CHAPTER II

GALL-BEARING PLANTS AND

GALL-INDUCING ORGANISMS

1. Plants that bear gall

Galls arise on nearly all the major classes of plants. Although no

group of plants is entirely free from galls, even casual observation
shows that galls are more common on some groups than among
others. Relatively few galls are found, for example, on Cryptogams
and Gymnosperms, but the Phanerogams, especially the Dicotyle-
donae, are remarkable for the abundance and great diversity of
their galls (fig. 5).
Algae.
Although the Algae would at first sight appear to be unlikely to
have galls, a number of interesting galls, caused by parasitic Algae,
Bacteria, Fungi, Nematoda, Rotifers and Copepodes, are known on
several species of Algae. Acarina and insects are not, however,
known to give rise to galls on Algae. Some of the better known of
these galls are found on the marine brown and blue-green Algae,

_GALLS ON CRYPTOGAMS &

GYMNOSPERMS
H···.:.:J GALLS ON MONOC01S
c:J GALLS ON DICOTS

Fig. 5. Relative abundance of galls on the major classes of plants.

but some typical galls have also been described on a number of

fresh-water Algae like Batrachospermum, Urospora, Coleochaete,
Vaucheria, etc. 12S7 . SCHMITZ1030 has described nodular galls,

M. S. Mani, Ecology of Plant Galls

© Springer Science+Business Media Dordrecht 1964
 13

caused by bacteria on the red Algae Chondrus crispus, Pelesseria

sanguinea, etc. BRAND l18 has given an account of bacterial galls,
in the form of irregular cellular outgrowths, on the filaments of
Chantransia. Reference may also be made to CHEMIN 186 for an
interesting account of several other bacterial galls on Algae like
Gracilaria confervoides, Cystoclonium, Saccorhiza bulbosa, etc. Other
galls on Gracilaria confervoides are caused by Gloreocolex pachydermus
and Gracilarophila oryzoides. Small galls are reported by LAGER-
HEIM 650 to be produced by Sarcinastrum u,rosporae LAGERH. on
Urospora mirabilis. Several examples of galls produced by parasitic
Algae on other species of Algae are also known. SAUVAGEAU 1010
has, for example, described a gall on Cystoseira ericoides, caused by
the parasitic Alga Ectocarpus valiantei. VALIANTE 1200 reports
Streblonemapsis irritans as causing nodular galls on Cystoseira
opuntioides. Among the fungal galls on Algae, the best known is
perhaps that of Sphacelaria and Cladostephus, produced by Olpi-
dium sphacelarum. ROTHERT 984 , TROTTER 1l64 and BENK0 79
have given interesting accounts of the galls produced by the rotifer
Notommata werneckii EHRENB. on Vaucheria spp. Swellings of the
air-bladders of the marine Algae Ascophyllum nodosum, Desemares-
tia aculeata and Rodymenia palmaza produced by the Nematode
Tylenchus fuscicola, were described by BARTON 58 in 1891 and by
DE MAN 246 in 1892. BARTON 58 has also described the nematode
galls on Furcellaria and Chondrus.
Fungi.
Galls are not really rare on Fungi, as may be supposed, but only
a relatively small number of these have so far been studied in any
detail. KUSTER 639 refers to Pleotrachelus fulgens ZOPF. (Olpidaceae)
as giving rise to irregular outgrowths on the mycelia of the fungus
Pilobolus crystallinus. The so-called Peronium aciculare, formerly
erroneously described as a distinctive species of fungus, is in fact
Saprolegnia, galled by OlPidiopsis saprolegniae. Certain parasitic
fungi are also known to give rise to galls on various Basidiomycetous
fungi. VUILLEMIN 1224 refers to the deformations on Boletus
granulatus caused by Sepedonittm chrysospermum. According to him,
Mycogone rosea gives rise to galls on the fructifications of Clitocybe,
Tricholoma, etc. Although the fleshy fructifications of most higher
Fungi are attacked by a great variety of insects, galls are, however,
formed only exceptionally on these fungal organs. VOGLER 1220 ,
RIEDEL 933 and RUBSAAMEN 992 have reported interesting galls on
the fructifications of different Basidiomycetous fungi. BOUDIER115
and THOM 1l33 have similarly described galls on certain well known
Agaricaceae. According to DANGEARD 24o , a Nematode produces
small galls on Ascobolus furfuraceus.
The large fructifications of Polyporaceae often bear on the lower
14

side numerous conical tubular galls. RUBSAAMEN 992 believes that

this tubular gall is produced by Diptera, which is perhaps Scardia
boleti F.981. ULBRICH 1199 has recently described the galls on Fomes
salicinus and Ganoderma applanatum from Europe.
Lichens.
The galls on lichens comprise the well known cephalodia and are
generally caused by the "foreign" Cyanophyaceae, which penetrate
into the thallus and give rise to horny or warty and often also other
kinds of malformations. BEIJERINCK 73 and FORSSELL 339 were per-
haps among the first to recognize the cephalodia as true galls. It is
also of considerable ecologic importance to remark that lichens are
the only Thallophyta on which Acarina are known to give rise to
galls.
ZOPF 1289 has given accounts of several galls on lichens like Rama-
lina kullensis, Ramalina scopulorum, Ramalina cuspidata, etc. Ac-
cording to BACHMANN 44 , 45, 47, a gall on Ramalina jraxinea is caused
by the caterpillar of a moth. He found the thallus developing into
a common gall chamber in Ramalina kullensis, but there are several
globose chambers in the gall of Ramalina jraxinea. The gall on
Ramalina kullensis is caused by Acarina. A number of different galls
develop on the squamae and podetia of Cladonia ochrochlora. BACH-
MANN 40 -43 , 45-48 has also given accounts of the structure and devel-
opment of several other interesting mycocecidia on the lichens
Cladonia ochrochlora, Cladonia jimbriata forma simplex minor, Cla-
donia cornutadiata, Cladonia jimbriata forma tubaejormis, Cladonia
degerens phyllophora, Cladonia gracilis and Cladonia pityrea holo-
lepis. The mycocecidia on Cladonia are remarkable for the enormous-
ly swollen cells of the alga of these plants. The walls of the fungal
cells of these lichens are also greatly thickened, with enlargement
of the cell spaces. The gall on Centraria glauca, caused by the fungus
A brothallus centrariae, has the thallus irregularly bulged above,
with a corresponding invagination below. The fungal galls on Par-
melia physodes are globose tubercles on the upper surface. In species
of Ramalina, the gall formation usually results in the branches of
the stock being tubularly swollen the whole of their length or only
basally.
Bryophyta.
A number of galls, like the swellings caused by N ostoc lichenoides
on Blasia pusilla, are known on liverworts. MARcHAL 748 has de-
scribed nematode galls on the growing tips of the mosses Lophocolea
bidentata and CePhalozia connivens laxa.
The leafy mosses bear several zoocecidia, caused particularly by
Tylenchus Spp.273, 762, 1018, 1019. Most of these galls are typically
artichoke-like swellings of the growing tips. GOODEy413 has given an
 15

interesting account of another nematode A nguillulina, causing galls

on the moss Pottia. Although different galls have been described on
over thirty genera of leafy mosses from Europe alone, only those on
Dicranum longifolium EHRENB., Cephalozia and Lophozia have been
studied in detail (fig. 49D).
Pteridophyta.
A number of galls on ferns, caused by fungi, Acarina, Hymen-
optera, Diptera, etc., have been described from different parts of
the world. The gall of Eriophyes pteridis NALEPA on the fronds of
Pteridium aquilinum KUHN (fig. 102D) is one of the best known of
these. Some interesting Diptera galls on the stem and leaves of
Hymenophyllum spp. are known from South America 535. These galls
are often remarkable for the dense covering of multicellular and
branched hairs 975. RUBSAAMEN 997 has described an interesting leaf
margin roll gall on the fronds of N ephrolepis exaltata SCHOTT,
caused by Eriophyes from Malaya (fig. 102C). Bulbil-like galls on
Selanginella pentagona are caused by Dipteral l l l . The gall-midge
Perrisia jilicina KIEFF. gives rise to leaf margin roll galls, with a
curious resemblance to an abnormal indusium. GIESENHAGEN 389 , 390
has also reported the finding of mycocecidia on ferns.
Gymnospermae.
As mentioned above, galls are more frequently met with on
Gymnosperms than on Cryptogams. The best known of these is the
an an as gall, caused by the aphids Chermes (Adelges) abietis LINN.
and Cnaphalodes (Adelges) strobilobius KALT. on Picea excelsa
(LAMK.) LINK. (fig. 52). Globose or fusiform galls on the stem of
Ephedra spp., caused by Coleoptera, Lepidoptera and Diptera
(fig. 47 D), are known from different parts of the world. Leaf galls

70
'"«u
«C)
60
...
«
'"~
50
iii
40 0Q.
~
30 0
u
20
10

Fig. 6. Percentage frequency of galls on the dominant Natural Orders from the
Holarctic.
16

on Gnetum are known from India and Java. Large galls on Juniperus
spp. are caused by fungi of Gymnosporangiae. Bacterial galls occur
on Pinus spp. A remarkable witches-broom gall on Abies alba is
caused by the fungus M elampsorella caryophyllacearum. Different
galls on J~tniPerus spp. are caused by Eriophyes quadrisetus j~tni­
perinus NAL., Schmidtiella gemmarum RUBS., Oligotrophus spp., etc.
A ngiospermae.
The Angiosperms bear not only the greatest bulk of all the known
galls from the world, but they are also remarkable for the enormous
diversity of their galls and the gall producing organisms associated
with them. In 1911, KUSTER 639 listed over 5500 galls on Angio-
sperms, mainly from Europe and North America. More recently
Ross & HE DICKE 982 reported 2900 galls on these plants from Central
and North Europe alone. HOUARD 520 has listed nearly 3200 galls on
the Angiosperms from Asia, Africa, Australia and Oceania and
about 1300 galls from South America 535 . FELT 330 f~stimated that
nearly 2000 galls are known on these plants from North America,
north of Mexico. DocTERs VAN LEEUWEN 307 described about 1500
galls on Angiosp~rms from Java, Sumatra and other parts of the
East Indies. The total number of galls on Angiosperms so far known
from all over the world is certainly not less than 14,750, representing
almost 98% of all the known plant galls.
The greatest bulk, viz. over 93% of the galls on Angiosperms,
are found on the Dicotyledonae. Although all Natural Orders of
Dicotyledons bear galls, it is usual to find them more frequently
on Cruciferae, Capparidaceae, Malvaceae, Sapindaceae, Anacar-
diaceae, Leguminosae, Rhamnaceae, Rosaceae, Geraniaceae, Vita-
ceae, Tiliaceae, Tamariscaceae, Myrtaceae, Combretaceae, Rubia-
ceae, Umbelliferae, Compositae, Convolvulaceae, Labiatae, Acan-
tha<:eae, Euphorbiaceae, Fagaceae, Salicaceae, Ulmaceae, Urtica-
ceae, Moraceae, etc. than on the others. The frequency of gall on
plants of different Natural Orders is naturally very different in
different parts of the world. Galls on Gymnosperms, Salicaceae,
Aceraceae, etc. are, for example, exceedingly rare in tropical regions.
There is a very pronounced abundance of galls on Compositae and
Fagaceae in North America and Europe. In these regions nearly
50% of the galls are found, for example, on Fagaceae, about 20%
on Compositae and 15% on Rosaceae (Fig. 6). Galls on Leguminosae
are particularly dominant in South America, parts of Africa and in
India. More than half the known galls from Australia are found on
Myrtaceae. Among the Monocotyledonae, the majority of galls
occur on Gramineae and Cyperaceae. Some galls have also been
described on Potamogetonaceae, Araceae, Liliaceae, Juncaceae,
Orchidaceae, Amaryllidaceae and Musaceae. For further accounts
of the frequency of galls on plant genera reference may be made to
 17

HEIMHOFFEN 54 and for their frequency in different Natural Orders

to KARNy 565 (vide Chapter IX).

2. Parts of plants galled

Galls are found on all the parts of plants, from the root tip to the
growing point of shoot, both on the vegetative and on the reproduc-
tive organs. The aerial parts of plants, as may be expected, bear
much larger numbers and greater varieties of galls than the under-
ground parts. Galls on roots and other subterranean parts are gener-
ally less evident than those on aerial parts and are therefore often
overlooked. In addition to the ordinary parts like roots, hypocotyl,
stems, buds, petioles, leaf blade, flowers, fruits, etc., galls are often
known to arise on other galls also. Not only the individual parts,
but not infrequently the whole of the aerial portion of a plant may
become galled in toto, giving rise to the familiar witches-broom or
hexenbesen galls. This is for example the case with Vaccinium vitis
idaea LINN., attacked by the fungus Exobasidium vaccini (FucK.).
Other common examples of galling of the whole of the aerial portion
of a plant are frequently found on Salix, Betula, Prunus, Syringa,
Pinus, Ononis, Euphorbia cyparissias LINN. and Euphorbia gerar-
diana. Some of these galls are caused by fungi, others by nematodes,
Acarina and still others by insects. Not all witches-broom growths
are the result of parasitic attack1192 .
Galls developing on other galls have been termed mixed galls and
are often of great complexity. HEDICKE 450 distinguishes anacecidia
from epicecidia, depending upon the degree of their complexity,
particularly on the anatomical and histological modifications brought
about by the second gall on the tissues of the first gall, on which
it forms. One of the commonest anacecidia is caused by the aphid
Byrsocrypta gallarum (GMELIN) on the galls of another species of
aphid on Ulmus. The gall of Diplolepis (= Rhodites) eglanteriae
(HTG.) on the gall of another cynipid Diplolepis (= Rhodites) rosae
(LINN.) and the gall of I teomyia capreae WINN. on the gall of Pontania
capreae LINN. on Salix spp. are common examples of epicecidia.
DOCTERS VAN LEEUWEN 308 has given an account of an interesting
gall caused by the thrips Gynaikothrips devriesii KARNY on the gall
of an unidentified midge on the leaves of Elastostema sesquejolium
HASK. from the East Indies. The gall of A ndricus inflator H TG. has
been reported to develop on the bud gall of Andricus globuli HTG.
on Quercus; not infrequently the gall of Andricus jecundator HTG.
arises on the gall of Andricus (collaris) curvator HTG. 639 •
Galls are not equally abundant on different parts. About 5% of
the known galls produced by cynipids on Quercus from the world
arise, for example on roots, 5% on buds, 22% on branches, 2% on
flowers, 4% on acorns and about 63°/r) on leaves (fig. 7). Over 80%
18

of the galls on Rosaceae are leaf galls. Nearly 70% of the South
American galls are on leaves of diverse plants, 20% on branches
and twigs, 7% on buds and only about 1-2% on flowers, fruits and
roots. About 60% of the galls from the Oriental Realm are also leaf
galls, about 15% are branch galls, 12% bud galls and the flower and
fruit galls amount each to about 6%. Almost 55% of all the galls
so far known from India arise on leaves; the stem galls represent
about 25% and flower galls 10%. From the world, the cynipid galls
on leaves amount to 80%, midge galls represent about 50%, Erio-
phyid galls 80%, Homoptera galls 90%. Considering the galls caused
by all agencies from the whole world, the largest number of galls
are leaf galls, viz. 65%. Next in importance of abundance are the
galls on twigs, branches and parts of the main axis, amounting to
about 19%, with the bud galls standing at about 10% and the
flower galls are about 4% and the root and fruit galls are each
only 1% (fig. 8). The relative abundance of galls on different parts
depends primarily on the species of the plant and the gall-maker
and is also influenced by a variety of other environmental factors.
It may also be observed that not all leaves, branches or buds of any
given plant become galled, but the physiological conditions and the
age of the organ seem to more or less greatly influence the develop-
ment of galls. Furthermore, different gall-makers seem also to have
become specialized to different degrees in their ability to induce
galls on different parts. Certain types of galls are, for example,
strictly found only on particular organs or even a part of an organ,
such as leaf margin, the upper or the lower surface of the leaf blade,
the petiole, etc. The development of a gall on a part presupposes the
capacity of the part to react in a specific way to the influence of the
gall-maker. As may be expected, all parts do not appear to be equal-
ly responsive in this respect and their capacity also seems to vary
with age and stage of development of the part. HOFMEISTER 479 was
of the view that not only the normally actively growing, young and

lJ..
<-
70 III
..J
60

50

40
~
III
30 I-
\II
20 l- I-
0 0 5
~
:::> a:
10 III lJ..

Fig. 7. Percentage frequency of galls caused by cynipids on different parts of

Quercus from the world.
 19

physiologically plastic structures, but also others that have ceased

to grow and are more or less dormant and physiologically relatively
inactive are capable of forming galls. THOMAS 1l35 believed on the

...<
LaJ
70 J

60
GSO
z
~40
0
::E
...~30 LaJ
I-
III
20
>-- I-
10 5 0
Q:
u. 0
cr

Fig. 8. Percentage frequency of galls caused by all agencies on different parts of

higher plants from the world.

other hand that gall formation is only possible as long as the organ
is still in a physiologically active state of development and growth.
KDsTER 639 has, however, cited numerous examples of galls by the
aphid Adelges jagi HTG. on old bark layers of beech. ZWEIGELT1297
recently reviewed the evidence on this important problem of the
age of the gall-bearing organ and of the cecidozoan as factors in gall
initiation and development. According to him, with increasing age
of the plant part and thus with differentiation of normal tissues, the
general complexity of the resulting gall decreases. In other words,
the degree of organization of the gall is inversely proportional to
the age of the gall-bearing organ. Old parts of plants do not show,
however, any demonstrable loss of capacity to react to the gall-
maker, but the latter only appear to be better adapted to the youn-
ger parts of plants than to the older ones.

3. Organisms which induce galls

The capacity to induce galls on plants is met with in widely
unrelated groups of animals and plants, besides border organisms
like bacteria and virus. Animals which give rise to gall are termed
cecidozoa and the vegetable organisms that give rise to galls on
other plants are cecidophyta. Not all species of a group cause the
formation of galls and gall forming species are often found among
genera and families, which are predominantly and typically non-gall
making types and in some cases are not even phytophagous in their
general habits.
20
i. Cecidozoa

The principal groups of cecidozoa are Trochelminthes, Nemathel-

minthes, Acarina and Insecta. With very few exceptions, the deve-
lopment of galls is closely bound up with the feeding activities of the
cecidozoa. The mode of feeding differs in different classes of ceci-
dozoa and these differences are correlated with fundamental differ-
ences in the structure of their mouth-parts. Some cecidozoa, like
the larvae of Coleoptera, Lepidoptera and some Hymenoptera, have
typic all y biting and chewing mou th-parts and others like Homoptera,
Heteroptera, Nematoda and Acarina have piercing and sucking
mouth-parts. Still others like Thysanoptera have rasping and suck-
ing mouth-parts. The larvae of the gall-midges (Itonididae: Diptera)
have very undeveloped mouth-parts and largely seem to absorb their
nourishment by suction. The salivary secretions of cecidozoa play
an important role not only in the feeding of the cecidozoa but also,
as we shall see later, in the initiation and development of the gall
itself (vide Chapter XIV). It is known, for example, that the salivary
glands of the gall-forming generation of the aphid Chermes are better
developed than in the non-gall-forming generation. Cecidozoa are
not capable of initiating the formation of galls throughout their life;
different groups of cecidozoa are able to influence the plant in such
a way as to produce galls at different stages in their life-history.
Most of them are able to do this only in their immature stages, but
some others are capable of initiating the development of a gall both
as larvae and as adults. It is also interesting to note that in some
species of cecidozoa, it is only one generation that is capable of
forming galls, while the succeeding generation in the life-cycle does
not produce any gall. Some of the outstanding characters of galls
caused by animals, viz. zoocecidia, are summarized in chapter IX.
T rochelminthes.
The only rotifer so far known to give rise to galls is N otommata
werneckii EHRENB., associated with unicellular galls on Vaucheria
spp., to which we have already referred. These galls are irregular,
unseptate, lateral outgrowths on the tubular green Algae 984 , often
expanded and with 1-4 tubercles apically.
Nemathelminthes.
The gall Nematodes belong to the order Rhabditoidea, often also
called Anguilluloidea. They comprise a large and important assem-
blage of mostly moderately sized worms, with the cephalic sense
organs reduced to papillae and the amphids reduced to pockets. The
pharynx generally presents one to two bulbs, one of which is a
pseudobulb and the other is a valvulated bulb. The excretory sys-
tem is usually asymmetrical. The rhabditoid Nematodes are terres-
 21

trial, sarcophagous or coprophagous and by transition to partial or

complete parasitism in plants and animals. The strictly gall-forming
species belong to the family Tylenchidae. This family is recognized
by its conspicuous buccal stylet, assymetrical form of the excretory
system, with canals on one side. Exhaustive accounts of Tylenchids
have been given by GREEF 416 , GOODEy404 and FILIPJEW & SCHUUR-
MANS-STEKHOVEN 331 . The species pierce the plant tissues with the
help of their buccal stylet and suck the plant sap. Their pharynx is
characterized by a muscular median bulb and a posterior glandular
dilatation. In the Aphelenchoid type the pharyngeal glandular
dilatation projects backward as a lobe over the anterior portion of
the intestine. All Tylenchoids are not, however, gall-formers. The
gall species, often also called eelworms, belong to the genera Tylen-
chus, Aphelenchus and Heterodera. Though parasitic in plants, most
of these gall nematode worms are capable of leading a free life in
soil for variable periods. The species usually give rise to galls on
roots, but often also on stems, leaves, flowers and fruits. They give
rise to galls predominantly on Dicotyledons, but sometimes also on
Monocotyledons, especially the grasses and occasionally on mosses
and marine algae. Most gall Nematodes are remarkably resistant to
desiccation and remain viable in the dry state for several years. The
dried Nematode galls on Triticum vulgare yield, for example, on
soaking, thousands of the first stage juveniles of Anguina tritici. The
dry cysts of H eterodera remain viable in soil for at least a whole year.
STEINER et alY04 have described a remarkable case of Tylenchus
polyhYpn1ts remaining alive for almost thirty years on herbarium
plants. Due to their capacity for active locomotion and also partly
due to their high fecundity, gall Nematodes are generally distributed
extremely rapidly. The genus Heterodera exhibits a high degree of
parasitic adaptations, variations and sexual dimorphism (fig. 9). The
second stage juvenile worm pierces a new host plant and sucks the
sap from the cells. This leads to the formation of a gall at the site
of attack. At the end of the third moult, the female becomes a
plump, pear-shaped or gourd-shaped bag, containing 300-500 eggs.
It then degenerates and finally leaves its cuticle as a protective
cyst for the eggs. These cysts become free in the soil by the disinte-
gration of the plant galls. The second stage juveniles develop within
these cysts and escape into the soil and are able to live for several
months without feeding, if a suitable host is not available. The males
pass through an additional moult, but do not become pear-shaped
bags. They escape from the decaying gall, seek a female and fertilize
it. The detailed structure, habits and life-histories of several species
have been investigated by a number of workers 199 , 406, 478, 1101,
1113, 1154. Most species are known to reproduce parthenogenetically
1196. The species very often exist in several distinct biological races,
structurally nearly indistinguishable from one another, but each
22

race attacks one set of host plants and does not breed on plants of
the other groups 1101, 478, 406, 198. Some species, like the common
Heterodera marioni (CORNU) GOODEY, give rise to galls on the roots

Fig. 9. Gall Nematodes. A-C. The second. third and fourth stage larvae of
Heterodera marioni (Cornu) Goodey. D. Adult female of Heterodera marioni (Cornu)
Goodey. E. Adult female of Heterodera schachti Schmidt.

of nearly 850 different species of plants. The family Allantonemati-

dae, recognized by their degenerate pharyngeal musculature, is
generally parasitic in diverse insects, but has a free-living phase
in plants attacked by the insect. The family is remarkable for con-
taining one species which gives rise to galls on Eucalyptus in Austra-
lia.
Acarina.
The gall mites belong to two families, viz. Tarsonemidae and
Eriophyidae. The Tarsonemid mites are generally about 0.25 mm
long and have four pairs of legs and a rounded body. The commonest
specie~ Tarsonemus phragmitidis SCHLECHT. is associated with
peculiar galls on Phragmites communis TRIN. Tarsonemus spirifex
MARCH. is associated with galls on grasses. Another related mite
Tenuipalpus geisenheyneri RUBS. (fig. 11 A) gives rise to small galls
in the angles of leaf veins 996 .
The Eriophyidae, an important group of cecidozoa, (fig. 10), are
minute, elongate, cylindrical mites, with only two pairs of five-
segmented legs and an annulated abdomen, ending in retractile
caudal lobes. The genital organs, epiandrium and epigynium, are
unpaired structures. Respiratory and circulatory organs and eyes
are absent. The mite is usually 0.08 to 0.28 mm long. Long setae,
found on different parts of the body, are of taxonomic importance.
 23
Most eriophyid mites are white, yellowish-white, yellowish-red or
brown. Relatively small numbers of eggs, each about 0.05 mm long,
are deposited by the female. The eggs hatch in a short time and the

Fig. 10. Eriophyes, a typical gall mite. Ventral view of the adult.

newly hatched larvae resemble the adult mites very much, but they
are much smaller, more weakly bristled and also lack the external
genital apparatus. On moulting, the larvae transform into nymphs,
which are larger and also have the external genital apparatus. The
adult female mite is as a rule larger than the male adult. The mouth-
parts of the eriophyid mites are adapted for boring into the plant
tissues and sucking their contents out in liquid form. The mandibles
are uniarticulate and needle-shaped, but them axillary palpi are
free and have three segments. Owing to their small size, most erio-
phyid mites can pierce single cells and suck its contents. NEMEC 861
has given an interesting description of the mode of feeding by these
mites. Eriophyids are remarkable for their relatively large salivary
glands. The salivary secretion is poured into the plant cell through
the wound at the time of sucking. Females are far more abundant
than males and reproduce commonly by parthenogenesis. The
greatest majority of gall mites hibernate on the plant, often below
the protective scales of buds or in the angles of branches and buds
and some species over-winter inside buds.
24

Not all species of eriophyids give rise to galls and some species
live free on leaves, others occur in some abnormal growth on plants
and still others are inquilines and commensals inside galls of other
species. The structure, habits and classification of eriophyids have
been placed on a sound basis by the monumental labours of NALE-
PA 837, 839, 842. Over 500 species of gall mites have so far been des-
cribed, the greatest bulk of which belong to the genus Eriophyes
(SIEB.) NAL. Other important gall mites are Phyllocoptes NAL.,
Anthocoptes NAL., Oxypleurites NAL. and Epitremerus NAL.

Insects
The princi;xll gall insects belong to Thysanoptera, Heteroptera,
Homoptera, Coleoptera, Hymenoptera, Lepidoptera and Diptera,
but isolated examples of galls, caused by species of some other orders
have also been described from time to time. Lestes viridis (Odonata),
for example, is reported to cause galls by thrusting its eggs under
the epidermis of various plants. The plant tissues immediately
surrounding the egg become more or less conspicuously swollen,
but as soon as the larvae hatch, undergo decay. Such galls, initiated
by oviposition and not by the feeding activity of the cecidozoa,
have been described as procecidia by THOMAS 1142 and PIERE 907. All
other insect galls are initiated when the larvae commence feeding.
Some species of Thysanoptera, Heteroptera and Homoptera are
capable of inducing the development of galls both in their larval and
adult stages.
Thysanoptera.
Thysanoptera are minute insects, ranging from 0.5 to 10 mm in
length, with a slender and depressed body and rasping and sucking
mouth-parts. They are usually found on foliage, buds and flowers.
They are readily recognized by their narrow, long-fringed membra-
nous wings and by the swollen, bladder-like arolia between claws.
Though Thysanoptera are known to give rise to galls on plants in
nearly every floral region of the world, they occupy on the whole a
minor place among the cecidozoa. The suborder Tubulifera con-
tains many gall thrips. K ARNy 561-564 has given excellent accounts
of the thrips commonly found in galls. HOUARD 520 records over
fifty species of gall thrips, belonging to about twenty-four genera,
chiefly from the Indo-Malayan region and Africa, representing
about 10% of the known species of cecidozoa from the area. In
South America the gall thrips represent hardly 0.5% of the insects
that induce galls. According to DOCTERS VAN LEEUWEN, over one
hundred and twenty-five species of gall thrips are found in Java.
The more important genera of gall thrips are Cecidothrips KIEFFER,
Cryptothrips UZEL, Dolerothrips BAGN., Gynaikothrips ZIMMERMANN,
 25
Mesothrips ZIMMERMANN, Thrips LINN., Euthrips TARG.-Toz.,
N eoheegeria SCHMUTZ, etc.
H eteroptera.
A small number of Heteroptera, belonging to the family Tingidae,
are remarkable for their characteristic galls on flowers of diverse
plants. The species belong mostly to the genera Copium THUNB.
(fig. 11 B), Paracopium, etc. LEACH & SMEE 654 have described the
capsid bug H elopeltis bergrothi as giving rise to gnarled galls on
branches of Thea sinensis.

Fig. 11. Some typical cecidozoa. A . Tenuipalpus geisenheyneri Rubs. B. Copium

clavi corne (Linn.). C. Ceutorrhynchus pleurostigma (Marsh.).
Homoptera
The order Homoptera are important gall insects nearly all over
the world. Gall species are found in several families like Cercopidae,
Jassidae, Chermidae (= Psyllidae), Aphidae, Phylloxeridae, Aleuro-
didae and Coccoidea. FRIEDERICHS 346 reports that the nymph of
the Cercopid Ptyelus (Aphrophora) spumarius (LINN.) causes galls
on certain plants. Procecidia are also produced by some J assids like
Tettigoniella J AC.
Chermidae.
Psyllids or jumping plant-lice are of considerable importance as
gall insects in nearly all parts of the world. The adult psyllids have
two pairs of wings, which are membranous and folded roof-like over
the dorsum. The nymphs have a characteristically depressed body,
usually coated with a waxy secretion and conspicuous wing-pads.
They are all usually sluggish. Not all psyllids are gall formers.
Gall forming species belong to Psylla GEOF., Psyllopsis Low,
Trioza Low (Fig. 12 C), Pauropsylla RUBS., Megatrioza, Cecidotrioza,
Neotrioza, Rhinocola CRAWF., Cecidopsylla, Dinopsylla, Livia, Ozo-
26

trioza, Phacosema, etc. The number of species of gall-forming psyllids,

so far described from the world, exceeds 350, of which almost half
are from the Oriental Region and Africa and one-fifth from South
America.

Fig. 12. Some typical cecidozoa. Homoptera. A. Pemphigus bursarius (Linn.),

alate fundatrix. B. Byrsocrypta gallarum (Gmelin), alate sexupara. C. Trioza
alacris Flor.

Aphidae.
Aphids are minute, winged or apterous, soft-bodied insects, with
3 ocelli in the alate form; antennae 3-6 segmented; fifth abdominal
segment bearing a conspicuous tubular siphon. A very large number
of aphids are known to give rise to galls, often of considerable com-
plexity of structure and development. In some regions, especially in
the temperate lands, the aphids appear to represent perhaps one of
the most important gall insects. The gall aphids reproduce parthe-
nogenetically and bisexually in alternate series of generations, in
spring and summer respectively. The alternation of parthenogenetic
and bisexual generations are often associated with changes in food
plants and pronounced polymorphism. These cyclic changes are also
associated with remarkable differences in the galls caused by the
individuals of different generations. The female and male adults
appear in autumn and after mating, the fertilized female lays. a
single egg. From this over-wintered egg an apterous female, called
the fundatrix or stem mother, hatches in early spring. The fundatrix
reproduces parthenogenetically and gives rise to several generations
of apterous females, which likewise reproduce parthenogenetically
and are thus virgins. Later in the season, alate females also appear
and towards the next autumn once again males also appear. The galls
produced by the same species of aphid on different parts of the same
plant are frequently very different. The same species may also give
rise to galls on different host plants in the course of its cyclic hetero-
genesis. Some of the more common gall aphids belong to Adelges,
 27

APhis, Byrsocrypta HALIDAY (= Tetraneura HTG.) (fig. 12 B), Forda

HEYD., Chermes (LINN.) BORN., Eriosoma LEACH, Hamamelistes,
Pemphigus HTG. (Fig. 12 A), Dreyjusia BORN., Cnaphalodes MAcQ.,
Pineus SCHIM., Peritymbia WESTW. (= Viteus SHIM.), Phylloxera,
etc. About 700 species of gall aphids are known so farfromtheworld.
Almost 50% of the species occur in the Holarctic and the gall
aphids occupy a minor place in tropical countries. Important contri-
butions to our knowledge of gall aphids may be found in MORD-
WILK0 817 , 818, INOUYE 539 , ZWEIGELT1292.1300 and others.
Coccoidea.
The female coccid is a degenerate sessile insect, which remains
permanently fixed to a suitable place on the food plant and sucks
copious quantities of sap. The males undergo a complete metamor-
phosis and have only a single pair of wings and non-functional mouth-
parts. Although several thousand species of coccids are known to
feed on various wild and cultivated plants all over the world,
exceedingly few species seem to give rise to galls. Coccids thus
occupy a minor place among gall insects in most faunal areas, but
in Australia they appear to have reached their maximum develop-
ment. Coccid galls are particularly abundant on Australian Eucalyp-
tus, Casuarina and Acacia. Amorphococcus, Aspidiotus BoucHE,
Asterolecanittm TARG., Chionaspis SIGN., Fiorina TARG. are some
of the more important gall forming coccids. The Australian Peg-
tococcids of the family Apiomorphidae (= Brachyscelidae) like
Apiomorpha RUBS., Ascelis SCHR., Cystococcus FULL., Opisthoscelis
SCHR. and Cylindrococcus MASK. are important gall makers; some
of them are found in America also. Important contributions on gall
coccids may be found in SCHRADER1034, FROGGATT 348 , 352, EHRHORN
317, etc.
Aleurodidae.
Aleurodids, popularly known as whiteflies, have two pairs of
similar wings, coated with waxy bloom. The early larvae are free,
but soon become sessile. There is a resting stage, within the larval
skin before the emergence of the adult. One or two species give rise
to pit galls on leaves. Aleuromarginatus tephrosiae CORB., from India,
gives rise to epiphyllous, hemispherical pouch galls on leaflets of
Tephrosia purpurea PERS. An unidentified species of Bemisia produces
a remarkably fleshy, cup-like, beautifully pink coloured emergence
gall from the epidermis on leaves of Achyranthes aspera LINN. from
India 745 .
Coleoptera.
Coleoptera are hard-bodied insects, with the fore wings modified
into horny or shell-like elytra, which serve to protect the hind wings.
The mouth-parts are biting and chewing. Development is holome-
28

tabolic. The larvae are variable and always have a distinct head. The
phytophagous beetles feed on leaves, or they tunnel inside various
plant parts and also mine inside leaves. Most gall-forming beetles
are Curculionidaeof the genera Anthonomus GERM., Apion HERBST.,
Baris GERM., Ceutorrhynchus GERM., (Fig. 11 q, Cleonus SCHONH.,
Dorytomus GERM., Lixus FAB., Gymnetron SCHONH., Nanophyes
SCHONH., etc. Some species of Cerambycidae, Chrysomelidae,
Buprestidae and Scolytidae have also been reported to give rise to
galls. Altogether about 150 species of gall-forming Coleoptera are
known from the world. Reference may be made to FROGGATT 349 for
gall Buprestidae and to SCHMIDT1026 for gall Curculionidae.

Hymenoptera
Hymenoptera have two pairs of membranous wings, with the
fore wings much larger than the hind wings. Venation in both the
wings is greatly reduced and often also wanting. Gall-forming species
are met with among Tenthredinoidea, Cynipoidea and Chalcidoidea,
which have all biting and chewing mouth-parts.
Tenthredinoidea or sawflies are remarkable for their saw-like
ovipositor. Their larvae superficially resemble the caterpillars of
Lepidoptera, but are distinguishable by the presence of 6-8 pairs
of prolegs. Not all Tenthredinids, however, give rise to galls. The
gall species belong to one of the following genera: Pont ani a COSTA
(fig. 13 A), Cryptocampus, Blennocampa HTG., Hoplocampa HTG.,

Fig. 13. Some typical cecidozoa. A. Pontania viminalis Linn. B. Isthmosoma.

Micronematus, Monophadnus HTG. and Selandria. The larvae occur

in their galls, but pupation is often in soil. Most sawflies give rise to
remarkably large, fleshy, hollow galls on leaves of various species
of Salix and some species also give rise to procecidia on Clematis,
Helleborus, Pteridium, f'iC. 319 , 272.
Cynipoidea.
Cynipoidea are well known as gall insects in Europe and North
America, where several hundred species give rise to often remarkably
 29
complex galls, especially on Quercus and Rosa. Cynipids are mostly
small or minute, hard-bodied insects, with compressed abdomen,
prothorax at the sides reaching back to tegulae, venation reduced
and antennae simple (not geniculate). Their larvae are about 3-4 mm
long, white, apparently apodous, but have distinct head and biting
and chewing mouth-parts. Throughout its development, the larva,
however, does not throw off any excrement, but at the time of pu-
pation the fecal matter may be ejected. Pupation is always in the
gall. Cynipids have well developed ovipositors, with the help of
which the eggs are accurately positioned inside buds, leaves and
other tissues. Many species reproduce parthenogenetically and have
also alternations of parthenogenetic and bisexual generations. Some
species of A ndricus H TG. reproduce exclusively parthenogenetically.
Very often the individuals of the two alternating generations are so
markedly different that they have in the past been described as be-
longing to different species and even genera. In some species there
is a change of the host plant associated with the alternation of
generations. The galls induced by the unisexual and by the bisexual
generations are usually fundamentally different. Some species do
not give rise to galls, but occur as inquilines in the galls of other
species and still other species are true parasites of various insects,
especially Diptera. Important contributions to our knowledge of
the structure, development, habits, ecology and classification of
Cynipids are by MAYR764-766, RIEDEL 932 , ASHMEAD 32 , DALLA TORRE
& KIEFFER237-238, KIEFFER 584, ADLERl, 2, BEIJERINCK 75 , FRUH-
AUF 354 , RAINER 923 , KINSEy594-597, WELD1239-1242 and others.

Fig. 14. Some common cecidozoa. A. Biorrhiza pallida (01.), male of the bisexual
generation. B. The same, female of the unisexual generation. C. The same,
female of the bisexual generation.

Chalcidoidea.
Chalcids are minute or small insects, with venation reduced to a
single vein in the anterior part of the wing, pronotum at the sides
not reaching back to tegulae, antennae elbowed at scape, ovipositor
conspicuous and issuing some distance in front of the tip of abdomen.
30

Although predominantly entomophagousparasites, many gall-

making species are met with among the families Agaontidae, Tory-
midae, Eurytomidae, Perilampidae, Pteromalidae, Encyrtidae and
Tetrastichidae. The more common gall chalcids include Blastophaga
GRAVENH., Isthmosoma (Fig. 13 B), Pareunotus GIR., Coelocyba
ASHM., Terobiella ASHM., Cecidoxenos ASHM., Eufroggatiana GAHAN,
Espinosa GAHAN, Perilampella GIR., Pembertonia GAHAN, Lisseury-
toma CAMERON, Brachyscelidiphaga ASHM., Systolomorpha ASHM.,
Trichilogaster MA YR, etc. GAHAN 356 , 357 has listed the phytophagous
chalcids from the world, some of which are gall chalcids. GAHAN &
FERRIERE 357 have given an account of some of the gall chalcids.
CROSBy 233 described some interesting gall chalcids from Africa. The
Agaontidae are the well known fig-insects, which breed in the figs of
various species of Ficus, both wild and cultivated and turn the
flowers into characteristic galls. Extensive studies of Agaontidae
from the world have been made by GRANDI 415 • For the taxonomy
of Chalcidoidea reference may be made to ASHMEAD 31 and SCHMIE-
DEKNECHT1029 .
Lepidoptera.
Lepidoptera are holometabolic insects, with two pairs of wings
and siphoning mouth-parts, with the body and wings clothed with
overlapping scales. Their larvae, the common caterpillars, have,
however, biting and chewing mouth-parts and 8 pairs of legs, of
which only' the first three are true legs and the rest are prolegs on
segments 6, 7, 8, 9 and 12.
About 100 species of gall-making Lepidoptera are known from
the world. WILLE 1267 has given an interesting account of the Teneid
Cecidoses eremita CURTIS, which forms a singularly interesting dehis-
cing gall on Schinus dependens. Certain species of Ectoedemia (Nepti-
culoidea: Nepticulidae a family of leaf-miners) are gall-makers in
America. The South American Ridiaschina (Tineoidea) has vestigial
mouth-parts and its larvae also give rise to galls. Other important
gall-forming Lepidoptera are A cella, Epiblema HUBN., Gelechia Z.,
Laspeyresia GERM., Lobesia, Nepticula Z., Orneodes LATR., Phalonia
HUBN., Trochilium SCOP., Cecidolechia, Grapholitha, Amblypalpis,
Gnorimoschema, etc. For further account of galligenous Lepidoptera
reference may be made to SORHAGEN 1073 , SOLOWIOW1072 and
SCHMIDT1027 .
Diptera.
Diptera are the most important gall insects nearly all over the
world. They are holometabolous insects, with only the fore wings;
the hind wings are reduced to halteres. The most important gall-
making Diptera are Itonididae (= Cecidomyiidae), Agromyzidae,
Trypetidae and Anthomyiidae.
 31

Itonididae or gall-midges (fig. 15) represent a large family of

Nematocerous Diptera, with about 5000 species described so far
from the world. They are readily recognized by their holoptic eyes,
short coxae, absence of tibial spurs, reduced wing venation and by
their characteristic antennae, with special sensoria called circumfila.
Though some of the Itonididae are fungus feeders, debris feeders or
are predaceous on various insects and mites, the bulk of the species
are phytophagous. Some of the phytophagous midges do not really
give rise to galls on their host plants, but the vast majority produce
more or less complex galls on all parts of plants, particularly on
Dicotyledonae. The adult midges usually never feed, but the larvae
are apodous maggots, characterized by the presence of a peculiar
sclerotic sternal spatula (fig. 15 C). The larvae have well-developed
salivary glands. In some species the larvae have the ability to jerk
and shoot up into the air by suddenly straightening the curved body.
The larvae are also responsible for the galls. Pupation is sometimes
in the gall, sometimes in small silken-lined chambers underground
in the soil. Parthenogenesis and paedogenesis are known in many
species. Gall-midges have been extensively studied by RUBSAAMEN
996, 998, 999, KIEFFER 588 , FELT 328 , BARNES & MANI744. Some of the
widely distributed gall-midges belong to the genera Lasioptera
MEIG., Neolasioptera FELT, Trotteria KIEFF., Misospatha KIEFF.,
Oligotrophus LATR., Dasyneura, Asphondylia Low, Schizomyia,
Perrisia ROND., Rhabdophaga WESTW., Contarinia ROND., Mayetiola
KIEFF., Itonida MEIG., etc.

Fig. 15. Some typical cecidozoa. Itonididae (=Cecidomyiidae). A. Asphondylia

sarothamni H. Low. B. Mikiola fagi (Htg.) C. Larva of gall-midge, showing the
characteristic sternal spatula. D. Pupa of gall-midge, sticking out of its gall on
grass flower, just before the emergence of the adult. E. Lasioptera falcata Felt.
32
Among the other Diptera, Agromyzidae are of some importance
as gall-makers. A few species are known to give rise to galls in stems
or leaf veins of different plants. Some fruitflies like Tephritis, Lipara,
Dacus, etc. occasionally give rise to galls in flowers and stems of most
Compositae 315 . A nthomyia is also reported to give rise to galls occa-
sionally.
Cecidophyta
The plant organisms that give rise to galls on other plants include
the Bacteria, Algae, Fungi and Phanerogams. Though the cecidophy-
ta are on the whole numerically inferior to the cecidozoa, some of
them often give rise to nearly equally complex galls.
Bacteria.
The best known gall bacteria are undoubtedly Rhizobium b'3ye-
rinckii and Rhizobium radicicola of the root nodule galls of Legu-
minosae. Bacterial galls are also known on a number of other groups.
of plants. Phytomonas tumefaciens (SM. & TOWN.) gives rise to the
well known crown-gall (vide Chapters XIV and XV).
Fungi.
The fungi are the most important among the cecidophyta. They
are remarkable for the great numbers of gall-forming species and the
wide variety of plants on which they produce galls. The mycelia of
the gall fungi may be intercellular or also intracellular in the host
plant. Intercellular parasitic fungi produce haustoria for absorbing
nourishment or their mycelia penetrate the cell wall in all
directions. The gall fungi are chiefly Chytridiaceae, Ascomycetes,
Uredinae, Ustilaginae, Exobasidiaceae and Peronosporinae. PZas-
modiophora brassicae W ORON. is perhaps the best known gall fungus.
Some like the monadine A phelidium deformans give rise to unicellu-
lar galls on Coleochaete 1297 • Of the class Basidiomycetes three orders.
are well known as gall fungi, viz. Exobasidinae, Uredinae and Usti-
laginae. The Exobasidinae are characterized by free basidial layers,
so that the spores cover the external surface of the host plant. They
mostly attack the leaves of the host plant, which become more or
less greatly swollen in consequence. Exobasidium rhododendri
CRAM. gives rise to large spongy galls on leaves of Rhododendron spp.
Other species of Exobasidium give rise to galls on Vaccinium spp.
(fig. 141 B-C). The Uredinae1079 produce different spore types, such
as ascidia, uredospores and teleutospores. Very often all these three
or at least two of these types of spores occur on the same host plant.
The ascidiospores occur chiefly in spring and early summer. They
are rounded, thin-walled, unicellular, rust-coloured, yellow or red
spores that arise in chains, close together in beaker-shaped groups.
The uredospores have an elongate stalk and are likwise unicellular.
 33

The teleutospores are, however, thick-walled and retain their via-

bility for much longer periods than the other two types (fig. 142 F).
They are multicellular. Several fungi produce small, hemispherical
or flat spermogonia or pycnidia, inside which small cells called
spermato- or pycnospores arise. Some of the best known fungus galls
are the witches-broom galls caused by the ascidial generation of
M elampsorella caryophyllacearum SeHRoT. on Abies alba LINN.; the
teleutospores of this fungus develop on leaves of Caryophyllaceae.
Species of Puccini a are known to give rise to some other common
galls on plants like Urtica, Origamtm, Berberis, j asminum, Carex,
etc. Gymnosporangium spp. give rise to extensive galling of the stem
of junipert£s. Uromyces spp. give rise to galls on Euphorbia and
Leguminosae. The teleutospore generation of Calyptospora is also
known to cause galls and its acidial generation develops in the nee-
dles of Abies. Ustilaginae produce spores in the interior of the host
plant tissue, such as the ovary of Gramineae, which thereby becomes
more or less greatly swollen and contain ultimately a sooty powder.
Very often Ustilagineae produce also pustular galls on leaves. The
Ascomycetous fungi are also frequently gall makers. Their spores
develop in the interior of elongate tubular cells called asci. Taphrina
gives rise to a variety of galls. Even the Chytridiaceae, which are
intracellular parasites, give rise to galls. The commonest fungus of
this group belongs to Synchytrium. The white rust fungus Albugo
(Cystopus) candida PERS. is associated with characteristic galls on
Cruciferae, especially Capsella bursa-pastoris. Reference may also
be made to Uromycladium that gives rise to galls on Acacia spp.
1002, 1003, 1159

Algae.
Gall inducing species of Algae are met with among the green-
algae, red-algae and brown-algae. Phytophysa treubi is reported to
give rise to large galls on Pilea (Urticaceae). Many species of algae
are also known to give rise to galls on other algae 242 . Ectocarpus
valiantei gives rise to galls on Cystoseira ericoides. The alga Rhodo-
chytrium spilanthidis gives rise to extensive galling on Hibiscus
sabdariffa887, 301.
Phanerogams.
Loranthaceae, especially Viscum, Phoradendron, Loranthus, Cus-
cuta, etc., parasitic on diverse other plants, give rise to large woody
galls, with insignificant anatomical and histological abnormalities.
For an account of the development of the haustoria of the parasite
Cuscuta in the gall it produces reference may be made to THOMPSON
1149. The haustoria arise underneath the epidermis of the parasite.
The outer region of the modified epidermal cells of the parasite
becomes closely applied to the epidermis of the host plant. Once this
34
contact is established, the middle group of cells dissolve away their
way through the host epidermis by the action of an enzyme and
thus constitute the prehaustoria. The haustorium actually resembles
a root in most respects but lacks a root cap. It follows the path
prepared for it by the prehaustorium and it also dissolves away the
host cells. The hyphae appear first from the haustorial dermis and
undoubtedly represent modified root hairs. These hyphae often
grow along the inner sides of the interfascicular cambium and come
into contact with the xylem elements, but do not attack the phloem.
The cells of the hyphae now become lignified, forming a string of
tracheids from the host xvlem to the haustoria. Phloem and sieve
tissues do not arise in the "haustorium. Cell proliferation in the host
tissue provides abundant ready-made nutritive material. It is not
therefore the phloem of the host, but the xylem that is the principal
path of supply of water and carbohydrates to the parasite. MANGE-
NOT 742 has recorded an extremely interesting case of the roots of
Thonningia coccina (Balanophoraceae) coalescing with the roots of
another tree in contact and forming large galls. 'with numerous
absorbing strands.