el al. 1976) an d

to ou r ow n observations, th e fol1owing spec ies and varieti es m ay be
recognized :
Penicillium gladioli M cffull och el Thom
Penicillium verrucosum Di erckx varo cyclopium (Westling) Samson, Stolk , el
Hadlok
Penicillium verrucosum Di erckx va ro verrucosum Samson , Stolk, el H adlok
Penicillium oerrucosum Di erckx va ro album (S m ith) Samson , Stolk , el H adl ok
Penicillium verrucosum Di erckx varo melanochlorum Samson , Stolk, el H adl ok
Penicillium verrucosum D ierckx varo ochraceum (T hom ) Sa mson, Stolk, el H adl ok

Pen icillium uerrucosum D ierc kx varo corymbiferum (Westling) Sa mson , Stolk , el

H adlok
Penicillium echinulatum Fassat iov
Penicillium expansum Link ex Gray em end . Thom
Penicillium italicum W ehmer va ro italicum Samson, Stolk el H adl ok
Penicillium italicum W ehmer va ro aoellaneum Samson el Gutter
Penicillium hispalense R amrez el M artnez
Penicillium griseo-fuluum Di erckx
Penicillium granulatum Bainier
Penicillium claviforme Bainier
Penicillium claviforme Bainier muto candicans Abe el U ra ex R amrez
Penicillium claoiforme Bainier muto olivicolor Ab e el Ura ex R amrez
Penicillium clavigerum D emelius
Penicillium hordei Stolk
Penicillium isariiforme Stolk el M eyer
Penicillium concenlricum Samson , Stolk el H adlok
Penicillium kojigenum Smith
P. marne.ffei Segretain, Capponi el Su reau ex R amrez

Dichotomous key to the sp ecies and varieties

Page
1a. Species producing sclerotia, at least in recent isolates, more abundantly produced at 25-30 oC, less abundantly at lower temperatures
P. gladioli McCulloch etThom
2
1b. Species not producing sclerotia
2a. Conidia smooth walled or nearly so
3
2b. Conidia rough walled
22
3a. Conidiophores strongly synnematous, forming true synnemata with
distinct stalks .
4
8
3b. Conidiophores mononematous and/or loosely synnematous
4a. Synnemata without distinct stalk and an enlarged head; conidiophore
stipe smooth walled but rough walled at the base
P. clavigerum Demelius
5
4b. Synnemata with a distinct stalk and a globose head
5a. Synnemata distinct, on CA up to 20 mm long, on MEA longer; stalk
normally pink or reddish
6
5b. Synnemata well developed on MEA, short, up to 1.5 mm long, but on
CA less distinct; stalks not reddish
P. concentricum Samson, Stolk et Hadlok
6a. Conidial areas developing green shades
P. clavitorme Bainier
6b. Conidial areas not as aboye
7

.... 451

.. .. 491

.... 498
. . .. 486

a. Conidlal areas whlte

P. clevitorrne Bainier mut. csndicens Abe et Ura ex Ramlrez . . . . 488

b. Conidial areas developing olivaceous to ochre shades
P. clevitorme Bainier mut. olivicolor Abe et Ura ex Ramrez . .. . 490
9
8a. Conidiophore stipe on both CA and MEA definitely smooth walled.
8b. Conidiophore stipe on MEA mostly rough walled, on CA smooth to
rough walled.
12
9a. Phialides short, less than 6.5 .tm long, with a very short, inconspicuous
neck; rami strongly divergent; not causing citrus rot.
P. griseo-fulvum Dierckx .. .. 481
9b. Phialides longer than 6.5 .tm, with conspicuous neck; rami appressed,
not divergent
10
lOa. Colonies yellow-green to blue-green; conidia ellipsoidal , sometimes
subglobose; producing rot of pomaceous fruits.
P. expansum Link ex Gray emend. Thom .... 471
10b. Colonies gray-green or avellaneous; conidia ellipsoidal to cylindrical;
causing citrus rot.
11
11 a. Colonies gray-green, not exceeding 35 mm in diameter within 14
days on CA.
P. italicum Wehmer varo italicum Samson, Stolk et Hadlok . .. . 474
11 b. Colonies avellaneous, exceeding 35 mm in diameter within 14 days on
CA.
P. italicum Wehmer var. avellaneum Samson et Gutter .... 477
12a. Conidiophore stipe on CA smooth-walled or nearly so; penicilli
comparatively elongate; causing a rapid blue rot of pomaceous fruits
P. expansum Link ex Gray emend. Thom .... 471
12b. Conidiophore stipe on CA usually rough walled; penicilli comparatively short; not causing a rapid rot of pomaceous fruits. Though a few
strains may produce a limited rot.
13
13a. Conidiophores in recent isolates predominantly loosely synnematous
but also mononematous; conidia definitely ellipsoidal, about 3.0-3 .5 .tm
in long axis; odour pungent, aromatic .
P. granulatum Bainier . . . . 484
13b. Conidiophores predominantly mononematous, sometimes loosely
synnematous, conidia globose to subglobose, mostly 3.0-4.0 .tm in diameter; odour usually mouldy or earthy, rarely absent.
(P. verrucosum)

14

14a. Conidial areas white, yellow-brown or in other, not true green,

colours
15
14b. Conidial areas in yellow-green, gray-green or blue-green shades.
16
15a. Conidial areas at first white, later pale pink or cream.
P. verrucosum Dierckx varo album (Smith) Samson, Stolk et Hadlok .. . . 461

15b. Conidial areas yellow-brown to pale greenish -brown.

P. verrucosum Dierckx var oocbraceum (Thom) Samson, Stolk et Hadlok .. .. 464
16a. Colonies mostly growing restrictedly (not exceeding 30 mm in diameter within 14 daysl; bright yellow-green coloured .
P. verrucosum Dierckx var. verrucosum Samson, Stolk et Hadlok . . .. 459
16b. Colonies usually not restricted; not coloured in bright yellow-green
shades .
17
17a. Conidiophores in recent isolates predominantly loosely synnematous,
yellow mycelium and orange-brown exudate usually present; conidial
areas gray-green, in some strains blue-green when young; common on
liIiaceous bulbs.
P. verrucosum Dierckx var . corymbiferum (Westlingl Samson, Stolk et
Hadlok . . . . 466
18
17b. Conidiophores not as aboye.
18a. Mononematous conidiophores predominant.
19
18b. Mononematous and synnematous conidiophores present; colonies of
recent isolates often appearing tufted or granular, especially at the
margin, in other strains sometimes almost lanose; conidial areas graygreen, dull blue-green or bright blue-green .
P. verrucosum Dierckx var . cyclopium (Westlingl Samson, Stolk et Hadlok . . . . 455
19a. Conidial areas dark green; colony centre in recent isolates often
covered by white aerial hyphae.
P. verrucosum Dierckx var. melanochlorum Samson, Stolk et Hadlok ... . 463
19b. Conidial areas developing other colours.
20
20a. Conidial areas developing pink shades on CA and deep red colours on
MEA; pathogen to animals and man o
P. marneffei Segretain, Capponi et Sureau ex Ramrez . . . . 503
20b. Conidial areas of recent isolates at first pale bluish green , then developing dark gray-green shades.
P. kojigenum Smith . . .. 500
21a. Conidiophores strongly synnematous with distinct stalk . Synnemata
very long , up to 15 cm in length .
P. isariiforme Stolk et Meyer . ... 496
21b. Conidiophores not as aboye.
22
22a. Conidia up to 5 .tm in diameter.
P. echinulatum Fassatiov . .. . 468
22b. Conidia not exceeding 4 .tm in diameter; vegetative hyphae yellow;
exudate orange-brown; rather common on cereals .
P. hordei Stolk .. . . 493
23. Conidiophores multi-stage branched.
P. hispalense Ramrez et Martnez . . .. 479

T he Fascicul at a do not reprcsent a n a tural gro u p . T h e spccies h ave only th e

fascicul a te h a bit in co m mo n, bu t th ey sh ow m arked differences in ot her m ore
importan t taxonomi c features, suc h as th e str uctu re of th e co n id iophores a nd
phialides.

Th c sub -scciion Fascicular a as understood by R ap cr and Thom (1949)

grollped 19 spec ics into nine seri es, which were a rra nge d m ainl y upon th e
co lo ny charactc rs , su ch as texture a n d colo ur. H ow cver , th es e cha ra cters a re
not co n sta n t, and th ey m a y vary co ns idc rably within th e same sp ecies .
M oreover, no sharp Iin es of demarcation ca n be drawn between th e
Fa scicul a ta a nd th e three o the r su b -sec tio ns of th e Asymmetrica: V elutina ,
Lan at a , a nd Funiculosa . H ow ev er, th e Fa scicul ata differ markedly from th e
VeI u tin a by th e struc ture of th e co n id iopho re a nd by their ch aracte r istic
od ours.
The fas cicul ate habit va ries m arkedly , depending on the sp eci es . In so rne of
th em th e colon ies co ns ist of m ononem at ous 0 1' single con id ioph ore s mixed with
sm a ll tufts of co n id iop ho res 0 1' Ioose sy n ne mata. In other sp ecies the colo n ies
con sist mainly of co ns p icuous , well-defined syn nem ata , which a re usually
better developed and larger upon MEA th an CA .
The conidiophores of th e Fa sciculata m ay r ange from two- to three- 0 1' foursta ge branched , a s in Penicillium griseojulvum, to multi-stage braneh ed sue h as
P. hispalense Ramrez et M artnez. The rami m a y b e eithe r closely ap pressed to
the m ain axi s 01' divergent. M any speeies have rough-walled eo n id iop ho res ,
this b eing m ore pronouneed o n M EA . Other speeies, sue h a s P. italicum
W ehmer, have smooth -walled eo n id io pho re s.
In mo st speeies th e phialides a re more 01' less eylindrieal , tapering to a sho rt
neek su eh as in P. expa nsum Link ex Gray em en d . Thom. In P. griseoj ulvum
Di erekx th e phialide s a re mueh small er than in any other form of th e gro u p a nd
th e neek is in eonspieu ou s 0 1' nearly so . In other sp eei e s th e phialides a re
tapering more g rad ually to a n a rrow ed n eek lik e in P. isariiforme St olk et M eyer.
A pron ouneed odou r is eharaete rist ie of seve ral sp eeies, usuall y referred to as
'mouldy ' 01' earthy. In other s, sue h as in P. expansum, a ro matie odo u rs
suggesting th at of a pples a re produced .
Strains c1 assifi ed by R aper a n d Thom (1949) in th e Penicillium viridicatum, P.
cyclop ium a n d partly in th e P. granulatum serie s can n ot be di stinguish ed on
sign ifica n t m o rphological charac te rs and th erefore , accord ing to Samson et al.
(1976) , a re co ns idere d h ere as one var ia ble sp ecies under th e re-introdueed
n ame Penicillium verrucusom Dierckx.
On th e o ther h and , thou gh Penicillium gladioli M cCulloch et Thom sho ws a
clos er rel a tionship to th e sclerotial species of th e P. thomii a n d P. raistricki i
se r ies, a n d to th e eo n id ial states of sp ecies of Eupenicillium (Pi tt 1974) , it is
co ns ide re d h ere toge the r with th e Faseicul at a as a matter of co nven ience .

159. Penicillium gladioli McCulloch el Thom: fig . 15 9 , colo ur pl ate 159 ,

a n d S EM plate 27, (centre: right) .
Penicil!ium gladioli McCulloch et Thom - Science . 67: 216-217. 1928. See also,

l.w.J.!WJ

10 jrrn
: . ~
r~.V.,~"
~,--,'

, \ ....,.;..J.< :j

e
1

"F
..

:.-:::~

L-.--J

100pm

Fig. 159. Camera lucida

drawings of Penicillium
gladioli McCulloch et Thom,
CBS 214.28. A , Detailed
drawings of the penicilli. B,
Matu re conidia. C, Habit
sketches.

McCulloch and Thom, J. Agric. Res. 36: 217-224. 1928. Independently published
by J.E. Machacek under the same specific name, Ann. Rep. Quebec Soc o
Protect. Plants. 19: 77-86. 1928; also Thom, 'The Penicillia', pp. 381-383. 1930;
also Raper and Thom, 'A Manual of the Penicillia', pp. 471 -475. 1949.

Colon ies upon CA (colour piate 159 , A and a) growing rather res trictedly,
attaining a diameter of about 25 mm in 14 days at room temperature
(20- 25 OC ), consisting of a fairly tough basal felt, with central area u sually
raised and deeply floccose, and wit h marginal area radially sulcated by shallow
fur rows, and surrounded by a very narrow fr inge of submerged hyphae
extending less than 1 mm beyond the aerial growth . Scierotia characteristically
produced, especia lly in recent isolates, varying in number in different isolates
and depending upon environmental factors, particularly temperature ; u suall y
produced at temperatures of 25 oC and aboye, scantily at 15 " C and below ;
sporing abundantly in blue-green shades (Sguy, PI. XXX, 438, 443) partly
obscured by the deep floccos e overgrowth of wh ite ae rial hyph ae ; exudate and

odo ur lackin g ; rcddish brown -col ou rcd pi gmcnt diffu sin g int o the su rr o u nd ing
uga r; eolony re ve rso (colour plat c 159 , a) devcloping rcdd ish bro wn shades; in
rccent isolates sclero tia norm all y appeari ng in abo u t 6- 7 days a nd developing
in succ ess ive conce ntric zo nes 0 1' as sec tors in locali zed areas, va ry ing greatly
in size , usu all y rangin g fro m 150 to 300 p.m but sometimes up to 500 p.m in
di ameter, a t first cream to pale pinkish tan, in age becoming tan 01' very pa le
br own , smooth, with walls co ns isting of th ick-w alled cells 8- 12 p.m in
d iameter; con id iophores aris ing primaril y fro m the substra tu m , m ostly as
independe nt structu res 0 1' m ononematou s but ofte n more 0 1' less aggr egated
in to definite fascicles, with wa lls eithe r smooth 0 1' ro ughe ned; gene rally few ,
scattere d, a nd in con spi cu ou s among th e sclerotia in new isolates , in older stock
cultures more abu ndant a nd co m mo nly aris ing from within th e floccose felt ,
often very long (u p to 2 m m ) and abo u t 2.5-4 .0 p.m in di ameter ; peni cilli
com paratively large , usu all y cons isting of one - 0 1' two -stage branch ed
stru ct ures (fig. 159, A), with 0 1' without one 01' two rami 20- 25 tut: by
2.8 -3.5 tut, bearing verticils of few m etulae 10-1 5 p.m by 2.8- 3.5 p.m ;
phialides 2-5 in th e vert icil, 9 - 12 p.m by 3- 4.0 tun , tap ering to a short nec k;
conid ia su bglobose, smooth wa lled 01' nearly so (fig. 159, E , and SEM piate 27.
centre : ri ght) ; 3.5 -4 .0 p.m in long ax is; con id ial cha ins often adheri ng in lon g
cha ins, more 0 1' less loosely parall el 0 1' tangled (fig . 159, C).
Colon ies upon C YA (colour plat e 159, E and b) spread ing broadl y , at ta ining
a d ia me ter of 55 mm in 14 day s at room temperatu re (20- 25 OC ), cons ist ing of
a deepl y floccose felt , radiately furrowed and rai sed towa rds th e centre , with a
thin eve n m argin surrou nde d by a narrow fri nge of sub me rge d hyph ae
extend ing 1 mm beyond th e ae r ial growth; sporing abu ndantly in mo use gray
shades (Sguy, PI. XXX V , 523), especially in marg inal zones; odo ur lack in g;
exudate ab u ndant , as la rge yellow a m be r-co lou re d drops; am ber-colou red
pigm en t di ffu sin g into th e surro u nd ing agar ; colony reverse (colour piate 159,
b) in reddish cinriamon shades; peni cilli essentially as described abo ye;
sclerotia as described abo ye, not produced in old stock cultu res.
C olonies upon M EA (colour plat e 159, C and c) attaini ng a d iame ter of
50 mm in 14 days at room temperature (20- 25 OC ), consis ting of a looser
tex tu red basal felt , plane , not fu rrowed , with centre slightly umbonat e , thinn er
th an whe n on the aboye su bs trata, floccose , with thin m argin surrou nde d by a
wide fringe of subme rge d h yphae ex tend ing abo u t 4 m m beyond the aerial
growth ; abu ndan tly spo ring in old stoc k cultures in pale yellowi sh shades
lending a gra n ula r appe arance to the colony su rface ; in recen t isolat es,
abundant p rodu cti on of sclero tia whi ch gene rally dominat e th e colony as pec t ;
odour fruity , aromatic; exudate lacking ; no colou re d pigm ent d iffus ing int o th e
surroundin g agar; peni cilli essentially as described above.
The spec ies is ofte n found to be th e ca u se of the ro t in gla d iolus corms .
The spec ies description is ce ntere d prirnarilv on the type strain C BS

1!i4

332.48 = IMI 34, 911 = AT C C 10,448 = NRRL

9 :~ 9

... QM 1955 isolat ed originally by

L. M cCulloch from cor m of Gladiolus sp. in 1927, W ashington, D. C ., and

upon strain C ES 214. 28 = IJ FM 7107, isolated b y J .E. M achacek at M acDonald

College, Queb ec, C anada, from co r m of Cladiolus sp . , 1927. The latter differs
from th e typ e in producing more restrict ed colon ies th at are more
cons picuo us ly floccose and produce com para tiv ely few sclerotia, eve n upon
MEA .
Thom (1930), asc ribe d priority of publication to M achacek , and th e same
u sag e wa s follow ed by R aper a nd Thom (1949) in th eir 'Manual ' , bu t
W ak efield an d M oore ( 1936), afte r revi ewing th e history of th e species,
co nclude d that it sho uld be cite d as Penicillium gladioli M cCulloch el Thom .
Penicillium gladioli, th ou gh included in th e p resent study within th c
Fascicula ta su b-section, primarily as a matter of convenience , sho ws , however,
a cer tain relationship to species of th e P. raistrickii series th at should not be
ove rloo ked, su ch as th e rather di varicat e characte r of its penicilli , th e ro u ghwalled coni d iop ho res, and particul arly th e production of sclerotia.
Penicillium uerrucosum Dierckx com p le x

Penicillium verrucosum Dierckx - Ann . Soco Sci. Bruxelles. 25: 88. 1901.
O UTSTAN DlNG FEATURES

C olonies upon C A gro wi ng res tricted ly or sp reading broadly . Usually

cons isting of a d en se felt of monon emat ou s con id iopho res mi xed with small
tufts of syn nem a tous conid iophores , particul arly produced in marginal arcas,
lending th e colo n y su rface a gran ula r a ppearance; in othe r strains the colony
su rface is predominantly velvet y or almos t floccose ; degen erated stra ins m ay
be floccose to la n at e . Colours rangi ng from yellow-g ree n, dark- green , graygree n to blue-green , seldom white or ochraceous . Aerial vegetative hyphae
usu all y absen t, sometimes yellowish or whitc cu shion -like m ycelial str uc tu res
are produced in cent ra l areas. Colony reverse rangin g from colou rles s to
yellow ish, pin k, orange or brown colou red. Odou r pron ounced , u su all y earthy
and pungent , sometimes aromatic or some what fruit y. Con id iopho res
100-400 tm b y 3.0-4 .0 urn , two-stage branch ed , some times three- or m ore
stage branch ed , rou gh walled (in sorn e stra ins of varo cyclopium smooth walled) ,
with rami a nd m etul ae appressed. Phialides bottle shape d, 7.0-9 .0 tm by
2.0-2.5 uit , with sho rt bu t co ns picuous neck . Con idia globose to su bg lobose,
sometimes ellipsoidal whe n yo u ng, smoo th wa lled or nearl y so , gree n ish,
3.0 - 4.0 tu in di amet er.
Colonies upon C YA with sim ila r growth , but usu all y spread ing more
b ro adl y.
Colo nies upon M EA with sim ila r growth, but u su all y thinner a nd less

fuscicul at e. O dou r usu ally mo re pronounccd th an llp OIl C A a n d C YA .

C o n id io phores a s llp OIl C A, hut usu all y more m arkcdly ro u ghe ne d .
' I'hc llam e Penicillium uerrucosum wa s int roduccd by R .P . Di erck x (1901) with
:1 sho rt d cscription. Biourge red e scri bcd th e sp ecie s ba sed o n a new isol at e a nd
th c or iginal drawings an d notes left uy Dierckx. R aper a nd T ho m (1949)
o nsid cred Dierckx ' s spec ies as a sy nonym of P. uindicatum Westli ng ( 19 11)
n nd state d th at P. oerrucosum ha s been commo nly in te rpre te d as representing a
iucm bc r of the P. uiridicatum se r ies .
Penicillium oerrucosum is th e oldest n ame of a se ries of ep ithe ts publish ed for a
ro m plex of fa scicul at e P enicilli a with rough-walled co n id iophores a nd globose
lo su bglobose, smooth - to slig htly rough-wall ed co n id ia. Strains formerl y
di sposed b y R aper a n d Thorn ( 1949) into the P. oiridicatum, P. ochraceum, P.
cyclopium se r ies a nd p artl y the P. expansum a n d P. granulatum se ries, also th e
spec ies of th e P. terrestre se r ies (sub-section Funiculosa) a n d so rne of th e P.
comm une se r ies (sub-sect ion Lanata) ha ve been in cluded b y Samson et al. ( 1976)
in Penicillium uerrucosum D ierckx.
Though th e species pl a ced in syno ny my w ith P. oerrucosum agree in th e
structure of the co n id iopho res , th ey m ay differ in th e co lo ny colo u rs .
Moreover, so rne va r ia tion in th e rate of growth, fa scicula ti on a n d ro ugh ness of
the co n idi op ho res m a y be e ncountere d . Accord ing to the aboye au thors, a
separa tion of P. oerrucosum into six var iet ies, album, corymbiferum, cyclopium,
ochraceum, melanochlorum a n d oerrucosum ba sed primarily on th e differences of
co lo ny colou r see ms 10 be rea sonably practicable .
Accordingly , Samson et al. (1976) c1assifica tion ha s been adopted in th e
cu rre n t stu dy.

160 . Penicill ium verrucosum Di erekx val'. cyclop ium (Westling) Sa mso n,

Stolk, el H adlok: fig. 160 , eolour plate 160 , a nd SEM plat e 27,
(bottom: left) .
Penicillium verrucosum Dierckx var. cyclopium (Westling) Samson, Stolk et Hadlok
- Studies in Mycology. 11: 37-41. 1976.

Synonymies
?Penicillium aurantiogriseum Dierckx - Ann. Soco Sci. Bruxelles . 25: 88. 1901.
?Penicillium puberulum Bainier - Bull. Soco Mycol. Fr. 23: 16. 1907.
Penicillium biforme Thom - Bul!. Bur. Anim. Ind. U.S. Dep. Agric. 118: 54. 1910.
Also Raper and Thom, 'A Manual of the Penicillia', pp. 437-439. 1949.
Penicillium cyclopium Westling - Ark. Bot. 11: 55.1911. Also Raper and Thom,
'A Manual of the Penicillia', pp. 493-497. 1949.
Penicillium palitans Westling - Ark . Bot. 11 : 55. 1911. Also Raper and Thom, 'A
Manual of the Penicillia', pp. 488-489. 1949.

~ ,;

..

~,.

\\\

~y~ 1
.

100jJm

fff..

Fig. 160. Camera lucida

drawings of Penicillium
verrucosum Dierckx, varo
cyclopium (Westling)
Samson, Stork et Hadlok,
CSS 690.77. A, Detailed
drawings of the penicilli . S,
Mature conidia. e, Habit
sketches.

Penicillium majusculum Westling - Ark. Bot. 11: 51. 1911.

Penicillium conditaneum Westling - Ark. Bot. 11: 52. 1911.
Penicillium solitum Westling - Ark. Bot. 11: 52. 1911. Also Raper and Thom, 'A
Manual of the Penicillia', pp. 453-455. 1949.
Penicillium casei Staub - Z. Bakt. Parasitenkunde. Abt. 2, 31: 454. 1911. Also
Raper and Thom, 'A Manual of the Penicillia', pp. 401-402. 1949.
Penicillium terrestre Jensen - Bul!. Cornell Univ. Agr. Exp. Stn . 315: 486. 1912.
Also Raper and Thom, 'A Manual of the Penicillia', pp . 450-453. 1949.
, enicillium aurantiovirens Biourge - La Cellule 33: 119. 1923. Also Raper and
Thom, 'A Manual of the Penicillia', pp. 503-505. 1949.
Penicillium martensii Biourge - La Cellule 33: 152. 1923. Also Raper and Thom,
'A Manual of the Penicillia', pp. 500-503. 1949.
Penicillium janthogenum Biourge - La Cellule 33: 143. 1923.
Penicillium flavidimarginatum Biourge - La Cellule 33: 150. 1923.
Penicillium porraceum Biourge - La Cellule 33: 188. 1923.
Penicillium olivinoviride Biourge - La Cellule 33: 132. 1923. Also Raper and
Thom, 'A Manual of the Penicillia ', pp. 487-488. 1949.
Penicillium flavoglaucum Biourge - La Cellule 33: 130. 1923.

Penicllltum btunneo-viotsceum Blourge - La Cellule 33: 145. 1923.

Penlcllllum surentiocendidum Dierckx - In Biourge 's Monographie, La Cellule 33:
116. 1923. Also Raper and Thom, 'A Manual of the Penicillia', pp. 442-443.1949 .
Penicillium eursntiostbidurn Biourge - La Cellule 33: 197. 1923.
Penicillium aurantiogriseum Dierckx varo poznaniensis Zaleski - Bull. Acad. PoI.
Sci. Sr. B. pp. 444. 1927.
Penicillium johanioli Zaleski - Bull. Acad. PoI. Sci. Sr. B. pp. 453. 1927.
Penicillium polonicum Zaleski - Bull. Acad. PoI. Sci. Sr. B. pp. 445. 1927.
Penicillium crustosum Thom - 'The Penicillia', p. 399. 1930. Also Raper and
Thom, 'A Manual of the Penicillia', pp. 516-518. 1949.
Penicillium lanoso-griseum Thom - 'The Penicillia', p. 327. 1930. Also Raper and
Thom, 'A Manual of the Penicillia', pp . 441-442. 1949.
Penicillium lanoso-coeruleum Thom - 'The Penicillia', p. 322. 1930. Also Raper
and Thom, 'A Manual of the Penicillia' , pp. 436-437. 1949.
Penicillium bordzilowskii Morotchkovsky - Nauk. Zapo Kyyiv. Derzh. Univ. 2: 71.
1936.
Penicillium casei Staub varo compactum Abe - J. Gen. Appl. Microbiol. Tokyo.
2: 101. 1956 (not validly published).
Penicillium viridicyclopium Abe - J. Gen. Appl. Microbiol. Tokyo. 2: 107. 1956
(not validly published) .
Penicillium pseudocasei Abe - J. Gen. Appl. Microbiol. Tokyo. 2: 102. 1956 (not
validly published): ex Smith - Trans. Brit. Mycol. Soco 46: 335. 1963.
Penicillium roqueforti Thom varo punctetum Abe - J. Gen. Appl. Microbiol.,
Tokyo. 2: 99. 1956 (not validly published).
Penicillium conservandi Novobranova - Nov. Sist . Niz. Rast. p. 233. 1974.

Colon ies upon C A (colou r pi a te 160, A and a) usu all y sp read ing b ro adl y ,
att aini ng a di amet er of 45- 50 mm in 14 d ays at room te m pe rature
(20-25 OC ), sometimes gro wing slower, with a di amet er of 30-3 5 mm ; in
recent isolates strongly granular with m any fascicula te con id iop ho res
p articul arl y in marg inal areas, which are thin and su r rou nded by a frin ge of
sub me rged h yphae ex te nd ing about 3- 4 mm beyond th e aeria l growth;
fascicul ati on is u su all y arranged in co nspicuo us conce n tric zo nes; in sorn e
strains th e colo ny su rface is almost velve ty or floccose , but sho wing sorne
fascicul ation ; heavil y spo ring through out in gray-green to dull blue-green or
bright blue-green , ran gin g from pal e blue to gray -blue shades, ap p rox imating
arte m isia, lily , celandine , gnaphaliu m, sage-gree n , (Sguy, PIs . XXVII , 404,
401, XXX, 448, 447 , 446, XXXI , 463, 462, 461) with th e ripening of con id ia ;
ve geta tive ae rial h yphae m ostly a bsent, in sorn e strains yellow ish or h yaline ;
ex udate m ostl y present in recent isolates as colou rles s droplet s; odo u r st rong,
earthy or m ouldy , som etimes aro matic, fruity , u su all y a p ple-like, th ough in a
particul ar strain , IJ F M 3865, isolated by us as a culture con ta m inan t in our

lab oratory , M ay 1977, (sce co lo u r plat o 167 , a nd lig. 167, p. 471) a fra gr ant od ou r
suggestive of Anan s satiuum wa s dct cci cd , th e co nidio phore wall was srn ooth ,
and a bright ye llow pigment was diffu sin g into th e su rro u nd ing agar upon C A
a nd CYA (colou r plate 167, A and B). H owever, this str ain duplicated th e type
strain in details 0 1' morphology . Colony re verse (co lou r plat e 160 , a) ora ngeb rown , yellow, pinkish , purplish to ligh t m a ro on , dep endin g upon th e strain;
yellow-colo ured pi gm ent some times diffusin g into th e surrou nd ing agar.
Penicilli la rge (lig. 160 A), a bo u t 50-60 fAm in len gth , asy m metric, bearing
loosely parall el conid ial cha ins to tangled chains 0 1' co nid ia in irregul ar m asses
up to 150 fAm lon g (fig. 160, C ), with con id iophores arising from th e
subs tra tu m, m ostl y 200-400 fAm in length b y 2.8 -3 .5 fAm in diamet er,
sometimes coarser , with wall s typi call y ro ughe ne d but in sorne strains
ap pearing smooth 0 1' nearl y so (lig . 167 , A); penicillus showing one 0 1'
occasionally m ore ram i, 20-30 fAm by 3.5 tun , 0 1't en ap p ressed, u su all y
bearin g sm all clusters 0 1' 2-4 m etulae 12-17 tui: by 2.4 -4.0 tuu; each
su ppo r ting a wh orl 0 1' 4-8 phialides, abou t 9- 12 fAm by 3.0-4 .0 ust , bottle
shaped, taperin g ab ru ptly to a very sho rt neck; co nid ia globose to su bg lobose,
mostly 3-4 .5 fAm in d iam et er, with wall s smooth 0 1' delicat ely roughened (lig .
160, B, and SE M plat e 27, bottom: left ); conid ia l chains usu all y loosely
parallel to tangled , some times forming well-de fine d di vergent co lu m ns (lig .

167, C).
C olonies u pon C YA (colour pl at e 160, B a nd b) attain ing a di amet er 0 1'
60 mm in 14 d ays at roo m temperature (20-25 OC ), sim ila r in texture as noted
aboye but heavier spo ring throu gh out in da rk er shades 01' gree n (Sguy, PI.
XXIX, 431) with coni d ial zo nes arranged in co nce n tric areas; odo ur mouldy ;
ex udate en meshe d in the m ycelial felt ; a mber-colou red pi gm ent diffusin g into
th e su rrou nd ing agar , in sorne strains (co lour pl at e 167, B) bright yellow
pi gm en t d iffus ing into th e su rrou nd ing agar; colony reve rse in reddish brown
(co lour plat e 160 , b) 0 1' yellow shades (colour pla te 167, b); penicilli esse nt ially
as described a boye.
Colonies upon M EA (colour plat e 160, C a nd e) sp re ad ing b roadl y ,
a ttaini ng a di ametcr 01' 65 mm in 14 d ays at roo m temperature (20-25 OC ) ,
co ns isting 0 1' a looser text u re d basal felt, plane , not furrowed , co ns picuo usly
zo nate, a ppearing more 0 1' less gra nula r, showing co nce n tric zo nes 0 1' fasc icles;
with thin margin surro unde d by a fringe 0 1' su bmerged h yphae ex te nd ing abou t
4 mm beyond th e ae rial growth ; heavil y sp oring th ro ugho u t in very deep bluegree n shades (Sguy , Pi s. XXIX , 43 1, and X XX , 446) ; odour penetrating ,
cart hy to m ouldy; exudate la cking; no coloured pigm ent diffusin g into th e
su rrou nd ing agar ; colony reverse colou rless 0 1' developin g orange shadcs;
pe nicilli as d escribed a boye but with th e con id iop hore wa lls more coarsely
ro ug he ned .
Sp ecies description is ce nte re d up on strain C BS 161.42 = CM ! 39,803 = ATCC

11 ,507

NJ{RL

i!)() .77

I"FM 7251, isolat cd by Ph . Biourgc , also on stra in C IIS

:n:>2, isolat ed lrom th c ai r 01' Las Palmas de G ra n C a naria ,

912

I,lFM

Cl na ry lslands , Spain , May, 1976 , and many othe r strain s, rcgarde d a s

l'l:pl'l:senting thi s spec ies, isolatcd in our laboratory . (Colour plates 243 , 244,
245 and 246 present tour representative strains 01' P. verrucosum val' . cyclopium
isola tcd by th e author , one from red pepper, two from th e a ir , and one from
so il, respc ctivcly .)
l'eniclllium verrucosum Dierckx varo cyclopium Samson , Stolk el Hadlok is
rc por tcd from m any differ ent substrata and is world-wide in di stribution . It
luis been isolat ed in H olland a nd in th e USA as the domin ant m ould in rotting
hulbs al' th e liliaceae family. It is amo ng th e m or e common spec ies isolated
lro rn va rious typ es 0 1' rip e 01' aging fruits, gr ain and cereal products , soil and
t1 ccayin g veget ati on , bee hives , mildewing tentage, cte .

I() 1. Penicillium verrucosum Dierckx val'. verrucosum Samson , Stolk el

Hadlok: fig . 161, colour plate 161, and SEM plate 27, (centre: right) .

'.:-.

.' .
....

:.-.

':.:.

~l-

~)~ V
'\

l.--..J

100

jrm

eR.

Fig. 161. Camera lucida

drawings of Penicillium
verrucosum Dierckx, var o
verrucosum Sarnson, Stolk
et Hadlok, CBS 222.71. A,
Detailed drawings of the
penicilli. B, Mature canidia.
C, Habit sketches .

Penicillium verrucosum Dierckx varo verrucosum Samson, Stolk et Hadlok Studies in Mycology. 11: 34-35. 1976.

Synonymies
Penici//ium verrucosum Dierckx - Ann. Soco Sci. Bruxelles. 25: 88. 1901 .
Penicillium musae Weidemann - Z. Bak. Parasitenk. Abt. 11, 19: 687. 1907.
Penicillium viridicatum Westling - Ark. Bot. 11: 53. 1911.
Penicillium stephaniae Zaleski - Bull. Acad. PoI. Sci. Sr. B. pp. 451-452. 1927.
Penicillium b/akes/eei Zaleski - Bull. Acad. PoI. Sci. Sr. B. pp. 441-444. 1927.
Penicillium psittacinum Thom - 'The Penicillia', p. 369. 1930. Also Raper and
Thom, 'A Manual of the Penicillia', pp. 448-449. 1949.
Penicillium /anoso-viride Thom - 'The Penicillia', p. 314. 1930. Also Raper and
Thorn, 'A Manual of the Penicillia', pp. 434-436. 1949.

Colonies upon CA (colour plate 161, A and a) growing usually restrictedly,

attaining a diameter of 25-35 mm in 14 days at room temperature, but in
sorne strains (colour plate 161, A) spreading broadly, attaining a diameter of
50-60 mm in the same days at room temperature (20-25 OC), consisting of a
tough felt up to 1 mm deep, azonate to broadly but indistinctly zanate, and
usually radiately wrinkled, with broad white margin surrounded by a fringe of
submerged hyphae extending about 4 mm beyond the aerial growth; heavily
sporing throughout in deep blue-green shades (Sguy, Pi. XXX, 448, 447),
with surface typically granular; little or no exudate produced; odour
pronounced, 'earthy'; no coloured pigment diffusing into the surrounding agar;
colony reverse (colour plate 161, a) colourless or in pale shades of yellowish
orange; conidiophores (fig. 161, A) with walls finely roughened , about 4 /lm in
diameter, arising either as short branches from trailing or ascending hyphae
mostly less than 50 /lm in length, or from the substratum and up to
200-250 /lm in length ; penicilli asymmetric, usually consisting of a main axis
with or without one or more appressed rami, sometimes simple (appearing
monoverticillate); metulae irregularly produced, commonly in groups of 2-4,
often at different levels, measuring 8-15 /lm by 2.5-3 tuu , occasionally up to
20 /lm in length ; phialides, closely packed and few in the verticil, 7-10 /lm by
3.0-3 .5 utt, bottle shaped, tapering abruptly to a very short narrowed neck;
conidia globose to subglobose, 4-5 /lm in long axis, with walls smooth or
irregularly and finely roughened (fig. 161, B, and SEM piate 27, bottom :
right); conidial chains forming loosely parallel columns or tangled (fig . 161, C) .
Colonies upon CYA attaining a diameter of 65 mm in 14 days at room
temperature (20- 25 OC), similar in texture and colony appearance to the
aboye description but more strongly radiately wrinkled and more conspicuously zonate (colour plate 161, B and b), with a thin margin surrounded
by a fringe of submerged hyphae extending about 10 mm beyond the aerial

growth ; hca vily spori ng throu gh out in decp hlu c-grccn shude (S guy, PI.
XXX, 442, 447) ; limit ed clca r exu da re pr oduccd ; odour stro ng, 'ca rthy";
pcni cilli more consisten tly asym mc tric an d la rger but ot he rwise as on C A; no
coloured pigm cnt d iffus ing int o th e su rrou ndin g a ga r ; colony reverse (colour
plat e 161, b) dev eloping pale yellow ish orang e lo reddish brown shades.
Colonies u po n M EA (colour pla te 161, C and c) atta ining a di ameter of
about 55 mm in 14 days at room tempera ture (20-25 OC ), consisting of a
looser textured basal felt , com parativ ely thi n , appearing velvety,
conspicuously zonat e , showi ng zo nes of fasc icula te conidiopho res, especially at
th e marg in , whic h is thin a nd su rro u nded by a frin ge of subm erged hyphae
cxte nd ing abo u t 5 m m beyond th e ae rial gro wth ; heavily sporing throu gh ou t
in dee p grecn colours (Sguy, PI. XXVII , 401); ex udate not produced ; odo ur
strong, ' earthy ' , suggest ing act ino mycetes; penicilli m ore abundantly
produ ced , esse ntially duplicat in g th ose u po n C A and CYA but with
con id iophores m ore cons picuously rou ghened ; no coloured pi gm ent diffu sin g
int o th e su rro u nd ing agar ; colony rever se (colou r pIale 161, e) developing
red dish brown shades .
Species descr iption is ce nte red prim arily upon strain CBS 265 .29 = ATCC
10,116 = NRR 932 = IJ FM 3864, isolat ed by Ph . Biourge as P. aureum Corda, and
stra in C BS 603 .74 = NR R L 965 , isolated by Biou rge and cons ide re d as th e
neotype of Penicillium verrucosum Di erckx .
P. oerrucosum val'. verrucosum ca n be di stingui shed from th e othe r va rieties by
its com m only restricted gro wth (ge ne rally Icss th an 30 mm in d iameter within
14 days), a nd its bri ght yellow -gree n colour upon CA. Althou gh th e strain CBS
265.29 (for me rly th e typ e strai n of P. psittacinus T hom) (see colour plate 161)
grows rather rapidly , the spec ies is cons ide re d by Samson el al. ( 1976) to be
identical to P. verrucosum va l'. verrucosum on acco unt of its identical
conid ioge no us structures.

162 . Penicillium oerrucosum Di erckx va l'. album (Smith) Samson , Stolk el

H adl ok : fig. 162, colou r plat e 162, and SEM plat e 28, (to p: left).
Penicillium verrucosum Dierckx varo album (Smith) Samson, Stolk et Hadlok
Studies in Mycology. 11: 35-36. 1976.

Synonymy
Penicillium cyclopium Westling varo album Smith - Trans. Brit. Mycol. Soco 34:
18. 1951.

C oloni es upon CA (colour plat e 162, A an d a) growing rapidl y , at aining a

di amet er of 50 mm in 14 days at room temperature (20-25 OC ), cottony,
white, becomin g pale pink 0 1' cream; with tufts of con id iopho res especially at

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Fig. 162. Camera lucida

drawings of Penicillium
verrucosum Dierckx varo
album (Smith) Samson,
Stolk et Hadlok, CBS
343.51. A, Detailed drawings
of the penicilli. B, Mature
conidia. e, Habit sketches .

the colony m argin , with colony surface almos t pl ane , sligh tly rais ed in
co nce n tric zones; odo u r lacking ; exudate not produced ; no colou re d pi gm ent
diffu sin g into th e su rrou nd ing agar; colon y rever se (colour plate 162, a)
developing cream rosy to pale ora nge shade s; 'conid iogenous structures
identical to the othe r variet ies (Fig. 162, A , B a nd C ) .
Co lonies upon CYA (co lou r pla te 162, B and b) sim ila r in cult ural
characte ristics, det ail s of m orphology and rat e of growth to th ose d escribed
up on C A but strongly radiately wrin kled, with m embranou s m argin , a bo ut
1 mm submerged ; odo u r and ex udate lacking; pal e yellow -colou re d pi gm ent
d iffusin g in to th e su rro u nd ing agar ; colon y reverse (co lour pIate 162, b)
developing pal e yellow -orange shades .
Colon ies u pon M EA (co lour plat e 162, C and c) attaini ng a di amet er of
50 mm in 14 d ays at room temper ature (20-25 OC ) , co nsisting of a looser
tex tu red b asal felt , ove rlaid b y an ab undan t ove rg row th of white ae ria l floccose
hyphae , lending a lanate a ppearance to th e colony su rface ; odo u r stro ng,
aro matic; ex udate lacking ; no colou re d pigment diffu sin g in to th e su rro u ndi ng
agar ; colo ny rev erse (co lour pl ate 162 , c) in yellowish orange shades.

T he vari cty descrip tion is ccntcrcd upon the typc strn in o us :H:1. 5 1 - C M I
- ATCC 111 , 2 :~ 2 - Q M ll, 1411 - IJFM 7 15G, isola tcd by Mrs S. Marcus fr orn
Musa sapicntum [ruit , a l th e London Schoo l 01" H ygien c a nd T ropical M edi cine ,
K, May, 1947.

ll !), :~ 12

1(j:s. Penicillium verrucosum Di erckx va l'. melanochlorum Sa mso n, Sto lk , et

Hadl ok : fig.1 63 , colou r plate 163 , an d SEM pIate 28 , (top : ri ght).

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8

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Fig. 163. Camera lucida

drawings of Penicillium
verrucosum Dierckx varo
melanochlorum Samson,
Stolk et Hadlok, CSS
487.75. A , Detailed drawings
of the penicilli . B, Mature
conidia. e, Habit sketches.

Penicillium verrucosum Dierckx varo melanochlorum Samson, Stolk et Hadlok Studies in Mycology . 11: 41-42.1976 .

Synonymy

Penicilliumpalitans Westling sensu Raper etThom - 'The Penicillia', pp. 488-489.

1949.

Colonies u pon CA (colour plat e 163, A and a) att ain ing a diameter of about
40 m m in 14 days at roo m temperat u re (20 -25 OC ), consisting 01" a den se

vclvcty laye r 01' m on ou cruatou s con idiopho res. soru etimes showing so rne lascieulate gro wth in the margin al arcas, ran gin g from ab out 300 to 500 .un dccp in
marginal to submarg inal zon es, and up lo 1- 2 mm in the colony cent re, with
su rfaee appearing granu lar 01' meal y but gen erall y not eo nsp ieuo usly
synnem at ou s, 01't en narrowl y zo nate with shallow radi al furrows, with centre
mostly eovered by a wh ite floeeose eushion 01' ae rial veget a tive hyphae partly
obseu ring th e othe rw ise heavily spo ring eonidial areas in dark gree n eolours ,
near Ame rican 01' Russian gree n (Sg uy, PI. X XX , 441 , 446) ; odo ur stro ng,
sug gesting ea m phor aroma; ex udate limited in amo u nt, as eolou rless droplets
enmes he d in the myeelial felt ; very pale am be r-co loured pigm ent diffusing into
th e su rro u nd ing agar ; eolony reverse (colou r pla te 163 , a) in pale eream shades
to eolourless; eonidial stru etu res esse ntially sim ilar to the othe r variet ies (fig .
163, A , B a nd C) .
Coloni es upon C YA gro wing more rapidly, attaining a diam eter 01' 60 mm
in 14 days at roo m tempera ture (20-25 oC) (eolour plate 163, B a nd b) ,
appearing granular, with a th in margin su rro u nde d by a frin ge 01' sub me rged
hyphae extending about 3 mm beyond th e ae rial gro wth , with eolony su rfacc
almos t entirely eovered by a eus hio n 01' whitish floeeose ae rial hyphae greatly
obse uring th e othe rw ise abu ndantly spo ring areas in dark blu e-gr een shade s
(Sguy, PI. X X X, 442), sulcated by shallow radial fur row s; odo ur faint,
sug gesting pot atoes; exu date limi ted , as eolourless dropl et s enmeshed in the
myeelial felt; no eoloured pi gm ent diffusin g into th e surrounding agar; colony
reverse (eolour plat e 163, b) developing reddish brown to einnamo n shade s.
Co lonies upon M EA (eolour piate 163, C and e) attain ing a di ameter 01'
50 mm in 14 days at roo m tem per ature (20- 25 OC), eons isting 01' a looser
textured basal felt , eom paratively thin , heavily spo ring, plan e, exee pt 1'01' a
slight floeeul ent overgrowth all over the eolony su rfaee, with very thin margin
surrou nd ed by a frin ge 01' sub me rge d hyphae ex te nding abo ut 4 mm beyond
th e aerial gro wth; eonidial areas in deep blu e-green shades (Sguy, PI. X XX ,
442); odo u r pen etrating, cam pho r-like ; exudate not produeed ; no eolou red
pigm ent di ffusin g into th e su rrou nd ing agar; eolony re verse (eolou r piate
163, e) developing yellowish shades.
The va riety de seription is eentered upon th e typ e culture CES 487 .75,
isolated in 1975 by R . H adl ok from meat produets, Gi essen , Fed eral R epublie
01' G ermany .

164 . Penicillium verrucosum Dierckx val'. ochraceum ( T hom) Samson, Stolk et

H adlok : fig . 164, co lo u r pl ate 164 , a n d SEM pI a te 28, (cen t re : Ieft) .
Penicil/ium verrucosum Dierckx var oochraceum (Thom) Samson, Stolk et HadlokStudies in Mycology . 11: 42-43. 1976.

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Fig, 164, Camera lucida

drawings of Pencllium
verrucosum Dierckx var.
ochraceum lThorn] Samson,
Stolk et Hadlok, CBS
246.32. A, Detailed drawings
of the penicilli. B, Mature
conidia . e, Habit sketches ,

Synonymies
Penicillium ochraceum Thom - 'The Penicillia', p. 309. 1930. (cited as P.
ochraceum (BainierrThom) . Also Raper and Thom, 'A Manual of the Penicillia',
pp . 477-478. 1949.
Penicillium ochraceum Thom varo macrosporum Thom - 'The Penicillia', p, 310.
1930.
Penicillium carneolutescens Smith - Trans. Brit. Mycol. Soco 22: 253. 1939. See
also Raper and Thom, 'A Manual of the Penicillia', pp. 479-481. 1949.

Colonies upon CA (colour plate 164, A an d a) attain ing a di ameter of

60 mm in 14 days at ro om temperature (20-2 5 OC ), in sorne stra ins m ore or
less floccose, 2- 3 mm deep , in othe rs almost velvety, az on at e at first but
co rnm only definitely zonat e in age, radiately furrow ed, bearing abundant
con id ia l str uc tu res particul arl y alon g th e intercolony m argins, in othe r stra ins
gro wing more restrictedly , attaini ng a diamet er of 30 -35 mm within th e sa me
period of time ; with thin m argin surro u nded b y a wide frin ge of su b merged

hyphac cx rc nd ing a bo u t :> 111111 be yond (he aeri ul gro w th; gro wing m argin
whitc to du ll bulf in colou r, conidia l arcas !>ecollling yellowish olivo l o dar k
greenish olive 01' buffy olive wh en mature in reccnt isolat es (Sguy, PI. XXIII ,
338, 33 7); odour pron ounced , pen etratin g , 'earthy; pale ycllow- coloured
pigm ent diffusing into th e surroundin g agar ; colony rcversc (colour plat o
164 , a) in yellow 01' vinaceous shades ; co n id ia l struc turcs (li g . 164 , A , B and
C ) as in th e othe r vari eti es.
C oloni es upon C YA (colour plat e 164 , B an d b) a tta ining a di ameter 01'
50 mm in 14 days at room temperature (20 - 25 OC), cons isting 01' a tough , wet
basal felt , cove re d b y a white floccose ove rgrowth 01' ae rial hyph ae and rop es 01'
h yphae , very poorl y spo ring; exudate a nd odo u r lacking; yellow coloured
pi gm ent d iffu sing int o th e surrounding ag ar ; colony rever se (co lou r plat e
164 , b) d eveloping yellow ish to reddish brown shade s .
C oloni es upon M EA (colour piate 164 , C a nd c) attain ing a d ia me te r 01'
45 mm in 14 days at ro om temperature (20 -25 OC), ge ne ra lly plane exce p t
slightly raised in th e cen tre, loose textured with a tcnden cy to become floccose ,
more 0 1' less za nate in margin al area s, co m mo nly becoming definitely
synnematous in age, spo rula ting a bundantly in recent isolat es , with
colouratio n in lighter shade s th an upon C A 0 1' C YA; in olde r strains poorly
spo ring and sho wing ab u ndan t ropiness (co lour piate 164 , C ); odo u r m ou ldy ;
exu date not produced ; no colou re d pi gm ent diffusing into th e su r ro u nd ing
agar; colon y reverse (colou r plat e 164 , e) colou rless .
The variety descr ipti on is ce ntere d upon th e typ e strain CBS 246. 32 = C M !
92, 264 = N R R L 3, 971, originally isolat ed b y G . Bainier , and up on R ap er and
Thom ' s notes (1949) that ade q uately present th e species whe n recen tly
isolat ed . The ty pe strain , al'ter so m any years under con tin uo us la boratory
cultiva tion ha s lost m any 01' its typi cal cultu ra l characteristics and ha s beco me
more pred ominantly funi cu lose and poorl y spo ring .

165 . Penicillium verrucosum Di erckx val'. corym biferum (Westling) Samson ,

Stolk el H adlok: fig. 165, colour plat e 165 , and SEM plate 28,
(centre : ri ght) .
Pencillum verrucosum Dierckx varo corymbferum (Westling) Samson, Sto lk et
Hadlok - Studies in Mycology. 11: 36-37. 1976.

Synonymies
Pencillum hrsu tum Dierckx - Ann. Soco Sci. Bruxelles. 25: 89. 1901 .
Pencllum corymbferum We stl ing - Ark . Bot. 11: 92. 1911 . See also Raper and
Thom, ' A Manual of the Penicillia', pp. 540-544. 1949.

C olonies upon C A (co lo u r plat e 165, A and a) gro wing rather rapidl y ,

Fig. 1650 Camera lucida

drawings of Penicillium
verrucosum Dierckx var o
corymbiferum (Westling)
Samson, Stolk et Hadlok,
CBS 135.41. A , Detailed
drawings of the penicill i. B,
Mature conidia . e, Habit
sketches .

at ta mmg a d ia meter of 60 - 70 mm in 14 days at room tc m pe rature

(20- 25 OC), u su all y za nate in m argin al areas , strongly fascicul at e , espec ially
in rece nt isolat es, with su rface appearing coa rsely gran ula r or rid ged and with
the m ajorit y of conidiophores aggregated int o clearIy defin ed syn ne mata;
heavily spo ring thro ughou t , developing pale yellow, pa ssin g throu gh yellowgrec n near pea gree n to sa ge green sha des with th e rip ening of con id ia, a nd
beco m ing sla te-olive gree n in age (Sguy, PI. X X IX, 434) ; with thin m argin
surrounde d by a wide frin ge of sub merge d h yphae ext ending about 5 mm
beyond th e ae rial growth; odo u r faint or lacking; exudate abu ndan tly
produce d as large brown drop s; yellow-coloured pigm ent diffusing into the
surrou nd ing a ga r; colony reverse (colour plat e 165, a) in deep purple brown
shade s; colony surface cove re d by a n ove rgrowth of yellow-coloure d vegetative
hyphae ; peni cilli csse ntia lly as in th e other varieties but with con id iophores
variable in len gth , ranging from 100 to 200 11m or m or e at th e m argin up to
1000 11m in th c stro ngly fa sciculate ce ntral areas (fig . 165, A , H and C), with
wall s closely a nd cons picuo usly ro u ghe ne d.
Colonies upon C YA attaining a di ameter of 60 mm (colour plat e 165, B and

b) in 11 days a t room tcmpcratu rc (20 -2!i OC), a ppca ri ng more loccosc an d

lcss fascicul at e than upon C A, with a wid c whit c m argin su rro u ndc d by l
frin ge 01' submerged hyphae exte nd ing about :; mm bcyon d th e ae r ia l growt h;
heavily spo r ing in radial sec to rs in deep blue- green sh ade s (Sgu y , PI. X XX ,
446), e1sewhere in pal e ye llow sha des; a m be r-coloured pi gment d iffu sing in to
th e su rrou n d ing agar ; colon y reverse in deep purpl e b rown sh ad es. (see colou r
piat e 165, B a nd b) .
Colon ies upon MEA (colour plat e 165, C a n d e) a tta in ing a di amet er 01'
45 mm in 14 d ays a t room temperature (20 -25 OC), cons isting al' a looser
textured b a sal felt , com pa ratively thin , plane excep t 1'01' a limited floccose
ove rg rowth all ove r th e colon y surface cons isti ng a l' ye llow ae r ia l hyphae , less
fasc icu la te than upon C A, m ore in tensely gree n n ear aspho del 0 1' pea gree n
(Sguy, PI. XX , 299); n o coloured pigment diffu sing into th e su rrou nd ing
agar; ex u date a n d odo u r lacking ; colon y rev erse (colou r plat e 165, c) in pale
re dd ish b rown shades .
T he variety description is ce ntered upon st rain CBS 143. 45, isolat ed fro m
Hyacinthus sp . a t th e London Sch ool a l' H ygiene a n d Trop ical M edicin e , 1945,
an d designated as the neot yp e , a n d upon st rain CBS 135 .41 = CMI 40 ,213 = ATCC
10,429 = N R RL 2, 032= IFO 6,092 = IJFM 7110 , isolat ed b y P . Beis in 1941 from
green fly in the Nethe rla nds.
Penicillium verrucosum val'. corymbiferum is ge ne rally accepted a s the ca usal
age nt a l' Penicillium rot a l' liliaceou s bulbs. It ca n b e di stinguish ed frorn th e
other varieties by th e ye llow-green colour al' its colonies and by the ye llow
vegetative h yphae . M oreover a brown exu date is usu all y p roduced a nd th e
phialides a re co m paratively larger , co m monly 15-1 6 p.m by 3- 4 ust, in stead
al' th e 7- 9 p.m by 2. 0-2.5 p.m u su all y fou nd in the othe r va rie ties.

166 . Penicillium echinulatum Fassatiov : fig . 166, co lo ur pl a te 166, a n d

SEM pl ate 28, (bottom : left).

Penicil/ium echinulatum Fassatiov - Acta Univ. Carolina Biol. 1974: 326-327. 1977.
Synonymies
Penicil/ium cyclopium Westling varo echinulatum Raper et Thom - 'A Manual of
the Penicillia', p. 497. 1949, (without a Latin diagnosis).
Penicil/ium crustosum Thom varo spinulosum Sasaki - J. Fac. Agric . Hokkaido
(imp.) Univ. 49: 158. 1950, (without a Latin diagnosis).
Penicil/ium palitans Westling var. echinoconidium Abe - J. Gen. Appl. Microbiol.
Tokyo. 2: 111-112. 1956, (without a Latin diaqnosis) .
Fassatiov's original diagnosis follows, since this adequately presents the species
in question:

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Fig. 166. Camera lucida

drawings of Penicillium
echinulatum Fassatlov. CBS
317.48. A, Detailed drawings
of the penicilli . B, Mature
conidia. e, Habit sketches.

'The colony is mealy, granulose or fasciculate on Czapek's agar, the conidiabearing layer being blue-green or yellow-green. The margin of the colony is white ,
1-2 mm wide. The colouration of the reverse side is yellow-orange, yellow-brown
to orange -brown and the pigment does not penetrate the agar . The exudate clear,
occurring in abundance. The conidiophores measure 50-400 Jlm in length, 3-3.6
Jlm in width, with a' granular or roughened surface. Sometimes the surface is
smooth. Branches of 11-24 and 3.4-5 Jlm. The height of the penicillus is 35-50 Jlm
(or more), that of the small penicillus (metulae + phialides) 17-24 Jlm. The phialides
are 7-11 .5 by 2.2-3 Jlm in size. Conidia globose with a thick wall, roughened or .
echinulate, 3.5-5 Jlm in diameter.
The original strain was obtained from Ottawa, Canada as a contaminant of an
agar medium (Raper and Thom 1949). In Abe's strain (1956) the origin is not given .

O ur not es follow :
Colo nies upon CA (colo ur pl at e 166 , A a nd a) a tta ining a d ia me te r of
45 m m in 14 days at room tem perature (20 - 25 OC ), consistin g of a basal felt
com posed of mostly monon em atou s co nidiopho res in a den se stand, with
loosely synnem a tou s conid iopho res in marginal areas, lending a gran ular

a ppcarance lo th e co lony su rface whi ch is almost e n tircly covcred b y a dcc ply

Iloccose overgrowth 01' whit c ac ria l hyph ae grcat ly inllucn cin g th e genera l
as pec t of th e colon y , whi ch a ppears as dccply lanat c ; hcavily spori ng
und ernea th th e floccose ove rg ro wth in gray-grccn shades (Sguy, PI. XXVII,
402) ; odour strong suggestive a l' Beta vulgaris L. aroma 0 1' ' earth y ' ; cxudu to
abu ndantly produced as large d eep oran ge -colou re d drops ex tr ud ing fro m th c
floccose felt ; p ale yellow-colou rc d pi gment diffu sing into th e su rro u ndi ng agur:
colony reverse (co lou r pla te 166 , a) developing ora nge to cin namo n shadcs :
penicilli (fig . 166 , A) cons isting al' con idio pho res m ostly m onon ernatous but
loosely sy n nemato us in marginal areas , rough walled , one a l' two stage
branch ed , with ra mi and metulae closely appressed to th e m ain a xis, wi th
sti pes ran ging from 150 to 400 11m in length b y 2 .5 - 3 .0 11m in di amet cr ; ra m i
about 12 - 20 11m by 3 .0 11m; m etulae in small ve r ticils a l' 2-3 ele men ts, eithcr
rou gh al' smooth walled, cylin d rical , 13- 20 11m by 2.8 -3.8 11m ; phialidcs
b ottle shaped, in clus te rs ol' 4 -8 ele men ts in th e verticil, about 11-1 5 11m by
2.5 -5. 0 11m , tape r ing to a sho rt co ns p icuous neck ; co n idia globose In
subglobose, co ns picuo us ly echinu la te, 4.5- 5 .0 11m (fig. 166 , R , a nd SEM
plat e 28, bottom : left) con id ia l chains loosely di vergen t (fig. 166 , C).
Colon ies upon CYA (colou r pl a te 166 , B a n d b ) sprcadi ng broadly, a ttain ing
a di amet er ol' 60 mm in 14 da ys at room temperature (20 -25 OC), co ns isti ng 01'
a close-t extured basal felt , stro ngly radiat ely wrin kled, w ith a th in whi tc
m argin su r rou n de d by a frin ge a l' submerge d h yphae ex te nd ing a bo ut 3 m m
beyond the ae r ial growth , with su rface growth, loosely ove rlaid b y a floccose
network a l' white ae r ial h yphae ; conspicuous ly za nate a nd hea vily spor ing
throughout in deep green shades (Sguy, PI. X X XIII , 486, 493) w ith th e
ri penin g al' con id ia, particul arly in m argin al a reas; odo ur strong, as d escribed
aboye; exu date ab u n dan tly produced as aboye; no coloured pigment diffusing
into th e su rrou n d ing agar; colony reversc de ve lopi ngreddish cin namo n shadcs
(colour pla te 166 , b ) ; penicil1i essentia lly as d escri bed a boye .
C olon ies upon M EA (co lour plat e 166 , C a n d c) at tain ing a di amet er 01'
abou t 35 mm in 14 d ays at room te mpe ratu re (20-25 OC), cons isting of a
looser textured b asal felt , zo nate, plane , sligh tly umbon at e in th e cen tre ,
appearing gran u lar, witho u t a ny floccose ove rg ro wth; wi th undulating margin
su rrou nde d by a narrow l'ringe al' submerged hyphae ex te n d ing a bou t 1 mm
beyond th e aeria l growth; hea vily spo ring th ro ughou t in deep green to oli ve
green shades (Sguy, PI. X XVII , 402); odo ur strong, aromatic but indefinite;
no coloured pi gmen t d iffusing in to th e su r ro u nd ing agar; exu date not
produced ; colo ny reverse (co lou r pla te 166 , e) develop ing reddish oran ge
shades in th e ce ntre, elsewhe re almost co lou rless wi th a green cas t showing
through ; penicilli esse ntially as described a boye, with con id iophores a r ising
prim arily frorn the substratum .
The spec ics d escription is ce n te red u pon th e typ e st rain CBS 3 17. 48 = C M !

10,02B - ATCC IO ,4:H "NI~I~ L 1,1 51 -IJI'M 7 163, as reccivcd from th e

Ccntraalbu rea u voor Schim me lcultu rcs , Ba arn , The Nethcrl ands , in 1979 ,
o riginally isolat ed in 1940 by G .A . Ledingham , Nati onal R esearch Counc il,
O ttawa, C a na da, as a cu ltu re con ta mina nt.
In our ob servation Penicillium echinosporum grown up on C A differ s fro m
Fassatiov's original description only by th e de eply floccose overgrowth of
white ae rial hyph ae a nd th e deep ora nge-colou red exu date (colou r plat e
166 , A).

800 0
0000
0 0

10jJm

ooo~ o\
O
L...--....J

100 jJm

Fig. 167. Camera lucida

drawings of Penicillium
verrucosum Dierckx varo
cyclopium Samson et al.
strain ananas -olens Ramrez,
IJFM 3865. A , Detailed
drawings of the penicilli . B,
Mature conidia. e, Habit
sketches.

168 . Penicillium expansum Link ex Gra y e me n d. Thom. fig. 168 , colour

pl at e 168 , a n d SEM pl ate 28 , (botto m : r igh t) .
Penicillium expansum Link ex Gray emend. Thom - Mago Ges. Nat. Freunde Berlin
3: 17. 1809. See also Thom, in Bull. Bur. Anim. Indo U.S. Dep . Agric . 118 : 27-28 .
1910. Also Thom , 'The Penicillia', pp. 402-405. 1930. Also Raper and Thom , 'A

Manual of the Penicillia' , pp. 512-516. 1949. See also Samson et al. Studies in
Mycology 11: 20-22. 1976.

at.

Fig. 168. Camera lucida

drawings of Penicil/ium
expansum Link ex Gray
emend Thom, CBS 150.45.
A, Detailed drawings of the
penicilli. B, Matu re conida.
C, Habit sketches.

Synonymies
?Penicillium glaucum Link - Mag. Ges. Nat. Freunde Berl. 3: 17. 1809; ex Pers. Mycol. Europ. 1: 40. 1822.

Penicillium crustaceum (Linneausl Fries - Syst. Mycol. 3: 407. 1832.

Coremium alphitopus Secretan - Mycol. Suisse 3: 539-540. 1833.
Coremium vulgare Corda - Prachtflora, p. 54. 1839.
Penicillium leucopus (Persoon) Biourge - C. R. Soco Biol. Paris. 82: 877-880.
1919.

Penicillium elongatum Dierckx - Ann. Soc o Sci. Bruxelles 25: 87. 1901.
Penicillium juglandis Weidemann - Z. Bakt. Parasitenk. Abt. 2, 19: 683. 1907.
Penicillium variabile Wehmer - Mycol. Z. 2: 195. 1913, and Ber. Bot . Ges. 31:
310. 1913. (Not Penicillium variabile Sopp - Videnskapselskapets Skrifter
Kristiania Math. Nat. KI. 11: 171. 1912).

Penicillium plumiferum Demelius - Verh. Zool. -Bot. Ges. Wien. 72: 76. 1923.
Penicillium malivorum Ciferri - Riv. Patol. Veg. Padova. 14: 77. 1924.
Penicillium resticulosum Birkinshaw, Raistrick et Smith - Biochem. J . 36: 829.
1942.

C olonics upon C A (colour plal c 16B , A and a) gro wing rapidly, a uaining a
di ameter 01' abo ut 50 mm in 14 days at room temperature (20 - 25 OC), with
surface plane 0 1' gene ra lly sho wi ng rad ial fur rows , 01'ten ranging frorn 0.5 to
~. O mm deep , with a thin , irregul arl y di ssected, white m argin surrounded by a
tringe 01' su b me rge d hyphae ex tend ing about 2- 3 mm beyond th e ae rial
g-rowth ; ve ry heavily sporing through out with con id iophores very abundant
a nd primarily a rising from th e su bs tratu m, in sorne strains in an alm ost
continuous dense sta nd, in others (colour piate 168, A) sho wi ng conid iopho res
almos t enti rely gro u pe d into fasc icles 0 1' synne mata with colony su rface
ap pearing gran ula r 0 1' showing conspicuo us zonatio n; conid ial areas in yellowgree n to sage gr een sha des (Sguy, Pis XXVII, 403, a nd XXVIII , 417) with
the ripening 01' co nid ia; exudate limited in a mo u nt, mostly as small dropl et s
cnrneshe d in th e m ycelial felt , colou rle ss; odo u r strong, mouldy , characteristic
01' rott in g a pples; reverse almost colou rless in sorne stra ins throu gh yellow brown to red di sh ora nge in othe rs (colour plat e 168, a); am be r-coloured
pigm ent diffusing in to the surround ing agar produced in sorne strains, not in
ot hers ; conid ial structures ve ry abundant an d , in m a ss, characterizing th e
colony , with con id iop hores m ostly 150-400 p.m in length in sorne strains, but
commonly 600 -700 p.m in othe rs, by 3-4 tut: in di ameter with wa lls smooth 0 1'
de lica tely rou gh ened , term inati ng in large penicilli usu ally measuring u p to
75- 100 uu: in length , typica lly asymmetrica l, two - 0 1' three-stage branch ed ,
with rami m ostl y 20-25 uit: by 2.5 -4 .5 tui , occasionally up to 60 p.m in
length , typi call y appresse d agains t th e main ax is; m etulae bo rne in verticils 01'
2-6 eleme nts, abo ut 10- 15 p.m by 2.5-3.5 p.m ; phialides bo ttle shape d, in
cluste rs 01' 2- 8 ele me nts in th e verticil, ranging fro m 8 to 12 p.m by
2.3-3 .0 tui , tapering grad ually to a conspicuo us short and narrowed nec k;
conid ia ellipt ical whe n first formed , th en becom ing su bg lobose, 3- 4 p.m in
long ax is, with walls smooth 0 1' ne arly so (fig. 168, and SE M plat e 28, bott om :
right).
Colonies upon CYA (colour pl ate 168, B and b) attaining a d iame te r 01'
abo ut 65 m m in 14 days at roo m temperatu re (20-2 5 OC ), sim ila rly textu red
as upon C A but m or e loosely synne matous and gra n ula r , rad iately wrinkled
and heavily spo ring throu gh ou t , especially in m argin al areas, in sage green
colours (Sguy, PI. XXVII , 403), su bmarginal a nd central areas with conid ial
layer obscu red by a floccose overgrowth 01' whi te ae rial hyph ae ; odo u r stro ng,
as aboye ; exu date limited in amo u nt, colou rless, 0 1' lacking ; am ber-colou re d
pigm ent di ffusin g in sorne strains, not in ot he rs ; colon y reverse developing
deep reddish orange shades (co lour plate 168, b) ; penicilli esse ntially as
described aboye .
Colon ies u pon MEA (colour plat e 168, C a nd e) attain ing a d iameter 01'
abo ut 65 m m in 14 days (20 -25 OC), cons ist ing 01' a loose- textured ba sal felt ,
pla ne, not furro wed , conspic uo us ly zonat e , with very th in margi n su rro u nded

by a wid e Irin gc 0 1' su b me rgcd hyphac cx tc nd ing about 7 01' more mm bcyond
th e a crial g ro w th ; heavily spo r ing throu ghout in gray-green shades (S guy , PI.
XXIX, 429), with conid iophores a rising dircctly Iro rn th c substraturn in
definite fascicles a rrange d in conspicuo us conce ntrical zon es; ex uda te lacking;
odou r strong , as not ed abo ye; no coloured pigm ent diffusin g int o th e
surro und ing agar; colony rev erse (colou r plat e 168 , e) colou rle ss; penicilli
esse ntially as described aboye .
The spec ies description is ce ntere d upon th e neotyp e stra in CBS 325. 48 = CMI
39,76 1 = N R R L 976 = ATCC 7,861 = IJ FM 5 14 1, isolated frorn apples by j .S.
Cooley , Bureau 01' Plant Industry , US Dep . Agri culture , 1926 , and m any
othe r strains isolated by the a ut ho r fro rn rotten apples in M adrid .
PeniciLlium expansum is one 01' th e oldes t and best known spec ies 01' PeniciLlium,
cons titutin g, with P. glaucum Link and P. candidum Link, th e three spec ies upon
whi ch H .F. Link esta blishe d in 1809 th e genus PeniciLlium.
Hundreds 01' stra ins bel ongin g to thi s spec ies hav e been isolated from
naturally infected products. They a re com mo n in or on organ ic m at erial s in
contac t with th e soil; they occasi on ally occ u r on grains and ce real products,
including bread . They have been isolated from eggs and m eat products in cold
storage, a nd fro m th e waste su lph ite liquors 01' th e wood pulp industr y . They
are occasionally isolat ed from fabrics and othe r deteri or ating military
equ ipme n t. They are, however, m ost com mo n a nd m ost characte ristica lly
isolat ed fro m a pples and othe r pom aceou s fru its, wh er e th ey typi cally produce
a rapid soft brown ro t, hi ghl y destru cti ve , com mo nly referred to as ' blue moul d ' rol. The spo res 01' th e fu ngus a re ca rried from th e soil on th e surface 01'
th e fruit , germ inating and ente ring throu gh cuts and abra sions in th e cuticle
a nd epide rm is .
When tested in th e laborat ory by piercing a nd inocul at in g sound apples,
typ ical strains 01' PeniciLlium expansum will produce areas 01' rot from 30 to
40 mm in di amet er , exte nd ing inward to th e core, wi thin a peri od 01' 8 - 10
days.

169 . Penicillium italicum W ehmer val'. italicum Samson , Stolk et Hadlok :

fig. 169 , co lo u r pl ate 169, and SEM pl ate 29, (to p: left) .
Penicil/ium italicum Wehmer var o italicum Samson, Stolk et Hadlok - Studies in
Mycol. 11: 28-30. 1976.

Synonymies
Penicil/ium italicum Wehmer - Hedwigia. 33: 211. 1894. See also Beitr. Kennt .
Einh. Pilze, Abt. 2, pp . 68-72. 1895. Also Thom, 'The Penicillia', pp. 412-414.
1930. Also Raper and Thom, 'A Manual of the Penicillia', pp. 526-529. 1949.
Penicil/ium aeruginosum Dierckx - Ann. Soco Sci. Bruxelles. 25: 87. 1901.
Penicil/ium ventruosum Westling - Ark. Bot. 11: 112. 1911.

'1-/:1

lJJ.illlluJ

10J.lm

C)

O O
0

0
B

O
Ot:0

OOCJO
0

00

C)

y~?t~
lOo~Jl

Fig. 169. Camera lucida

drawings of Penicillium
italicum Wehmer varo
italicum Samson et al.,
CSS 136.41. A, Detailed
drawings of the penicilli.
S, Mature conidia. C, Habit
sketches.

Colonies upon CA (colour plate 169, A and a ) growing very restrictedly ,

a ttain ing a di ameter of 20 mm in 14 da ys at room temperature (20- 25 OC ),
consisting of a loose-textured basal felt, with ce ntral are a 1.0-1.5 mm wid e
commo nly raised up to 2 mm deep , m arked in sorne stra ins by shallow
furrows, with m argins usu all y thinner surrounded by a fringe of deeply
su bme rged hyphae ext ending abou t 1-2 mm beyopd/ th e ae rial growth, in
sorne stra ins almost plane but showing evide nce of fascicul ation , in othe rs
stro ng ly fasciculate , with fasciculation arrangd in conce n tric circles (colour
plat e 169, A) , sp orulating irregularly , mor e or less abundantly , with con id ial
are as typi cally in pal e gra y-gree n in sorne strains , in othe rs more yellowish
gree n near pea green (Sgu y , PI. XXIX , 422) or blue-gr een sh ad es (S guy, PI.
XXXI, 462) ; odour fragr ant, suggesting perfume; exudate lacking or very
limited in amount ; pale amber- coloured pigment di ffusin g into th e
surrounding agar in sorn e strains, not in othe rs; colony rev erse (colour plate
169 , a) colou rless to orange-brown ; conid iophores sometimes mononemat ous ,
more freque n tly loosely synne matous, especiall y in th e marginal a re as; the
synnem ata sometimes ari sing from bundles of ve geta tive hyphae submerged in
th e substratum, va riable in length up to 1 mm or more , in sorn e strains,
com mo nly exte nd ing out seve ral millimeters beyond th e colony margin bel ow

't 11>

the surface 01' th e agar bc fore assuming un cree ! or ascc nd ing position (colour
plate 169, A) ; peni cil1i asymmetric (fig . 169, A), with con id iophores sm ooth
wal1ed , two-stage branched , usu ally with rami closcly a ppressed; stipes
ranging from 100 to 250 flm by 3.5 -5. 0 flm ; ra m i usually m ca suring
15-2 5 flm by 3.5-4.5 flm ; m etulae mor e or less cylind rical, borne singly or in
grou ps 01' 2-4 cle me nts in th e ve rticil, abo u t 15-20 tu by 3- 4.5 tui , bu t
frequentI y larger or smaller , sometime s ari sing a t different levels in the
peni cillu s; phialides slende r, mor e or less cylind rical, with short but
cons picuo us neck , com mo nly 10-1 3 flm by 4 flm and not infrequentI y up to 20
or m ore flm in length; con id ia typi call y cylind rical wh en young but be coming
definitely ellipt ical or eve n su bglobo se at m aturity , ex tre mely va riable, but
generally 5-6 flm by 3.0 -3.5 tun , with indi vidual spo res wh en elliptical
ranging u p to 9 flm by 5. 0- 6.5 tui , with wall s smooth or nearly so (fig . 169, B,
and SE M plate 29, top: left ) ; conid ial chains ta ngled (fig . 169, C ).
Colon ies upon C YA (colour plate 169, B and b) spre adi ng bro adl y , a ttaining
a di amet er 01' abo ut 65 m m in 14 days at roo m temper ature (20-25 OC ),
consis ting 01' a close-t extured b asal felt , cons picuo us ly za nate a nd heavily
spo ring , with conidio pho res aggregated in to well-de fined synne mata arrange d
in conce ntric zones; coni d ial areas in dee p gree n shades (Sg uy, PI. X X IX,
422); exudate limi ted , colourless; od our pro no unce d, ' mo uldy'; am be rcoloured pi gment d iffus ing int o th e su rrou nding agar; colony reverse (colour
pIate 169, b ) cons picuo us ly zonate, develop in g dark b rown zones alte rnating
with reddish bro wn arcas; penicilli esse ntially as desc ribe d above .
Colon ies upon MEA (colou r plat e 169, C a nd c) attain ing a diamet er 01'
abo u t 55 mm in 14 days at roo m tempera ture (20 -25 OC), cons ist ing 01' a
loose-textured basal feIt , cons picuou sly za nate ow ing to the fascicul at e conid ioph or es , with a very th in m argin su rro unde d by a wide fringe 01' submerged
hyphae exte nd ing abo ut 5 mm beyond th e aerial gro wth; heavily spo ring
th ro ughout in th e sa me green shades as above; odo ur strong, aro matic,
suggesting perfume; exudate not produce d; n o colou re d pi gm en t diffusin g into
th e surrounding agar; colony reverse (colour plate 169, e) cons picuous ly
zonate, developing da rk b rown colou rs; pen icilli esse nt ially as described
above.
The spec ies description is ce ntered upon th e neot yp e strain C BS 339.48 = C M I
39,760 = ATCC 10,454 = NRRL 983, isolated by Dr H .S . Fawcett, Ci trus
Experiment Station, Ri versid e , California, fro m citr us fruits, in 1930, a nd
m an y othe r strains isolat ed fro m citru s frui ts in Spain, by th e aut ho r. (colour
pIate 247 presents a representat ive strain 01' P. italicum var. italicum isolat ed by
th e autho r fro m Cayenne pe pper.)
Penicillium italicum va r. italicum is rarely found in n ature except on citr us
fruits, wh ere it ca uses a characteristic soft rot, appeari ng in advanced stages 01'
fru it dec ay as a bluish green m ould cove ring th e rind as a velvety a l' tu fted

~ 77

lnycr 01' powdcry spores. The rol ca use d by thi s spccics is clearly distin ct Irom
rluu ca use d by P. digitatum Sa ccardo . Both sp ecies m ay occ ur o n the sa me
individual fru it. The char ac te r ist ic so ft rot is generall y referred lo as ' b luemould ' citrus rot. P. italicum val'. italicum ca n be distinguish ed from P. digitatum
by th e stru cture a n d colo ur o f their co lo n ies, those of th e former are
synnem at ou s a n d blue- green , while th ose of the latter a re thin , velvety a n d
olivaceou s . M oreover , P. italicum gro ws better upon C zapek ' s agar th an P.

dZttitatum .
170. Penicillium italicum W ehmer v a l'. avellaneum Samson et G u tter: fig .

170, colour plate 170, a n d SEM plate 29 , (top : right).

-!{~\

,..\;1j

fR .
~

100j.lm

Fig. 170. Camera lucida

drawings of Penicillium
italicum Wehmer varo
avellaneum Samson et
Gutter, CBS 719.73.
A, Detailed drawings of the
penicilli. B, Mature conidia.
e, Habit sketches.

Penicillium italicum Wehmer varo avellaneum Samson et Gutter - Studies in

Mycology. 11: 30-31. 1976.
Colo n ies upon CA (co lou r pl ate 170, A a n d a) grow ing restrict edly,
a ttaini ng a di amet er of abou t 10-1 2 mm in 14 d a ys a t room ternperature
(20-25 OC), co ns isting of a close-te x tu re d b a sal felt , ap pear ing velvety ,

11"{11

some timo s obscurcd by whit c locc ose acri al hyphae es pec ia lly in th c colon y
ce ntre; co n id ia l a rcas pale brown, near cinnarnon, becorning d arker, nca r
ave llane ous (Sgu y , PI. IX, 134) ; odour aromatic; exu d ate not produ ccd ;
conid iopho res mon on ematous 01' syn ne mato us, with synnemat a especially
present in m arginal areas, ofte n d evel oping from bundles of submerged
hyphac ; co ni d iophores smoo th walled , at least in th e upper parts, two-stage
b ranched (fig . 170, A), with rami usually elos ely a ppressed; with penicilli
duplicat in g in det ails of mi cr oscopi cal morphology , in size and shape those of
th e va l'. italicum; con id ia cylind rica l, sometimes ellipso idal, smooth walled (fig,
170, B, a nd SEM plat e 29, top : ri ght) , pal e brown in m ass , 4. 5. ,tm by
2.8- 4.0 ustv ; no coloured pi gmcnt diffu sing into th e su rrou nd ing agar; colon y
rever se ve ry dark violet, almost bl ack colou red (colour plat e 170, a); conid ial
chains loo sely tangled (fig . 170, C).
C olonics upon C YA (colour plat e 170, B and b), sp read ing broadly,
att aining a diamet er of abo u t 65 - 70 mm in 14 d ays at room temperature
(20- 25 OC), co nsisting of a dense basal felt , plane in margin al areas and
stro ng ly furrowed in th e cen tre in a radial pattern , with a thin margin
surrou nde d by a fringe of su b merge d hyphae ex te nd ing a bo u t 3 mm beyond
th e ae rial growth ; heavily spo ring th rou ghout in avella neous shades, show in g
sorn e zonat ion owing to th e syn ne matous co nid iopho res arranged in co ncen trical cireles; odo u r aro matic; ex udate not produced ; am ber-colo u re d
pi gm ent diffu sin g into th e surro unding agar; colony reverse (colour pl ate
170, b) in de ep reddish brown shades; penicilli essentially as described abo ye
for th e val'. italicum .
Colonies upon M EA (co lo u r plat e 170, C a nd c) attaining a di amet er of
abo u t 55 mm in 14 d ays at room tempera ture (20 -25 OC), co ns ist ing of a
looser textured basal felt , a p peari ng velvety to ' mcaly', plane , not furrowed ,
very sligh tly umbon ate in th e cen tre, with very thin m argin su r ro u nde d by a
frin ge of su b me rge d hyphae ex te nd ing abo u t 3- 4 mm beyond th e ae ria l
gr owth ; heavil y spo ring th rough out in th e sa me shades as a boye ; ex udate not
produ ced ; odo u r aro matic; no coloured pi gm ent d iffus ing in to th e surrou ding
agar; colon y rever se (colour pla te 170, c) d evel opin g yellow ish shades;
con id ioge no us structures identical with th ose on C A and CYA .
T he va riety description is ce n te re d upon the type stra in CBS 719. 73 = IJ FM
7193, isolated from Citrus sp. in Bet D a gan , Israel , by Y . G u tte r in 1973.
The varie ty diffcrs fro m th e val'. italicum m ainly b y th e avella neou s colour.
V al'. avellaneum ca n be poten tially a serious post-harvest ca use of infecti on
and su bse q ue n t decay of citrus fru its.

Penicillium italicum W ehmer val'. album W ei:

Nanking J. 9: 241. 1940 .

Penicillium italicum W eluncr val'. al/mm W ei is a un gu s dcscribcd in 1940 th ut

is helicved lo be a whitc vari cty 01' P. italicum W ehmer. H ow cvcr, it has not
Ilee n includ ed in the kcys to spc cies since no typ e cu ltu re was av a ila ble 1'01'
rnmpari son with th e aboye describcd va ricties, a nd all a tte m pts to obta in it
1'J'() 1II 1nt ernati onal C ulture Collections fail ed .
We i's original di agn osis follows:
'The fungus produces vigorous white mycelium on agar medium. Both velvety
fruiting layer and coremia were formed with the latter forming a ring near the
center. Fructification are al! pure white. The morphology of the conidiophores
resembles closelv that of Penici/lium italicum with sterigmata (phialides) measuring
8.4-12.6 by 1.8-4.2 .tm. Conidia are ovate to oblong, mostly ovate, measuring
3-8.4 by 2-5.7 .tm.
The present fungus is, however, a mutant of P. italicum to which it agrees both in
morphological characters and reaction to ternperature.'
(The aboye description was presented by the author without providing a Latin
diagnosis) .
The type cultu re 01' Penicilli um italicum va ro album We i wa s isolated in 1940 by
C .C . W ei fro m decayin g sweet oranges in Szechuan Pro vin ce , China , a nd is
pr ob abl y lost. It h as been cited by R aper and T ho m (1949), Kulik (1968), and
by Samson el al. ( 1976).
171 . Penicillium hispalense R amre z el M artnez: fig. 171 , colo ur plate
171.
Penici/lium hispalense Ramrez et Martnez - Mycopathologia. 74: 163-171. 1981.

Colonies upon C A (colour pIate 171, A and a) gro wing rap idly , attaini ng a
d iameter 01' abou t 45 mm in 14 days a t roo m temperature (20- 25 OC ) ,
consisting 01' a dense , stro ngly syn ne matous, felt , with synne mata a rrange d in
a circula r pattern, with colony surface appearing coarsely gran ula r and with
th e great maj ority 01' conid iopho res aggregated into c1early defin ed fascic1es,
1- 2 mm deep in central and subcentral areas, not furrowed , with thin m argin
su rrou nde d by a fringe 01' su bmerged h yphae extending abo ut 3- 4 m m beyond
the ae rial gro wth; hea vily sporing throu gh out in deep blue-gr een shades
(Sguy, PI. XXX , 447) with th e rip ening 01' con id ia; exudate lacking ; odo ur
fragra nt, aro m atic; pale yellow-coloure d pigm ent slowly di ffusin g in to th e
surrou nd ing agar; colony reverse (colour plat e 171, a) developing yellowish
sha des with a green cas t sho wing throu gh ; penicilli (fig . 171, A) typi call y
as ym me tr ic, with conid iophore s coarsely rough en ed , inc1udin g the first rami ,
the ot he r ele me nts a re smooth wa lled, usu all y two- , three- 01' mo re stage

:.~. !

o
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0 0 00
0 0 0
O 00

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r

/......

."
~ :/
"

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.'

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....

fR.,

Fig. 171. Camera lucida

drawings of Penicillium
hispalense Ramrez et
Martnez, IJFM 5940.
A , Detailed drawings of the
penicilli. B, Mature conidia.
Habit sketches.

e,

branched , with all eleme nts multisep tat e so as to m ake its differ entiati on difficult ; all eleme nts of th e pen icillu s are closely appressed; stipes of th e
conidiophores va ry ing in length , rangin g from 50 to 200 p.m in len gth by
4.0 -4 .5 uit: in di ameter; b ranc hes or ram i va ria ble in form and size, usu ally
becoming small er at successive levels away fro m th e m ain ax is; me tulae not
readily di stingui shabl e from th e u pp ermost rami , m ostly 4.0-11 .0 p.m by
3.0 - 5.0 p.m ; stipes and first rami appearing cons picuo us ly warty , with
rem aining conid ial struc tu res smooth walled ; phialides bottle shape d,
com paratively sm all , mostl y 6.0-8 .0 p.m by 2.5 -3 .0 psi , with a conspicuo us
bu t sho rt neck ; conid ia globose to su bg lobose, 3.0- 4.0 p.m by 2.5- 3. 5 p.m,
smooth wall ed or nearl y so (fig. 171, B) .
Colonies upon C YA (colour plate 171 , B a nd b) attaining a di amet er of
55 mm in 14 days at room temperature (20- 25 OC ) , sim ila r in texture and
colony appearance to those gr own upon C A, but showing more abu ndant
fascicul a tion , with colony su rface radi at ely furro wed and za nate in m argin al
areas; h eavil y spo ring th roughout in blue-gr een sha des (Sguy , PI. XXX ,
447 ) ; exudate and odo ur lacking ; pale amber-coloured pigment diffusin g in to

the surrou nd ing aga r ; colo ny reverso (colour plale 171, b) developin g yellowi sh
cin narnon shadcs; peni cilli essc n tially as cl cscribcd aboye .
Colonies upon M EA (co lou r plat e 171, C and e) a tta in ing a diamet er of
55 mm in 14 d ays at room temperature (20-25 OC ), cons istin g of a loosetextured ba sal felt , co ns picuo usly zonate ow ing to th e co nce n trica lly a rra nged
syn ne mata; heavil y spo ring throughout in deep blue-gr een shades (Sguy , PI.
XXX, 446); ex udate lacking ; odo ur fragrant , fruity ; no colo ured pigment
diffusing into th e surrounding a gar ; colony reverse colou rless with a yellow
cast showing th rou gh (colour plate 171, e) . Penicilli essen tially a s described
a boye.
The Penicillium hispalense description is centered upon the typ e strain IJ FM
5940, isolat ed in ou r laboratory as a cult ure con taminan t in 1978. It is ac tually
kn own only fro m th e typ e strain.
Penicillium hispalense is a di stincti ve species characte rize d m ainly by its
see m ingly polyv erticillate penicilli , owing to th e multiseptate characte r of th e
e1ements of its penicilli , and by its strong ly synnematou s colo n ies .
The correc t pl acement of thi s species is sti ll ope n to questi on since it ap pears
as an intermedi at e between th e Asym m etric- fasc icu la ta and th e Polyverti cillat a
sect ions (Raper a nd Thom 1949). H ow ever , for th e time being , it is tentati vely
assigne d to the for me r sectio n, near P. granulatum Bainier.

172 . Penicillium griseoJulvum Dierckx : fig . 172 , co lo ur p late 17 2 , a n d

SEM pl ate XXIX , (ce n t re: left) .
Peniclllium griseo-fu/vum Dierckx - Ann. Soco Sci. Bruxelles. 25: 88. 1901. See also
Biourge , La Cellule 33: 164-167. 1923; also Thom, 'The Penicillia', p. 371. 1930;
also Raper and Thom, 'A Manual of the Penicillia', pp. 534-539. 1949; also
Samson, Stolk, and Hadlok, in Studies in Mycol. 11: 24-25. 1976.

Synonymies
Penicillium patu/um Bainier - Bull. Soco Mycol. Fr. 22: 208. 1906.
Penicillium urticae Bainier - Bull. Soco Mycol. Fr. 23: 15. 1907.
Penicillium f/exuosum Dale - Ann. Mycol. 24: 137. 1926.
Penicillium ma/tum Hori et Yamamoto - Jpn. J. Bacteriol. 9: 1105. 1954 (not
validly published) .

Co lon ies upon C A (co lour plate 172 , A a nd a) growing restrictedl y ,

a tta ining a di am et er of a bo ut 25 mm in 14 d ays at room temperature
(20-25 OC), radiat ely sulcated by numerous furrows in m ost strains, with
m argin s ab ru p t a nd with cen tral area oft en raised and folde d in a crate rifor m
pattern , ranging from 0.5 to 1.0 mm deep in m arginal areas to 2.0 -3.0 mm
deep in colony ce n tre ; surface di stinctl y gr anular in most recent isolat es, wi th

o
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0 0 000 O

O 0 0 00
o O 8

O O

00 O
0 0
O
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jrrn

10

Fig . 172. Camera lucida

drawings of Penicillium
griseo-fulvum Dierckx, CBS
384.48. A, Detailed drawings
of the penicilli. B, Mature
conidia. C, Habit sketches.

prominen t fasc icles u suall y produced at lea st in m argin al areas; in olde r

isolat es ofte n with ba sal felt ove rla id by a floccose ove rg rowth of wh ite ae ria l
h yphae (colour pla re 172, A) , partly obsc uring th e con id ial areas; heavily
sporing th rou gh out in sorne strains, less heavily in othe rs in arte m isia gree n
shades (Sguy, PI. XXVIII , 418) ; exudate not produce d in sorne strain s,
abu ndantly in othe rs as colou rless large d rops, ofte n largely embe dde d in thc
m ycelial felt; odour distincti ve a nd fra gr ant in sorn e strains, not pronounced in
othe rs; reddish a m ber-coloured pigmen t d iffus ing into th e surrou nd ing agar ;
colony revers e (colour piate 172 , a) at first dulI yelIow becomi ng orange
cinnamon to reddish brown colou re d; peni cilli looscly diver gen t (fig . 172, A),
com paratively large but extre mc ly variable in patt ern and di me nsions, with
conid iophores m on on em at ou s or loosely synne matous, espec ially in m arginal
areas, smo oth wa lled, irregul arl y two- to four-stage branched , with rami
stro ngly divergent ; stipes u su all y undul at e , 400 - 500 p.m by 3.0- 4 .0 p.m ; ram i
mostl y 15- 20 p.m by 3 .0 -3.5 tui , occasion alIy up to 30 p.m in len gth ; m etulae
com para tively sho rt, mostly 7-9 tut: by 3. 0 -3 .5 tun , usu alIy in grou ps of 2 - 4

'IIU

closcly a pp rcsscd elcmc n ts in th c verticil ; phi alidcs co m pa ra tivcly vcry sho rt ,

1 .:- () .5 Ul! by 2 .5 -:~. 0 uu, with a vcry short incon spi cu ou s neck; co n id ia
cllipsoidal, sometimos su bg lo bo se, smooth wall ed , 2.5-3 .5 11m by 2. 3-2. 5 11m
(lig . 172, B, a nd SEM plate 29, ce ntre : left); co n id ia l chains more 0 1' less
c1i vergcnt to tan gled (fig. 172, C) .
C olon ies upon C YA (calour pIat e 172, B a nd b) attaining a di amet er of
:)0 mm at room temper ature (20-25 OC), sim ila r in texture to th ose grown
lIpon CA but less ab ru ptly raised , with a stro ng ly wrinkled basal felt in a radial
pattern , some times ove rlaid b y a floccose overg ro wth; heavil y spo ring
throu gh out in gray-gree n shades (Sguy, PI. XVIII, 419) ; ad our faint ,
ind efinite ; ex udate not produced ; reddish a m be r-colou re d pigment diffu sin g
into th e surrou ndi ng agar; colony reverse (co lour pl at e 172 , b) developing deep
rcddi sh b rown shades ; penicilli esse ntially as described aboye .
Colonies upon MEA (colou r pl at e 172, C and c) att a ining a di ameter of
a bo u t 45 mm in 14 d ays at room temperature (20- 25 OC), co nsis ting of a
loa ser textu re d b as al fclt, mostly pl ane , with a ve ry thin m argin su rro u nde d by
a fringe of submerged hyphae ext ending a bou t 3 mm beyond th e ae rial
growth; heavily sporing through au t in dark gree n shade s (Sguy , PI. XXV II,
401) in sorne stra ins show ing co nid ia l arcas partly ob scured by a deepl y
floccose ove rg ro wth of white vegeta tive ae rial hyphae (colour plate 172 , C);
yellow-colou re d pi gm ent diffusin g into th e su r rou nd ing agar; colo ny reverse
(colour plat e 172, c) showing reddish brown colours; penieilli esse n tially a s
described a boye, with th e co nid iophares arising primarily fro m th e
su bstratum.
The spec ies description is ce n te re d upon strain CBS 124.14 = C M I
92,2 73 = NR R L 992 = VKM F-320 = IJ FM 1863, isolat ed by M iss Dal e in 1926 a s
P. flexuosum an d reg arded b y Thorn (1930) a s representing P. griseo-fuloum, and
many othe r stra ins isolat ed by A.T. M artnez from a n ando sol in th e Sp anish
P rovince of Navarra, 1977 . (Colou r pl at e 251 presents a representative strain
of P. griseo-fuloum isol at ed b y th e a u tho r from packed m ash ed potat oes. )
Penicillium griseoj ulvum Di er ckx ap pears to be widely di stributed but not
particul arl y a b u ndant in nature . It s typi cal habit at is soil, but it has been
isolat ed from d eca yin g vegetation, shee p dung , a nd othe r su bs tra ta.
When inoculated into sou nd citrus fru its, P. griseo-fuloum produces black to
dark b rown necrotic arcas 10 to 15 mm in di amet er in abo u t 10 d ays, and
throughout the centra l area a deep purplish colouration , but without marked
evi de nce of tis su e destruction .
Biochemicall y , th e principal interest of th e sp ecies was cente re d arou nd th e
p roducti on of an an tibiotic rep orted by Anslow et al. (1943) to be produced b y
stra ins of Penicillium patulum Bainier, hen ce design at ed patulin. T he sa me
an tib iotic has been rep orted fro m othe r sp ecies of Penicillium and Aspergillus
under different names by othe r in vestigators (c1a viformin and clavacin) .

4114
H owever, since patulin is app rcciab ly toxic to a nimals (Abraham and Florcy,
1949), it is regarded ac tually as a mycotoxi n .

173 . Penicillium granulatum Ba ini er : fig. 173 , colour plat e 173 , a nd SEM

plat e 29, (centre: righ t).

$V

10 um

00 O

O OO ~OC)()
00 0 0 C)
00 O

'----'

100,JJm

Fig . 173. Camera lucida

drawings of Penicillium
granulatum Bainier, CBS
171 .44. A, Detailed drawings
of the penicilli . B, Mature
conidia. e, Representative
penicilli as seen under low
power. D, Habit sketch
showing the cha racteristic
synnematous arrangement of
the con idiophores.

Penicil/ium granulatum Bainier - Bull . Soc o Mycol. Fr. 21: 126. 1905.

Synonymies
Penicil/ium divergens Bainier et Sartory- Bull. Soco Mycol. Fr. 28: 270. 1912.
Penicil/ium schneggii Boas - Mycol. Z. 5: 73-83. 1914; and Z. Bakt. Parasitenk .
Abt. 2, 44: 695-696. 1916.
Co loni es u pon C A (colour pl at e 173, A a nd a) attaining a di ameter of abou t
40 mm in 14 days at room temperature (20 -25 OC), com pa ratively deep , up to
2-4 mm , m ore 01' less floccu lent wit h abu ndant sterile hyphae white to dull

'111:>

yc llow co lo urcd , with surlacc irrcgularly tuft cd ami radi at cl y wrinklcd , with a
I hin m argin su r ro u nde d by a narrow Iringc 01' su b mer ged h yphae exte nd ing
nhou t 1 mm beyond th e aerial growth; m edium sporing in pale blue-green
shades (Sguy , PI. XXIX , 428) ; ex u date limited to abundantIy produced ,
clca r lo pal e yell ow ; odour a romatic, usu ally pronounced ; colony reverse
usually in dull ye llow to orange 01' reddish brown shades (colour pia te 173, a),
wirh a gr een cas t showing through in marginal areas ; penicilli as ym m etric (fig.
173, A) , comparatively large, with con id iop hores sometimes m ononematous,
but more com m on ly synnematous to loosely synnematous a nd in zones ,
cspecially in m arginal areas (fig . 173, D), consp icu ou sly rough-walled , two- to
ihree-stag e branched; stipes variable in length, from 100 to 200 utt: up to very
long in synnemata, about 3.5- 4 .0 p.m in diameter; rami variable, com m only
12-30 p.m by 3.0 - 3.5 iui, somewhat appressed against the main axis;
metulae m ore 01' less cylindrical, usually rough wall ed , few in the verticil,
about 8-12 p.m by 3.0- 3.5 p.m ; phialides bottIe shape d , with a short but
consp icu ou s neck, in small crowde d clusters , usually about 7 -10 p.m b y
3.0-3 .5 p.m; conidia strongly elliptical when you n g, in age elliptical to
subglobose, 4-5 utt: by 3-4 p.m in di ameter, with walls smooth 01' nearly so
(fig . 173, B, and SEM plate 29, ce n tre: right) ; conidial ch a in s tangled 01'
divergent (fig . 173, C).
Colonies upon CYA (colour plate 173, B and b) a tta in ing a diameter 01'
45 mm in 14 days at room temperature (20-25 OC ), similar in texture as
described ab oye but poorly spor ing in brownish shades, with basal felt ov erlaid
by a dense overgrowth 01' whitish ste r ile aerial hyphae ; with a thin margin
surrounded by a fringe 01' submerged h yphae ext ending a bo u t 2- 3 mm b eyond
th e ae r ial growth ; colony surface strongly radiat ely furrowed a nd irregul arly
fold ed in th e cen tre; od our strong, suggestin g a m m onia ; exu date n ot
produced ; no colou re d pigment .diffusing into th e su rrou nd in g aga r ; co lony
rever se (colour plate 173, b) in reddish orange sh ad es ; penicilli esse n tially as
described aboye .
Colon ies upon MEA (colou r pl at e 173, C a n d e) gro wing variously , in sorne
stra ins a ttain in g a diamet er 01' 40-50 mm in 14 d ays a t room temperature
(20-25 OC ), in others restrict ed a n d not exceed ing 25 mm in th e same period
(colou r plate 173, A) , plane , not furrowed , with su rface consp icuo us ly
gr anular from the production 01' abundant fas cicles up to 1 mm 01' more in
length; heavily sporing throughout, in dull gra y-green shades (Sguy,
PI. XXIX , 426) , with surface partIy ea ve re d by an overgrowth 0 1' white
floccose ae r ia l h yphae , obsc u rin g more 01' less th e fru iting areas; exu date not
produced ; odour arom ati c; penicilli essen tially as d escribed aboye ; n o colou re d
pi gm ent diffusing into th e su rrou nd ing agar; colon y rev erse (co lou r pia te
173, e) in yellowish orange shade s.
The specie s description is centered upon th e n eo typ e strain C BS