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Q: Analyze how historical botanical expeditions contributed to current understanding and classification of Mexican pines.

Historical botanical expeditions, such as those by Roezl and Sessé, significantly contributed to the current understanding and classification of Mexican pines by collecting and documenting diverse taxa across various regions. These early collections, referenced in taxonomic revisions and monographs, provide baseline data for morphological comparisons and phylogenetic studies. The meticulous collection and naming practices have informed contemporary taxonomic frameworks, despite challenges in classification consistency and missing localities, underscoring the lasting impact of historical botanical research .

Q: What is the significance of seed dispersal mechanisms in the survival and proliferation of different pine species?

Seed dispersal mechanisms are crucial for the survival and proliferation of different pine species. Many pines have adapted to specific environmental challenges through specialized seed dispersal mechanisms. Closed-cone pines in areas with frequent and intense fires have developed serotinous cones that only open under extreme heat, which allows seeds to be dispersed and take advantage of cleared, fire-affected areas for regeneration . This is critical in fire-prone ecosystems where rapid colonization is necessary for survival. In contrast, some pinyon pines, such as those in juniper woodlands, have evolved mutualistic relationships with birds like the pinyon jay and Clark's nutcracker for seed dispersal. These birds transport and store seeds, aiding in their spread and subsequent germination, particularly important in habitats where other forms of seed dispersion might be limited . Additionally, species like P. maximartinezii, with large, hard-scaled cones, suggest possible adaptation to different dispersers, possibly birds like ravens, which might handle seed dispersal differently than the smaller seeds of pinyon pines . Overall, these dispersal strategies not only enhance the ability of pine species to regenerate in suitable areas but also maintain genetic diversity and resilience in varying ecological zones . Such mechanisms allow pines to effectively occupy diverse environments characterized by climatic and ecological extremes.

Q: Evaluate the effects of environmental changes on the morphological diversity observed within pine forests.

Environmental changes, such as alterations in precipitation patterns, fire frequency, and temperature, significantly impact morphological diversity within pine forests. Species like P. elliottii, with specific fire-adaptive traits, exhibit morphological adaptations that regulate community composition and structure. These changes shape growth forms, leaf characteristics, and reproductive strategies, influencing species distributions and interactions. The inability of some species to adapt quickly can lead to shifts in dominance, affecting ecosystem function and biodiversity conservation .

Q: Discuss the implications of resinous versus non-resinous buds in the adaptation strategies of pine species.

Resinous buds in pine species are associated with adaptation to withstand hydroclimatic variability, such as prolonged droughts. The resin provides a protective barrier against desiccation, pests, and diseases, offering a survival advantage in these conditions . Meanwhile, non-resinous buds may develop in species that do not require such protection, possibly due to their adaptation to less harsh environments or different ecological niches where the threat from pests and extreme weather is minimized . The presence of resin also differentiates pine species into hard and soft groups, with hard pines being more resinous and utilized extensively in turpentine production . This trait affects the pines' economic use and ecological roles depending on their adaptive strategies in specific environments.

Q: Compare the morphological and karyotypic similarities and differences observed among species in subgenus Strobus, particularly between P. flexilis and P. ayacahuite.

In subgenus Strobus, P. flexilis and P. strobiformis share similar morphological characters and karyotypes, suggesting a closer genetic relationship. However, P. ayacahuite differs noticeably both morphologically and karyotypically from these two, displaying variation that enhances adaptation to different environmental conditions. These differences emphasize the importance of integrating karyotype analysis with morphological studies for effective classification and understanding evolutionary relationships .

Q: What role do geographical and edaphic factors play in the distribution of pine species across different regions?

Geographical and edaphic factors significantly influence the distribution of pine species by affecting their ecological requirements and growth patterns. P. hartwegii, for example, is found at the highest elevations near the treeline in Mexico and Guatemala, where it often forms pure stands due to its adaptation to high-altitude environments . Similarly, P. cembroides and P. culminicola can be found in open woodlands and montane chaparral in northeastern Mexico, where specific soil composition, such as calcareous rock covered with deeper soils, supports their growth . These variations in pines at different altitudes and soil types underscore the importance of geographical and edaphic factors in their distribution.

Q: How do seed characteristics and cone morphology influence the ecological roles of pinyon pines?

Seed characteristics and cone morphology significantly influence the ecological roles of pinyon pines. Larger seeds with thicker coats and high starch and fatty oil content, as seen in P. maximartinezii, affect dispersal mechanisms and ecological interactions. Unlike the scrub jay-associated seeds of P. cembroides, these larger seeds suggest potential roles for other dispersers, like ravens. This differentiation in seed and cone attributes also impacts the adaptation to fire frequencies and habitat types, from open pine savannas to closed montane canopies .

Q: How do geographical clines in morphological characters demonstrate the complexity of taxonomic classification in pines?

Geographical clines in morphological characters demonstrate the complexity of taxonomic classification in pines by showing continuous variation across different regions, which may or may not justify taxonomic recognition. Taxonomic decisions must consider this clinal variation, particularly in widespread taxa, making it challenging to delineate clear species boundaries. This variation, often seen in migration histories and environmental responses, exemplifies the difficulty in interpreting chemical and morphological data taxonomically, as debated in historical surveys and modern classification approaches .

Q: What are some morphological differences that distinguish Pinus leiophylla from P lumholtzii, and why are they significant?

Pinus leiophylla and P lumholtzii differ in several morphological traits such as resinous versus non-resinous buds, leaf quantity per fascicle, leaf length, number of stomatal lines, and hypodermis uniformity. These differences are significant as they justify their classification within the subsect. Leiophyllae, reflecting evolutionary adaptations and ecological niches, despite the general similarity in their deciduous fascicle sheaths, which is an instance of parallel evolution in the genus .

Q: In what ways do anatomical features of Pinus leiophylla contribute to its identification and classification?

Anatomical features of Pinus leiophylla, such as bark thickness, leaf cross-sectional shape, hypodermis layer structure, and resin duct placement, are crucial for its identification and classification. The unique triangular cross-section of leaves, the number of stomatal lines on leaf faces, and detailed cone morphology with complex phyllotaxis provide identification markers that are essential for distinguishing it from related species and supporting its taxonomic placement .

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Organization for Flora Neotropica

Pinus (Pinaceae)
Author(s): Aljos Farjon and Brian T. Styles
Source: Flora Neotropica, Vol. 75, Pinus (Pinaceae) (Nov. 21, 1997), pp. 1-291
Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica
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.. .. .....

... . .. ....~~~~~~~~~~~~~~~~~~~~~~~~Wiiii~ii
..

... ..... . .. .~~~~~~~~~~~~~~~~~~~:

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

Brian T. Styles
Sepember1934 -27 Jue 199
26

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FLORA NEOTROPICA

MONOGRAPH75

PINUS (PINACEAE)
ALJOS FARJONAND BRIAN T. STYLES

PIC OP 1
1O CANCr

8^^ * -

FLORA(
NEOTROPICA

OP //

CAPIOCOIM

r11PIC
Publishedfor
Organization for Flora Neotropica
by
The New York Botanical Garden
New York
Issued 21 November1997

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InternationalStandardSerial Number0071-5794
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Printedin the United States of America using soy-based ink on recycled paper

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Library of Congress Cataloging in Publication Data

Flora neotropica.-Monograph no. 1-New York: Published
for Organization for Flora Neotropica by the New York
Botanical Garden, 1968-
v.: ill.; 26 cm.

Irregular.
Eachissuehas distinctivetitle.
no. 40.
Separatelycataloguedandclassifiedin LC beforemonograph
ISSN0071-5794= Floraneotropica.

1. Botany-Latin America-Classification-Collected works. 2. Botany-

Tropics-Classification-Collectedworks. 3. Botany-Classification-Collected
works. I. Organizationfor FloraNeotropica. II. New YorkBotanicalGarden.

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Pinus (Pinaceae)

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Flora Neotropica

Publishedfor
Organizationfor Flora Neotropica
by
The New YorkBotanicalGarden

Ariane L. Peixoto, Presidlent,UniversidadFederal Ruraldo Rio de Janeiro,Caixa Postal 74852, Seroped-

ica, 23851 Itaguaf,Rio de Janeiro, Brazil (1996).
Stephan Beck, Vice-President,HerbarioNacional de Bolivia, Casilla 10077, La Paz, Bolivia
(1996).
Enrique Forero, Executive Director,Ex Qfficio, Institute de Ciencias Naturales, Universidad Nacional de
Colombia, ApartadoAereo 7495, Bogota, Colombia.
Renee Fortunato, Herbario,Institute do Recursos Biol6gicos, I.N.T.A., 1712 Castelar, Buenos Aires, Ar-
gentina (1999).
Paul J. M. Maas, Treasurer,Instituteof Systematic Botany, Heidelberglaan2, P.O. Box 80.102, 3508 TC
Utrecht, Netherlands(1999).
Ghillean T. Prance, Royal Botanic Gardens,Kew, Richmond,SurreyTW9 3AB, United Kingdom (1996).
Laurence Skog, Secretarn,U.S. National Herbarium,Dept. of Botany, NHB-166, Smithsonian Institution,
Washington,D.C. 20560, USA (1996).
Rodolphe Spichiger, Conservatoireet Jardinbotaniquesde la Ville de Geneve, Case postale 60, CH-1292
Chambesy/Geneve,Switzerland(1996).
Wm. Wayt Thomas, The New York Botanical Garden,Bronx, New York 10458, USA (1999).
Paulo G. Windisch, UniversidadeEstadualPaulista(UNESP), Depto. de Botanica,Caixa Postal 136, 15001
S. Jose do Rio Preto, Sao Paulo, Brazil (1999).

Staff Committee
EnriqueForero,Chairman
Paulo G. Windisch, Ex Officio

Board Representativeto UNESCO

Alicia Lourteig,Ex Officio

EditorialCommittee
James L. Luteyn, Editor
S. Rob Gradstein,Editor

Claudio Delgadillo M. (1996-2001) MariaMercedesArbo (1993-1998)

LaurenceJ. Dorr(1996-2001) RodolpheSpichiger(1993-1998)
Jose R. Pirani(1996-2001) Olga Yano (1993-1998)

ORGANIZATION FOR FLORA NEOTROPICA

Founded and Conductedunderthe Auspices of UNESCO

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The Scientific PublicationsProgramat The New York Botanical Garden
thanksthe following donors for their generous supportof this publication:

MetropolitanLife Foundation
The TinkerFoundationIncorporated

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PINUS (PINACEAE)
ALJOS FARJON AND BRIAN T. STYLES

With Accounts of Wood Anatomy by IAN D. GOURLAY,

Pollen Morphology by MARIEH. KURMANN,
and Monoterpenes by JACQUELINES. BIRKS

CONTENTS
Abstract/Resum n ...........................................................................................................................................................
Preface (by Ghillean T Prance) ............................................................................................ 3
Introduction .................................................................................................................................................................... 4
Historical Survey ............................................................................................................................................................ 5
Morphologyand Anatomy ............................................................................................. 13
Wood Anatomy (by an D. Gourla ) ............................................................................................................................ 30
.......................................................................................................................................................................
KaryologyKaryology 35
Pollen Morphology(by Marie H. Kurmann)........................................ ............................................. 36
The Role of Monoterpenesin the Taxonomyof Mexican and MesoamericanPines (by Jacqueline S. Birks) ......... 38
ReproductiveBiology .................... ........................................................................... 41
Phylogeny and Evolution ............... ............................................................................... 42
Distributionand Ecology .............. ................................................................................. 49
Uses and Conservation................ ........................................ ........................................ 59
Systematic Treatment.....................................................................................................................................................
TreatmentSystematic 62
Subgenus Pinus ..........................................................................................................................................................
PinuSubgenus 71
Subgenus Strobus ............. ................................................................................. 198
Doubtful Names and ExcludedTaxa ........................................ ........................................... 255
Acknowledgements ........................................................................................................................................................ 256
Literature Cited ..............................................................................................................................................................
CitedLiterature 257
NumericalList of Taxa .................. ............................................................................ 267
List of Exsiccatae ...........................................................................................................................................................
ExsiccataeList 267
Index of Local Names ................ ................................................................................ 285
Index of Scientific Names ....................................................................................... 285

ABSTRACT

Farjon, A. (HerbariumDivision, Royal Botanic Gardens, Kew, Richmond, Surrey

TW9 3AB, U.K.) & B. T. Styles (Oxford ForestryInstitute, Departmentof Plant Sci-
ences, University of Oxford, South Parks Road, Oxford OXI 3RB, U.K.). Pinus (Pina-
ceae). Flora NeotropicaMonograph75: 1-293. 1997.-A revision of the pines native to
Mexico, CentralAmerica, and the Caribbeanis [Link] the 47 species recognized,
most are endemic to this region, but a number extend into Mexico from the United
States, while P. elliottii var. densa is included because it occurs on the Florida Keys in
the [Link] extensive synonymy has been largely resolved by (lecto-)typification.
Some taxonomic and nomenclaturalproblems have been addressedin previous papers;
only one new combination,P arizonica var. cooperi (C. E. Blanco) Farjon,is proposed
here. All species are fully [Link] introductioncovers all aspects of pines that
are of interest to both taxonomists and more general readers, and includes specialist
contributionson wood anatomy (I. D. Gourlay), pollen morphology(M. H. Kurmann),
and taxonomy based on monoterpenes(J. S. Birks).

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2 Flora Neotropica

RESUMEN

Farjon, A. (HerbariumDivision, Royal Botanic Gardens, Kew, Richmond, Surrey

TW9 3AB, U.K.) & B. T. Styles (Oxford Forestry Institute, Departmentof Plant Sci-
ences, University of Oxford, South Parks Road, Oxford OXI 3RB, U.K.). Pinus (Pina-
ceae). Flora Neotropica Monograph75: 1-293. 1997.-Se presentauna revision de los
pinos nativos de Mexico, Centroamericay el Caribe. La mayoria de las 47 especies
reconocidasson endemicas a esta regi6n, pero algunas se extienden hacia Mexico desde
los Estados Unidos, y se incluye P elliotii var. densa porque se encuentraen las islas
de Florida en el Caribe. La sinonimfa muy extensiva en el genero, se ha resuelto en
gran parte por medio de (lecto-) [Link] problemastaxon6micos y de no-
menclaturafueron discutidos en varias publicaciones anteriores;se propone aquf sola-
mente una combinaci6n nueva: P arizonica var. cooperi (C. E. Blanco) [Link]
las especies estan [Link] capitulos preliminaresincluyen todos los
aspectos de los pinos de internsparatax6nomos tanto para los lectores mas generales, e
incluyen contribucionesescritas por especialistas sobre anatomia de la madera (I. D.
Gourlay), morfologia del polen (M. H. Kurmann)y aspectos taxon6micos con relaci6n
al anilisis de monoterpenoides(J. S. Birks).

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PREFACE

Many people, tending to think of pines as a genus that is largely distributedin the
forests of the temperate region, are unawareof the importanceof the tropical pines.
Mexico and Central America have a great richness of this ecologically and
economically importantgroup, and so it is vital that they be monographedfor Flora
Neotropica. Twenty-five years ago the vision of this task came from the Oxford
ForestryInstitute,and in particularfrom Dr. BrianT. Styles, who was also co-authorof
the treatment of Meliaceae for Flora Neotropica (Pennington, 1981). He began his
work on the neotropical pines in 1970, and it was his greatest ambition to finish the
monographhe considered would be his magtnumtopus. During his years of studying
pines he travelled widely in Mexico and CentralAmerica and collected a large amount
of valuable material that has helped to resolve the taxonomy of several difficult
species-complexes. He also studied a vast amount of herbariummaterial and placed
this on a [Link] he publishedseveral papersabout the pines,
much of the informationhe had accumulatedremainedin his head, to be committed to
paper during the next few years. His notes and the drawings of each species which he
had had prepared by artist Rosemary Wise enabled the synthesis of his taxonomic
ideas. Unfortunately he was not to see the completion of this project during his
lifetime because of his untimely death in 1993, at the age of 58.
We are very fortunatethat a second specialist both existed and was free to devote
time to the completion of this [Link] too many taxonomic projects die along with
the specialist. However, in this case, Aljos Farjon stepped into the breach and pulled
together the many loose ends to produce this fine monograph. It is interesting and
fortunatethat Farjonagreed closely with Styles about the basic taxonomic framework
of the neotropical pines. Although much of the compilation and a great deal of
additional research has been carried out by Aljos Farjon, this monograph also
representsthe taxonomy that would have been producedby Brian Styles.
This commendable, and regrettably enforced, collaboration has produced an
outstanding monograph that is a most significant contribution to our knowledge of
tropical pines. It will stand as a fitting memorialto a generous person who was always
helping other people and who gave so much of his time to teaching botany to a large
numberof students, including me during my undergraduateclasses in plant taxonomy.
There are many of us who are indebtedto Brian Styles for his help, and so we are also
grateful to Aljos Farjon for taking up the difficult challenge of completing someone
else's work.
Ghillean T. Prance

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INTRODUCTION
Pinus (Pinaceae,Pinales), an exceptionally natural Mexico from the United States not laterthanthe Mid-
genus almost entirelyconfinedto the NorthernHemi- [Link] (1967) suggested that they entered
sphere, is representedin Mexico, Central America, Mexico along the Sierra MadreOccidental from the
and the Caribbeanby nearly one-half of its roughly North American Cordillera during the Late Creta-
110 species. The constituenttaxa show more features ceous or at the beginning of the Tertiary. Later
in common with each other (synapomorphies)than invasions may have been connected with floral re-
they share with any other genus in the family Pin- treatsfrom glaciations in the northernpartsof North
aceae, so the genus is distinctly monophyletic. In America and associated southwardshifts of biomes.
fact, during the last 50 years, only one species, P. Conversely, Axelrod (1980) and Axelrod and Cota
krempfiiLecomte-a rare, localized endemic in the (1993) postulated a Mexican pre-Miocene origin of
high mountains of Vietnam-has been placed in a the closed-cone pines (Pinus subsect. Oocarpae),
separategenus DucampopinusA. Chev., on account on the basis of plate tectonic history and present
of its paired, broad, flat leaves that lack a fascicle distributions. Millar (1993) assumed that Central
sheath at maturity,though otherwise it fits well into America may have been one of the refugia of pines
subgen. Strobus. The mature ovuliferous cone is duringthe Late Eocene cooling of the climate. Unfor-
typical of this subgenus, and in most recentrevisions tunately, fossil pines are virtually unknown from
it has been properlyincluded in the genus Pinus. Mexico, and none have been found in Central
The genus is distinguished from other members America or on any of the Caribbeanislands (Mirov,
of the Pinaceae by its needle-like secondary leaves, 1967;Axelrod, 1980; Millar, 1993). The earliest finds
borne singly or more commonly in fascicles of 2-8 in Mexico are Late Oligocene pollen found in amber
on short shoots (dwarf shoots), with a fascicle sheath deposits in Chiapas, which may themselves be of
of bud scales, present at least when young. Primary pines but cannot be definitively determinedas such.
leaves are retained in mature plants, but modified Pines are mostly pioneerspecies and probablyspread
into scariouscataphyllswhich are helically arranged, quickly on the lava flows thatcame from the volcanic
as are the leaves of the other genera, subtending activity occurringthen throughoutthe greaterpartof
the needle fascicles. The mature ovuliferous cone Mexico.
consists of few to many woody seed scales with It is not surprising, in a genus with such an
thickened distal parts (apophyses), each of which is extensive worldwide distributionas Pinus and with
subtended by a small, insignificantbract. All pines the diversity it displays in several of its characters,
are woody, and many form large forest trees, some thatthere is little agreementon the numberof species
up to 70 m tall. They frequently grow in stands, extant. Shaw's (1914) exceptionally conservative
forming extensive forests composed of a single spe- monographacknowledged only 66, whereas Gaus-
cies or a mixtureof several. sen's (1960) treatmentincluded 120. Critchfieldand
Although Florin (1963) held that the Pinaceaeand Little's (1966) geographic distribution maps show
the genus Pinus originated in the Northern Hemi- 94, while Mirov(1967) recognizedsome 103 species.
sphere during the Jurassic, subsequent evidence For Mexico the situation is not much better: Shaw
seems to point at a possible origin in the Triassic (1909) recognized only 19 species for Mexico, but
(Miller, 1977). The genus representsthe earliest evi- 40 years later Martinez (1948) counted 39 species
dence of the family. However, soon after this time there. Loock (1950) accepted 28 species for Mexico
the two major pine groups, Haploxylon (subgen. and Belize, and finally Perry (1991) recognized 58
Strobus)and Diploxylon (subgen. Pinus) had already species for Mexico and Central America. There is
differentiated. More recent interpretationsof many little doubt, given the number of new pines being
Jurassic fossil cones ascribed to Pinus have cast described each year, that the species concept is be-
doubts on this north-temperateorigin and have dated coming narrower still. Taxa formerly described at
the emergence of the genus later, perhaps at the the level of varietyor subspecies have been elevated
beginning of the Cretaceous(Axelrod, 1986; Millar, to the rankof species. This trend is ascribed mainly
1993). So far as the Mexican pines are concerned, to the use of chemical information,where if a popula-
their ancestors may have existed in North America tion differs from the species as a whole in the level
since the lower Cretaceous,where fossil cones simi- of a certain compound or in the absence of an
lar to those of P devonianahave been found(Chaney, enzyme, that population must be a different taxon.
1954). According to Miller (1977), the pines reached But chemical constituentscan be as variable as any
4

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HistoricalSurvey 5

other characters,and difficulties arise in interpreting has been so intensively studied as the genus Pinus.
them taxonomically,especially if such qualifications The taxonomic literature alone is extensive; with
are based on compound data (Birks, this volume). references to related disciplines added to this there
Much of this "splitting"has been proposed by non- are thousandsof relevanttitles (Farjon, 1990b). The
taxonomists, who do not seem to appreciatesuffi- synonymy to be dealt with is also substantial:for 47
ciently that widespreadtaxa often show geographical species and 20 infraspecific taxa here recognized,
variationsthat may or may not be worthy of taxo- more than 400 names have been proposed. This
nomic recognition. Given the known history of mi- monographis the first critical revision of the pines
grationof many pines, it is considered-and, indeed, of Latin America since Shaw (1909); it is the most
observed when assessed-that the variationin many comprehensiveassessmentof morphologicaland an-
of them is clinal in nature. Perhaps the most im- atomical data to date. It cannot definitively solve all
portant reason for the disagreement about species taxonomic problems: in particular,the delimitation
limits lies in the morphology of pine plants them- of taxa and their interrelationshipsin the species-
selves; taxonomistsare forced to base theirdecisions complexes of P. montezumnae, P ponderosa and P
on the variationfound in leaves and matureovulifer- pseudostrobus,and the relationshipsof disjunctpop-
ous cones. Leaves (needles) of pines show compara- ulationsand taxa in P cembroidesand its close allies
tively little variation, compared with the leaves of merit further investigation. However, new ap-
angiosperms,and may be greatly subjectto extremes proaches such as cpDNA variationanalysis may be
of [Link] cones are extremely similar more promising than furtherinvestigations into the
in most species. Thus the seed cones remain as the apparentlypolymorphicmorphologicaland chemical
chief source of taxonomic information. characters which have resulted in the conflicting
No othergroup of woody plantsof equivalentsize taxonomies.

HISTORICAL SURVEY

HISTORYOF COLLECTING F. L. von Schlechtendaland A. von Chamisso (1830,

AND DESCRIPTION 1831). A decade later,anotherGerman,C. G. Ehren-
berg, collected pine specimens in Mexico; the new
The earliest period of incidentalcollecting of pines, species were again described by Schlechtendal
1785-1914, is here brokendown into two periods. (1838) and by Gordonand Glendinning(in Gordon,
1858). A fourth German, a gardener named K. T.
I. 1785-1850 Hartweg,set out for Mexico to gathernew plantsfor
the HorticulturalSociety of London;he made further
Botanical collecting in both the Caribbean and additions,and it was throughhis specimens, widely
Mexico startedwith a few late seventeenth-and early distributed,that Mexico became known to host a
eighteenth-centurycollections, mainlyby Dutchtrav- diversity of these trees. The plant collection as a
ellers (Adams et al., 1987; Divila A. & GermanR., whole was enumeratedand described by Bentham
1991). The earliest botanical exploration started in (1840, 1842), who describedonly one new pine from
Mexico with the expeditions led by Martinde Sess6 Hartweg's gatherings, using a later homonym (P
in 1786 and 1788-1795, but no pine specimens from tenuifolia) to name it. Several more were described
these collecting activities are known. There may be by J. Lindley, Secretaryof the HorticulturalSociety,
some specimens of pines from the "Royal Botanical who was instrumentalin the distributionof some of
Expedition to New Spain" in 1790-91 in Madrid Hartweg's pine material to "subscribers"or to his
(MA). The oldest collections we have seen are two own botanical acquaintances,e.g., S. Endlicher in
specimens collected by M. A. Bonplandand A. von Vienna. F M. Liebmanncollected pines mainly in
Humboldt,now in Paris(P-Bonpl.).The earliest pine Oaxaca; B. C. Seemann collected several specimens
specimen from the Caribbeanwas collected by Olof on his second trip into the interiorfrom Mazatlanto
Peter Swartz, a pupil of Linnaeus filius, on Hispan- [Link] these was P lumholtzii,which was
iola in 1785. Althoughabundantin many areas,pines not recognized as a distinct species until Robinson
were apparently of little interest botanically until and Fernald (1894) described it based on Hartman
1828-1829, when C. J. W. Schiede & F. Deppe made 541 from near Coloradas, Chihuahua,collected in
several collections in Mexico, to be describedby D. May 1893. Liebmann's specimens went to various

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6 Flora Neotropica

herbariain the United States, and those of Seemann P engelnmannii (Wislizenus)

went to England (BM, K); no new species were P greggii (Gregg)
described from any of these. J. Gregg discovered a P hartwegii (Ehrenberg,Galeotti, Hartweg, Kar-
new species, which was named after him by G. winski)
Engelmann and described by Parlatore(1868). He P herrerae (Liebmann)
also travelled to northernMexico. The far north of P leiophylla (Ehrenberg, Gregg, Liebmann,
thatcountrywas exploredfromthe UnitedStates on a Schiede & Deppe, Seemann)
boundarysurvey,on which F A. Wislizenuscollected P. leiophylla var. chihuahuana (Seemann, Wis-
specimens of Pinus strobiformis,describedas a new lizenus)
species by Engelmann(1848). P. lumholtzii(Seemann)
These modest early collections from a vast and P mnaximinoi (Bonpland & Humboldt,Hartweg)
largely unexplored territory nevertheless yielded a P n ontezumae(Bonpland& Humboldt,Hartweg,
substantial portion of the species now known to Parkinson,Schiede & Deppe)
occur in Mexico. The pine forests of Mesoamerica P. montezumaevar. gordoniana (Hartweg)
(Chiapas and the Central American countries) re- P. occidentalis (Jaeger, Swartz)
mained virtually unknown-only Hartweg had col- P. oocarpa (Ehrenberg, Liebmann, Schiede &
lected a few specimens in Guatemala,described by Deppe)
Bentham (1842) as Pinus tenuifolia (= P max- P patula (Ehrenberg,Galeotti, Gregg, Hartweg,
iminoi). In the Caribbeana pine was known from Liebmann,Schiede & Deppe)
Hispaniola by two collections (P occidentalis), but P. pinceana (Ehrenberg, Ghiesbreght, Kar-
none were collected in Cuba and the Bahamas. winski?)
1. Collectors in Mexico 1785-1850, with number P pseudostrobus(Hartweg)
of discrete collections: P pseudostrobusvar. apulcensis (Hartweg)
P quadrifolia (Parry)
1806 M. A. Bonpland& A. von Humboldt(2) P strobiformis(Wislizenus)
1827 F. Karwinski(3)
P tecunumanii(Liebmann)
1828-1829 C. J. W. Schiede & F. Deppe (28) P teocote (Ehrenberg, Gregg, Hartweg, Lieb-
1837-1840 C. Ehrenberg(25)
mann, Schiede & Deppe)
1838, 1845-1847 K. T. Hartweg(22)
1839 J. Parkinson(I) By the end of 1850, a total of 21 species and 3
1840 H. G. Galeotti (2) varieties out of 47 species and 20 infraspecifictaxa
1841-1842 F. M. Liebmann(15) presently recognized had been collected-that is,
1843 A. B. Ghiesbrecht(1) nearly one-half of the species now recognized were
1846 F. A. Wislizenus (5) known in the first 50 years of botanical exploration
1847-1849 J. Gregg (9) of the region. More were named by then, but a
1850 C. C. Parry(1) numberhave been reduced to varieties (3 taxa, here
1850 B. C. Seemann (6) mentioned)or lost into synonymy.
2. The only collector in Central America during Although some collectors-e.g., K. T. Hartweg-
took care to collect pines purposefully,none of the
the period 1785-1850, was K. T. Hartwegwho made
2 discrete collections in Guatemala,1841-1842. early collectors had a special interest in them, and
3. Collectors in the Caribbean 1785-1850, with they were collected incidentallyby botanistsor pro-
fessional collectors with other, or at least much
number of discrete collections and place of collec-
tion: wider, interests in plants.

1785 0. P. Swartz (1, Hispaniola)

1828 B. Jaeger (1, Hispaniola) II. 1851-1914

4. Species collected 1785-1850 (in their current Early in this period, the curious efforts by B.
Roezl (for a more detailed account, see under P
taxonomy), with collectors names in parentheses:
montezumae)resulted in a substantialcollection of a
Pinus ayacahuite (Ehrenberg,Liebmann) limited number of species from Central Mexico.
P cembroides(Ehrenberg,Gregg, Hartweg, Kar- However, since they were almost all presented as
winski, Schiede & Deppe, Seemann) new species, it was never meant to be a collection
P devoniana(Gregg, Hartweg,Schiede & Deppe) to investigate the variationwithin species. The exact

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HistoricalSurvey 7

dates of his gatheringsare unknownbut seem to fall University who describeda new species: P. nelsonii,
between 1854 and 1858. His material was sent named after its discoverer. His monographon the
mostly to Europe as "vouchers" for seed batches, pines of Mexico (Shaw, 1909) was based primarily
which were sold to subscribersin the Horticultural on their collections. Shaw himself went to Mexico
Society of London and to well-known gardenersand in 1904, travelling with Pringle, to study his spe-
plantsmenlike George Gordon in Englandand E. A. cialty, the genus Pinus (McVaugh, 1972). His is the
Carrierein [Link] had given them names and, first trip in the history of pine collecting in Latin
in most cases, descriptionsin his catalogues (Roezl, America specially dedicated to the study of the ge-
1857, 1858). Both Gordon and Glendinning(1862) nus. Because Shaw entertaineda very broad species
and Carriere (1867), ill equipped to judge on the concept (Shaw, 1909; and especially Shaw, 1914)
taxonomyof Pinus in Mexico and perhapsalso eager there are several new species among the gatherings
to accept"noveltiesfor the garden,"acceptedmost of of Goldman,Nelson, and Rose which were only later
them. Schlechtendal(1857-1858) translatedRoezl's recognizedand described in the next period, usually
descriptionsfrom French into Latin; without having based on new collections.
seen a single specimen he merely commented that No pines had been collected from countries in
so many new species in Pinus seemed unlikely even Central America other than Guatemala until after
for Mexico. F. Parlatore,when in London in August the First World War. The collections made on the
1862, boughtRoezl's specimens from John Standish, CaribbeanIslands during this period were still few.
gardener of the Duchess of Gloucester at Bagshot J. A. Shafer, gatheringin Cuba, made an exemplary
Park, Surrey: on 14 October 1862, a total of 435 exception with 30 collections of the two species
specimens of conifers were entered in the registerof occurring in western Cuba; for the first time in
accessions to the Botanical Museum in Florence. the history of pine collecting there was a collector
Parlatore(1868) made short shrift of Roezl's species gatheringmaterialof a species in sufficientquantities
and put them all into synonymy. to assess morphological variation among popula-
Very few pine collections were made until 1885- tions. Othercollectors before him had only acciden-
1886, when two professional plant collectors, E. tally, not purposely, made multiple gatherings of a
Palmer and C. G. Pringle, started making general single species. This was to change fundamentally
plant collections on a large and systematic scale. in the next period: although incidental collecting
Palmer's plants were enumeratedand described by continued until the present, a more systematic ap-
Sereno Watson at HarvardUniversity, but the pines proach was adopted by many collectors who now
were entrustedto Engelmannat the MissouriBotani- took a special interestin the pines themselves.
cal Garden, who described Pinus radiata (as P in- 1. Collectors with -10 discrete collections in
signis) [Link] from Palmer's 1875 gatheringson Mexico, 1851-1914, with numberof discrete collec-
GuadalupeIsland. Pringle was a most prolific plant tions:
collector, and it was his custom to make sets of 60
duplicates whenever possible (McVaugh, 1972), but 1854-1858 B. Roezl (84)
sets of 12, 15, or 20 are also common, and there (1875) 1885-1908 E. Palmer (26; 3 with C. C.
are even (unnumbered)unicates. As a consequence, Parry)
Pringle's specimens were distributedto many herb- 1886-1907 C. G. Pringle (96)
aria. His earliest pine specimens are from Baja Cali- 1898-1908 E. H. Goldman(28)
fornia (P quadrifolia, collected in 1882). Very few 1892-1903 E. W. Nelson (106; 3 with E. H.
collections of pines were made by him in the south- Goldman)
ern states of Mexico (one only, from Oaxaca, has 1897-1911 J. N. Rose (48; 10 with R. Hay; 5
come to our attention),and he did not visit Central with W. Hough; 8 with J. H. Painter)
America. New species were described by Shaw 1904 G. R. Shaw (47)
(1905a, 1905b, 1905c), of which P pringlei is here 1903-1913 C. A. Purpus(18)
accepted. E. W. Nelson-sometimes, as in 1897, 1905 J. G. Lemmon (13)
with E. H. Goldman and J. N. Rose (the latter 1907-1912 G. Arsene (24)
numberedhis specimens separatelyas he worked for 1910-1911 A. Arsene (10)
the U.S. National Museum)-collected for the U.S.
Departmentof Agriculture,and those main sets of 2. Collectors with >5 discrete collections in Cen-
specimens are kept in Washington(US). It was again tral America, 1851-1914, with number of discrete
G. R. Shaw at the Arnold Arboretumof Harvard collections:

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8 Flora Neotropica

1876 C. G. Bernoulli & R. Cario (8) P pseudostrobusf. protuberans(Pringle)

1890-1892 J. Donnell-Smith(6) P radiata var. binata (Palmer, 1875, Rose, 1911)
1895-1896 E. W. Nelson (6) P tropicalis (Clinton-Baker,Shafer, Wright)
1904-1907 H. von Tuerckheim(5)
1905 W. R. Maxon & R. Hay (6) By the end of 1914 an additional 15 species and
1905-1907 W. A. Kellerman(14) 9 infraspecifictaxa had been collected. More often
than duringthe previous period, these nomenclatural
3. Collectors in the Caribbean, 1851-1914, with innovations involved taxa only much later to be
numberof discrete collections: recognized as distinct, e.g., Pinus douglasiana, P
1853 R. H. Schomburgk(1) durangensis, P praetermissa, and the infraspecific
1856-1865 C. Wright (8; I with C. C. Parry,C. taxa in P. arizonica and P cembroides.
The contemporaryperiod (1915-present) of sys-
Wright& Brummell)
1887 H. F A. von Eggers (1) tematic collecting of pines is broken down into two
1903-1912 J. A. Shafer (30) periods.
1903-1905 G. V. Nash & N. Taylor (2)
1904-1914 HermanoLe6n [brotherLe6n] (2)
III. 1915-1950
1910 H. von Tuerckheim(1)
1911 H. Clinton-Baker(1) We characterizethe time after the FirstWorldWar
1914 E. L. Ekman (3) to the present as the period of systematic collecting
1878-1914 e.g., L. J. K. Brace; J. K. Small & of pines. With few exceptions (Shaw, Shafer), speci-
J. J. Carter(16, Bahamas) mens of the genus collected in the foregoing period
had been gathered by people who were out in the
4. Additionalspecies collected 1851-1914 (in their
field making general collections of plants, often fo-
currenttaxonomy), with principalcollectors' names
in parentheses: cusing on entirely different taxa. Even Roezl, who
at one time was involved with pines, is much better
Pinus arizonica (Nelson, Palmer) known for the great number of orchids and other
P arizonica var. cooperi (Pringle) plants he collected, for which he undertook much
P arizonica var. stormiae (Palmer) wider exploration than he did for the pines. To be
P. ayacahuite var. veitchii (Roezl, 1857, lost, sure, general plant collecting continued, and these
Pringle) efforts brought in pine specimens as well. But in-
P caribaea (original materialdescribedby More- creasinglywe encountercollectors who are interested
let 1851, lost; Rowlee, 1901, Le6n brothers) in the genus Pinus itself, primarilyor exclusively.
P caribaea [Link](earliestBrace, 1878) Native-borncollectors are beginning to play an im-
P caribaea var. hondurensis (original material portantpart in the collecting; it is no longer solely
described by S6enclauze 1868, lost; Ber- the concern of Europeansor Americans.
noulli & Cario, Donnell-Smith, Kellerman, Partly of Mexican descent and later living and
Maxon & Hay) working in California, Y. Mexia made an extensive
P cembroides var. bicolor (Nelson, 1892) trip to the western states of Mexico in 1926-1927
P cembroidessubsp. lagunae Nelson & Goldman, making general collections for the University of
1906) Californiaat Berkeley (UC). She was the first female
R contorta var. murrayana(Goldman, 1905) plant collector to make substantial collections of
P cubensis (Ekman, Shafer,Wright) pines in Mexico. E. Matuda, a Mexican collector,
P douglasiana (Nelson, Rose) was very active in Chiapas from 1936 to 1950 and
P durangensis (Pringle, Rose) collected almost all of his specimens of pines there.
P flexilis var. reflexa(Nelson) Yet he did not discover P strobus var. chiapensis,
P jeffreyi (Goldman, 1905) and his collections cover the common species. One
P lambertiana(Goldman, 1905) of these is P tecunumanii,not yet recognized; his
P lawsonii (Pringle, Purpus,Roezl, Shaw) collections of this species were considered to be of
P nelsonii (Nelson, Pringle, Shaw) P. oocarpa or P patula.
P oocarpa var. trifoliata (Rose) M. Martinez(1888-1964), who was Professor of
P praetermissa (Palmer,Rose) Botany at the Universidad Nacional Aut6noma de
P pringlei (Nelson, Pringle) Mexico in Mexico City and a prolific writer on the

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HistoricalSurvey 9

Mexican flora, did not collect pine specimens him- 1926-1938 Y. Mexia (20)
self. They were obtained for him by other persons, 1933-1937 G. B. Hinton (21; 4 by Hinton et al.)
but he assigned his own numbersto the specimens 1933-1939 C. H. Mueller (25; 14 with M. T.
"for ease of citation and reference" (McVaugh, Mueller)
1972). He rarely mentioned the person who made 1935-1950 H. S. Gentry 18; 3 by Gentryet al.)
the gathering,an exception being C. E. Blanco, for 1936-1950(-1959) E. Matuda(110)
whom he named a variety of P arizonica. The pine 1937-1944(-1954) M. Martinez(216, not by him-
collections date from 1937 to 1950, with a few to self)
1954. Their collection numbers correspond in no 1938-1941 S. S. White (28; 1 with R. M. Chat-
way to these dates on the labels. The informationis ters)
written in Martinez's hand and no original labels 1929-1930, 1938-1948 I. L. Wiggins (10; 6 with
exist; obvious errorsencounteredmake it clear that D. Demaree in 1930; 3 with J. W. Gillespie in
the information is not entirely trustworthy,but it 1929)
cannot be checked. The collections (both unicates
2. Collectors with '5 discrete collections in Cen-
and duplicates) are in MEXU and in part also in
tral America, 1915-1950, with number of discrete
herbariain the United States. Martinez described a
collections:
few new species and numerous new infraspecific
taxa based on these collections. Most of the latter 1922, 1928-1950 P. C. Standley (53; 2 with P.
appearto be mere local forms or variantsof limited Chacon; 1 with L. O. Williams)
taxonomic meaning; in part this is due to the fact 1932 J. B. Edwards(5)
that Martinez,working during the difficult years of 1933-1934 A. F. Skutch (8)
the Second WorldWar,had no access to collections 1938-1939 H. von Schrenk(5)
outside Mexico and limited access to literatureand 1939-1942 J. A. Steyermark(30)
few contacts with other specialists. However, his 1943-1944 J. Valerio(3; 3 with Rodriguez)
lasting contributionsto the taxonomy of Pinus in 1945-1946 A. J. Sharp(8)
Mexico are several (some of which are listed below), 1946-1950 L. O. Williams (2; 5 with A. R.
while others (new varieties of P arizonica and new Molina)
species P douglasiana, P durangensis, P herrerae, 1947-1948 L. R. Holdridge(5)
and P michoacana = P devoniana) were described
3. The only collector with 25 discrete collections
by him but had in fact already been collected and
in the Caribbeanduring the period 1915-1950 was
deposited in herbariaoutside Mexico well before his
time. E. L. Ekman, who made 20 collections from 1915
In Central America, it was the collecting and to 1927.
floristicwork by P. C. Standleyand J. A. Steyermark 4. Additionalspecies collected 1915-1950 (in their
currenttaxonomy), with principalcollectors' names
during this period, culminating in their Flora of
Guatemala (1958), published in Fieldiana, which in parentheses:
established the basic taxonomic knowledge of the Pinus attenuata (Martinez)
genus in that country and beyond. Collections were P coulteri (Martinez)
also made in Belize, Honduras, El Salvador, and P culminicola (Hinton, Mueller)
Nicaragua,but these yielded no new taxa of Pinus. P monophylla(P. Litchey 1936, Martinez, 1939,
We now know that all of CentralAmerica's pines,
Wiggins, 1946, Gentryet al., 1950)
even P caribaea var. hondurensis, also occur in P muricata (Gentry,Martinez)
Mexico. P patula var. longipedunculata(Martinez, coil.
Only E. L. Ekman gathered more than a few E. E. M. Loock)
numbersof Pinus in the Caribbeanduringthis period. P ponderosa var. scopulorum (E L. Wynd &
He collected pines in Cuba and Hispaniola; they C. H. Mueller, 1936, J. M. Tucker, 1952)
were treated by Pilger (1926) and later by Florin P remota(Mueller)
(1933), who described the distributionof the pines P strobus var. chiapensis (Martinez)
on these islands.
1. Collectors with >15 discrete collections in In this period, the additions to the Mexican flora
Mexico, 1915-1950, with numberof discrete collec- resulted mainly from the finding of more localized
tions: taxa, notably the extension of four Californianspe-

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10 Flora Neotropica

cies into Baja California(P attenuata,P coulteri, P 1. Collectors with >20 discrete collections in
monoophylla, and P muricata)and the subalpinespe- Mexico, 1951-present, with numberof discrete col-
cies P culminicola, the latter not recognized as new lections:
until 1961. The first four were collected for Martinez 1949-1975 R. McVaugh(67; 3 with W. N. Koelz)
by anonymous collectors; of the last he had no 1951-1955 J. H. Maysilles (40)
knowledge (Martinez, 1945, 1948), even though it 1951-1964 E. Matuda(49; 3 by Matudaet al.)
had been collected years before by two independent 1952-1982 R. Moran(82; 6 by Moranet al.)
collectors. Two varietiesare taxa describedby Marti- 1952-1985 J. T. Marshall(24)
nez which are recognizedhere;the remarkableoccur- 1952-1989 J. Rzedowski (95; 2 with G. B. Hin-
rence of a variety of P strobus in southernMexico
ton; 7 with R. McVaugh)
and Guatemala is the more noteworthyof the two. 1955 B. Zobel (21)
The occurrenceof P ponderosa var. scopulorumjust 1957-1960 L. Kaplan(19; 4 with E. L. Davis)
inside Mexico is known from the two collections 1957-1961 J. H. Beaman (19; 9 with J. W. An-
mentioned;all other collections under P ponderosa dresen)
from Mexico studied by us appeared to be other 1959-1980 X. MadrigalS. (165; 22 by Madrigal
taxa. S. et al.)
1960-1963 E. L. Little (170)
IV. 1951-Present 1960-1974 L. Vela G. (26)
1962-1980 D. E. Breedlove(54; 21 by Breedlove
Of ca. 8500 collections in our database, the vast et al.)
majority, roughly 6000, has been gathered since 1963-1970 A. May Nah (64)
1950. More recently, since around 1970, they have 1967-1982 R. HerndndezM. (8; 17 by Hernandez
increasingly been accumulatedas vouchers to seed et al.)
collections by collectors working for forest geneti- 1970-1971 M. A. Capo (35)
cists or by the researchersthemselves. They have 1971-1984 R. A. Bye (23; 5 by R. A. Bye et al.)
been deposited in a limited number of herbaria, 1972-1974 B. T. Styles (85; 14 with R. F. Kemp)
e.g., FHO in Oxford and NCSC in Raleigh, North 1974-1983 D. K. Bailey (25; 19 by Bailey et al.)
Carolina. One recognizes such specimens collected 1975-1984 A. A. Ventura(20)
for this purpose often by their bound-up foliage, 1977-1978 T. Eguiluz (24)
which makes them virtuallyworthless for herbarium 1977-1981 W. L. Mittak(102)
studies. Emphasis is on species and "provenances" 1978-1981 J. W. Stead & B. T. Styles (145 + ca.
of these, which have a putativepotentialin plantation 240 as "populationcollections"; 7 by Stead et
[Link], 5-20 trees were sampledof a species al.)
in a stand or otherwise limited area ("population 1979-1985 J. C. Soto N. (7; 30 by Soto N. et al.)
collections"; such collections have been enumerated 1981-1986 M. Nee (11; 65 by Nee et al.)
with serial numbersin the list of exsiccatae). Because 1982-1991 J. A. Perez de la Rosa (25)
pines extend into the tropics in our region, there 1983-1984 K. Taylor & M. Nee (22)
was much interest in collecting taxa from the more 1983-1985 P. S. McCarter (7; 24 with C. E.
southernparts of their ranges, which might be used
Hughes; 33 with B. T. Styles + ca. 100 in
in other tropical countries. The data presented here
"populationcollections")
for this period may therefore be biased, despite 1983-1993 C. E. Hughes (34; 19 with B. T.
thoroughscrutinyof collections in many of the major Styles)
herbaria,especially by B. T. Styles. 1984 K. D. Rushforth(20)
Botanists were active, too, and they are largely 1984-1986, 1991 B. T. Styles et al. (40)
responsible for the collecting of the remainderof 1992-1994 A. Farjon(5; 33 with D. Mejia)
species and for the discovery of new ones. In the 1993 S. Higman et al. (52)
late 1980s and early 1990s the emphasis returnedto
botanical collecting, although some organizations- 2. Collectors with >20 discrete collections in
Central America, 1951-present, with numberof dis-
e.g., CAMCORE, Raleigh, North Carolina (not in-
cluded in list of exsiccatae)-continued to make crete collections:
vouchercollections of seed-samplingefforts through- 1951 F. Schwerdtfeger(36, Guatemala)
out the ranges of a number of timber-producing 1956-1975 A. Molina R. (28; 12 by Molina R.
species. et al.)

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History of Classification I1

1957 H. A. Luckhoff
HISTORYOF CLASSIFICATION
1969-1972, 1977 R. H. Kemp (123)
1972, 1986 B. T. Styles (5; 82 with R. H. Kemp) The genus Pinus was established by Linnaeus
1972-1975 A. Greaves (20) (1753, 1754) with 10 species, of which only five are
1975-1979 G. Chaplin (32; 1 with A. Greaves) truepines (Pinus [Link].),the othersbelonging to Abies
1975-1981 W. L. Mittak(308 + ca. 150 "popula- (2), Cedrus(1), Larix (I), and Picea (I) in Pinaceae
tion collections"; 5 with E. Espinosa; 13 with (Farjon, 1990a). The earliest segregation of Lin-
J. P. Perry,Guatemala) naeus's species in Pinus [Link]. was by Miller (1754).
1977-1982 J. W. Stead (132 + ca. 250 "population Numerousproposalsto furthersubdividethe genus
collections"; 160 with B. T. Styles) Pinus have been made, the firstof these by Duhamel
1979, 1985-1987 A. Grijalva(7; 1 with M. Ara- du Monceau(1755) on the basis of numberof leaves
quistain; 13 with E. Sediles, Nicaragua) per fascicle. Mirov (1967) discussed some major
1981-1983, 1993 C. E. Hughes (68; 4 with B. T. classifications and gave lists of species with each.
Styles) Little and Critchfield (1969), in their well-docu-
1983-1988 P. S. McCarter(55; 20 with C. E. mented classification,gave a review of the previous
Hughes;64 with B. T. Styles + ca. 100 "popula- principalclassifications. A review of 10 classifica-
tion collections";6 with Hughes & Styles) tions, from Shaw (1914) to van der Burgh (1973),
was furthergiven by Landry(1978). We think it is
3. Collectors with 210 discrete collections in the
appropriateto refer to these reviews here instead of
Caribbean, 1951-present, with number of discrete
repeatingthem entirely, but a few salient examples,
collections and place of collection:
pertainingto two majorsystems of subdivision"com-
1956-1957 H. A. Luckhoff (13, Bahamas) peting" up to the present time, may be highlighted.
1966-1971 J. Bisse et al. (13, Cuba) These two systems differ in that one recognizes a
1968 A. F. A. Lamb (30, Cuba) basic binarydivision in the genus, whereasthe other
1980-1988 T. A. Zanoni et al. (20, Hispaniola) divides the genus into more than two groups at the
1988 W. B. H. Baylis (28, Bahamas) highest subgeneric rank recognized. The singular
segregation of P krempfii,an aberrantspecies also
4. Additional species collected 1951-present (in placed in a separate monotypic genus, in otherwise
their current taxonomy), with principal collectors' binary classifications, is not counted as a deviation
names in parentheses: from that system.
Pinus cembroidessubsp. orizabensis (C. T. Hart- Secondly, as already accentuated by Little and
Critchfield(1969), there are nomenclaturalimplica-
weg, before 1850 [lost]; M.-F Robert, 1970; tions which, despite a comprehensive treatmentby
Bailey, 1980)
these authors, are still being ignored in much of
P jaliscana (S. Carvajal,1982; Perez de la Rosa,
the more recent literatureon pine phylogeny and
1983; Farjon& Mejia)
[Link] most often encounterednomencla-
P maximartinezii (Rzedowski, 1964; Bailey,
tural error is the failure to recognize the ruling of
Hughes, Farjon& Mejia) Article 22 of the Code (e.g., see Greuter et al.,
P rzedowskii(MadrigalS., 1968; Farjon& Mejia)
1994), which prescribesthe use of the genus name
Despite the enormous increase in the number of ("autonym")as the epithet for thatpartof any subdi-
collections made since 1950, only four additional vision of a genus that includes the type of thatgenus.
taxa were discovered, all in Mexico. Of Hartweg's The most widely acceptedsubdivisionof the genus
original collection nothing remains; this subspecies Pitnusis the binarydivision into two groupsof equal
of P. cembroides was rediscovered by Bailey, al- rank,each usually subdividedfurtherinto groups of
though it had earlierbeen collected by M.-F Robert species. The firstauthorto proposethis was Parlatore
without recognizing it as distinct. Two very remark- (1868), who recognized within Pinus subgen. Pinus
able new species were discovered, both narrowen- the two sections ("sectio")Pinea and [Link]-
demics: P maximartinezii and P rzedowskii; the mann (1880a), treatingthe genus in its strict sense,
latest new discovery is P jaliscana, a species allied divided it (in a key to the species) into two sections,
to P [Link] in the "pinyonpines"other Strobus and Pinaster. It is importantto note that
new taxa have been describedduringthis period,but Engelmanntried to use several independentcharac-
we have not recognized them as distinct from those ters (which admittedly did not all fit equally well)
which had been named earlier. insteadof only one (see below). Section Strobuswas

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12 Flora Neotropica

characterizedby (terminology modernized here) a Martinezabandonedthe binaryarrangementsof pre-

terminalumbo, deciduous fascicle sheaths, leaves in vious authorsand returnedto the system of the early
fascicles of 5, microsporophyllswithout a terminal period of classification.
crest, and wood less dense and less resinous. Section An entirely differentschool of thoughtwas devel-
Pinaster had a dorsal umbo, usually persistentfasci- oped by Campo-Duplan(1950) and de Ferre (1953,
cle sheaths, a variableleaf number,microsporophylls 1965), based on ideas of evolution in conifers held
with a semi-orbicularcrest, and denser, more resin- by [Link] led to detailedobservationalstudies
ous wood. Notably, this led to his grouping of P of those charactersbelieved to have evolved much
cembroides and its allies, as well as P balfouriana, more than others, e.g., charactersof pollen (size and
with sect. Pinaster; these taxa have been classified shape) and seedlings (cotyledons), and to the ignor-
with (subgen.) Strobus in most later binary classifi- ing of others believed to be primitive, e.g., the
cations. This binary treatment of species groups morphologyof the female reproductiveorgan. For a
within the genus was accepted by the authors of critique see van der Burgh, 1973. The genus was
major"NaturalSystems"of plantclassification,such accordinglysubdividedinto two subgenera(Campo-
as Bentham and Hooker (1880) and Eichler (1889), Duplan, 1950) on the basis of pollen morphology,or
as well as by Sargent(1897) in his authoritativetree into four subgenera (de Ferr6, 1953, 1965) on the
manual, which certainly stimulated its acceptance. basis of seedlings, the latter with further subdivi-
Lemmon (1888, 1895) was the first author to use sions. Landry(1974, 1978, 1994) reasoned contrary
the rank subgenus for the two main subdivisions to Gaussen(1960), who had rejectedcone characters,
designated previously as sections by Engelmann,for and believed in a Linnaeanprincipleof classification:
which the currentcorrectcitations are Pinus subgen. since the greatmasterhad used flowers,the reproduc-
Strobus (Sweet ex Spach) Lemmon and Pinus sub- tive organs of pines must give the clue to an unam-
gen. Pinus [not Pinaster (Engelmann)Lemmon]. biguous classification. Although Landry has been
Koehne (1893) discovered that a binary division correct in recognizing some distinctive taxa (Pinus
of the genus was supportedby the presenceof either leiophylla: Landry,1992; P nelsonii: Landry,1994),
one or two vascular bundles in the leaf and conse- his classification,like those from the school of Gaus-
quently proposed yet another set of names for two sen, must be rejectedas entirely artificial.
sections: Haploxylon(I vascularbundle)and Diplox- Few other contemporaneousinvestigations using
ylon (2 vascular bundles). According to this single a limited number of charactershave proposed new
characterscheme, P balfouriana and P. cembroides classifications for the genus. We mention Hudson
have to be classified with Haploxylon (= Strobus). (1960) on wood anatomy (non-binary) and Jahrig
These two "appropriatelydescriptive names" (Lit- (1962, 1968) on leaf anatomy (binary). The con-
tle & Critchfield, 1969) are still widely used, espe- trasting method, phenetic classification in the sense
cially in non-taxonomicliteratureand at subgeneric of Sokal and Sneath (1963), has never been applied
rank,despite their illegitimacy underthe rules of the to the genus as a whole, althoughclassifications like
Code. those of Little and Critchfield (1969) and van der
Non-binaryclassificationshave been proposedand Burgh (1973) are phenetic in the sense that they
were adoptedas alternativesto the basic division into group taxa on overall similaritycomparing multiple
two groupsas outlinedabove, and indeed precededit. characters.
Early examples are Spach (1841), with a division Increasingly, evaluation of special characters is
into four sections; Endlicher(1847), with six sections being used to assess phylogenetic relationshipsand/
in Pinus [Link].; and Koch (1873), with six "Gruppe," or to test previously proposed "traditional"classifi-
presumablycorrespondingwith the rank of section. cations. Most papersinvestigate only a limited num-
Nevertheless, all classificationsof the (entire) genus, ber of species traditionallyclassified in a section or
proposed between 1890 and 1940, recognized the subsection of the genus; these are not considered
binary division of the genus, either as subgeneraor here, as we are interested in the history of the
sections, albeit undera numberof (now illegitimate) classification of the genus as a whole. Most studies
names. of monoterpenes,for instance,fall into this category.
Martinez(1945, 1948) arrangedthe Mexicanpines Yet there are a few chemical studies that did find
into nine sections, some of which were furthersubdi- differences between the two subgenera,such as Erdt-
vided into "groups."None of these were formally man (1959) on heartwood polyphenols. Molecular
proposed with a Latin description and some had techniques,constantly improving,have made it pos-
Mexican, not Latinized, names. Yet it is clear that sible to study genetic variationanalysing allozymes

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Morphologyand Anatomy 13

and even more directly in the DNA itself. Examples study using morphologicalcharactersof the pinyon
of allozyme studies are those by Karalamangalaand pines, arrivedat a similarconclusion. Classifications
Nickrent (1989), who looked at variationamong 14 ought to be based on phylogeny, investigated by
Mexican and southwesternU.S. species in subgen. means of the evaluationof a numberof independent
Pinus ("Diploxylon"),and by Shurkhalet al. (1991), characters,consistent with a testable theory of the
who studiedEurasianrepresentativesof both subgen- evolution of the genus. The recent molecularevalua-
era. A phylogenetic analysis based on nuclear DNA tions have been, and promise to continue to be,
(ribosomal DNA) was carried out by Govindaraju helpful test cases for existing classifications as well
et al. (1992) assessing 30 species. The chloroplast as sources for useful taxonomiccharacters;a synthe-
genome, paternallyinheritedin conifers, appearsin sis with anatomical and morphological data seems
these studies to give the most reliable results on the way forward.
relationshipsbetween taxa, includinghybridrelation- A final remark on a recent(?) trend in conifer
ships (Wang, 1992). A numberof papershave studied taxonomy and classification is relevant here. There
chloroplast-DNA polymorphisms, using restriction has been a consistentnarrowingof the genus concept,
fragmentlength analysis, in a (random)selection of both in less well-studied families such as Podocar-
species assigned to several traditionalgroups (sub- paceae and in intenselystudiedones such as Pinaceae
genera, sections), and evaluatedthese using a cladis- (e.g., Page, 1989, 1990). How well this can be
tic method (usually presenting parsimony clado- justified in the light of the evolutionary history of
grams) to investigate the phylogeny of that sample conifers generally is not a point of discussion here.
of taxa. Examples of such cpDNA analyses are Whatmust be questionedis the assumptionthatsince
Szmidt et al. (1988), with 20 species studied, Strauss the (morphological)differencesbetween some of the
and Doerksen (1990), with 19 species tested, Wang segregate genera appear to be small, a large and
and Szmidt (1993), with 18 species, and Perez de la polymorphicgenus such as Pitins ought to be broken
Rosa et al. (1995), using 12 species. Only the last of up as well. Divergence, caused by the "accumula-
these studied Mexican pines. tion" of autapomorphies,may lead to relative differ-
The majorityof these investigationstends to sup- ences between otherwise perfectly monophyletic
port a binary classification of the genus; at lower taxa. Contraryto the "rule"still maintainedby some
levels of hierarchy,several supportLittle and Critch- cladists, that each clade (sister group) should be
field (1969) and similar classificationsgenerally,but given equal rank or that all nodes representnatural
give indications that some groupings, such as the taxa and should consequentlybe ranked,it may well
heterogeneoussect. Parryanae,are paraphyleticand be valid not to do so (Stace, 1989).
need to be revised. Malusa (1992), in a phylogenetic

MORPHOLOGY AND ANATOMY

This section comprisesan accountof the morphol- ally, resulting in a hierarchical,symmetricalsystem

ogy and anatomy of habit, shoots, vegetative struc- of branchingfrom the centre toward the periphery,
tures, bark, and [Link] and known as "Rauh's growth model" (Halle et al.,
pollen are described in separatechaptersby special- 1978). The characteristicsare an erect leading shoot,
ists, in acknowledgementof their responsibilityfor assurgent higher-orderbranches, and a non-plagio-
these contributions;their treatises immediately fol- tropic disposition of the branchesof all subsequent
low this chapter. orders, usually 4-5. Normally, shoot elongation is
uninodal, i.e., during the growing season a single
GROWTH FORM AND system of apical and lateral primordia of stem,
leaves, and reproductiveorgans, initiatedin a previ-
BRANCHING PATTERN
ous season and forminga terminaland severalsubter-
The genus Pinus consists of perennial,arborescent minal "winterbuds," elongates and develops at the
species, many of which attain great age and size. ends of stem and branches;growththen ceases and a
The principalbranchingpatternis monopodial,of a new set of primordiais initiatedbefore the dormancy
type in which the lateral branches are (pseudo-) period. In a number( 10 in our region) of species
[Link] apical shoot meristemdominates over of subgen. Pinus, multinodal shoots occur or are
the laterals;this patternis repeated,usually season- [Link] are likewise initi-

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14 Flora Neotropica

ated in the winterbuds (see below), and development very low spiral), but they are cyclic (whorled) in
of the vegetative organs, as well as of the reproduc- Pinus. Primary leaves continue to develop on the
tive organs, especially ovuliferous cones, occurs si- branchingshoot each season in Cedrus, Larix, and
multaneously on these internodes within a single Pseudolarix, whereas in mature Pinus only cata-
growing season. Shaw (1914) believed there was in phylls (also termed"scale leaves" or "bracts")appear
the genus an evolutionary tendency toward (Thomson, 1914). Dwarf shoots cease to grow api-
multinodal growth, most prevalent in the "seroti- cally at a length of ca. 250 aum(Mirov, 1967) and
nous" pines. Multinodalshoots are not a fixed char- terminate in a dormant bud. After defoliation, this
acterstate in any species; rather,they are a traitthatis bud may produce a second fascicle of secondary
either rare (occurringonly in strong leading shoots), leaves. In some cases, shoot proliferationoccurs as
common, or prevalent;this type of occurrencecould a response to damage (e.g., after damage to the long
support Shaw's view if it appears to be congruent shoot by herbivory).In most pines the initiation of
with the phylogeny of species inferred from fixed long shoots from shortshoots is not the normalmode
derived [Link] occurs primar- of branching,as it is in the otherthreegenera(Farjon,
ily fromthe cambiumlayers, formingbarkand wood, 1990a). Nevertheless,the distinction is only gradual,
although reiterationfrom dormantbuds may occur, with the condition in Pinus as the most advanced
especially in a few species that also coppice, e.g., and specialized of a derived state in the family.
Pinus leiophylla. A subapical meristem commonly The branching shoots (internodes) develop from
assumes dominance upon damage to or elimination single terminaland multiple subterminal,rarely lat-
of the leading shoot; this, rather than reiteration, eral, buds by elongation (the whole of a season's
causes irregularstem shape and branchingpatternin growth, vide Shaw, 1914) and are defined by a
older trees. In a few species, the plants remain leafless base and a terminal node of buds. As de-
small and shrub-like,primarilyby limitationof apical scribed above, several species in our region develop
dominance of the main (stem) shoot; however, all multinodal shoots, but the prevalent mode of shoot
species in our region, even P culminicola, remain development is uninodal. Shoot elongation com-
monopodial, and no truly multistemmedbranching monly concurs with elongation of secondary leaves
patterns(nor layering)has been observed in Mexican (see below), but in P maximartinezii,P nelsonii,
shrub-likepines. and, to a lesser extent, P cembroides and its allies,
growth of the leaves is postponed until the shoots
have reached full length. Shoot elongation, apart
SHOOTS
from apical dominance,is influencedby environmen-
The genus is characterizedby extreme shoot di- tal factors and is generally greater in species re-
morphism: branching shoots and dwarfed foliage stricted to a mesic environment.A few taxa in our
shoots. Shoot dimorphism occurs in all genera of region, most notably P elliottii var. densa, exhibit
Pinaceaeto a greateror lesser degree (Frankis,1989), arrestedapical growth of the apical meristem in the
with examples of strong dimorphism in Cedrus, seedling stage ("grassstage"). Instead,the hypocotyl
Larix, and Pseudolarix and weak dimorphism in is greatlyenlargedand the root system develops first,
Abies and Picea. In Pinus, the short foliage shoots while the foliage undergoes the normal develop-
are "extreme"in thatthey neitherelongate repeatedly mentaltransitions(see below). Once foliage and root
nor producemore than a first set of leaves normally, system are sufficient to sustain it, rapid growth of
as is the case in Cedrus, Larix, and Pseudolarix the stem lifts the young tree and its foliage above
(numberof leaves indefinite).They are thereforebest the normal zone of grass fires, where it is relatively
defined as dwarf shoots (brachyblasts)to distinguish safe and starts lateral branching.A comprehensive
them from the short shoots of the other [Link] study of shoot morphologywas publishedby Thom-
initial growth and formationof a definite numberof son (1914).
green leaves, they remain on the plant for the dura- Pubescence is rarein our region and of very short
tion of the leaves (2-20 years or more, but in our duration;several species commonly have glaucous
region not longer than8 years) and thereafterbecome to pruinose shoots. Shaw (1914) saw a correlation
deciduous with them, leaving in some cases a few between glaucousnessand aridity,but in the Mexican
basal remnantsof sheath(= bud) scales. In the other species thatgrow in the most aridclimatic conditions
genera, the short or spur shoots are persistent and it is absent or only weakly developed. Glaucousness
only the leaves fall. The secondary leaves are also is also stronglycorrelatedwith age and vigour, being
helically arrangedin the former three genera (in a much moreobvious on shoots of young, fast-growing

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Morphologyand Anatomy 15

trees and nearly absent on shoots of senescent trees primary leaves of species in subgen. Pinus is on
of most species. average somewhat higher: respectively, 1-5 and 2-
The cataphyll bases (pulvini) of most species are 4. There are 2-3 resin ducts in the primaryleaves;
persistentuntil after the leaves have fallen; these are the average number is slightly higher than in the
either short decurrent(subgen. Strobus, sect. Stro- cotyledons. Primaryleaves are usually producedfor
bus), short to intermediate(subgen. Strobus, sect. only the first season and only on the main stem in
Parryanae) or long decurrent (subgen. Pinus), in most species, but in sect. Parrvanae,and more rarely
which case they form ridges between the leaf bases in other sections, they continue to be producedfor
(Fig. IA-B). A furtherdifference between the two severalyears(up to 20 years in P maximartinezii)and
subgenera is the way the pulvini disintegrate: in also on lateral shoots. They may also be produced
subgen. Strobusthey erode slightly and become al- again on injury-responseshoots from dormantfasci-
most imperceptibly incorporated in the growing, cle buds and adventitiousshoots of matureplants in
smooth bark (but some exceptions occur in sect. several species in sect. Parranae. De Ferr6(1952)
Parryanae);in subgen. Pinus they become scaly and published a comprehensive study of the juvenile
break away from the underlying bark, which then stages of Pinaceae, including Pinus. from which she
attains a rough and scaly surface. inferredphylogeneticlinages and a classification.
The buds of long shoots, as definedhere, are really The secondary leaves begin to appeartoward the
"compound buds" (also known as "winter buds") end of the firstgrowing season or early in the second
consisting of the compacted,helically arrangedbuds in many species, but in other species (notably in
of dwarf shoots with secondary leaves, each subsect. Parryanae)not untilthe thirdor [Link]
tended by a bracteatecataphyll; the latter are more secondary leaves develop on dwarf shoots initially
or less imbricateand envelop the primordialleaved axillary to primaryleaves, later subtendedby cata-
shoot (Fig. IC). phylls, which are homologous with primaryleaves,
on (decurrent) pulvini. Thomson (1914) reported
transitionalstages between primaryleaves and cata-
LEAVES
phylls in several species. Cataphylls are alternate
The leaves of pines are of four types when ac- (helically arranged) non-chlorophyllous primary
counting for all developmental stages in the plant: leaves; their relative separationincreases with shoot
cotyledons, primary (juvenile) leaves, cataphylls elongation and secondary growth (girth). They are
(scale leaves), and secondary leaves (needles) (Fig. relatively small, subulatein most species, lanceolate
ID-G). Cotyledons appear epigeous in a whorl at in a few species, thin, soon scarious, and often
the apical end of the hypocotyl of seedlings and are recurved. In several species the margins are ciliate
linear,entire, and [Link] vary in number and hyaline, whereas the central part is thicker and
from 4 to 24 (P maximartinezii),the highest number pale brown to reddish brown or dark brown. Cata-
observed in any [Link] are (0-)2-3 resin ducts phylls are often stomatiferouson the adaxial side
in the cotyledons and either one (haploxyl) or two (epistomatic) (Florin, 1931). They are small, with
(diploxyl) vascular bundles. According to Masters erose-hyaline margins and early deciduous in most
(1904), some diploxyl pines in the maturestate are species of subgen. Strobus; they are larger, with
haploxyl in the juvenile stage (both cotyledons and ciliate margins and persisting longer in species of
primary leaves). A similar condition is noted by subgen. Pinus (Fig. IF).
Florin (1931) for the earliest secondaryleaves in the Secondary leaves (needles) are the permanently
extralimitalP halepensis Miller. Masters'sinference green, metabolically active leaves of the evergreen
about the evolution of diploxyl pines from haploxyl pines. They are borne on dwarf shoots axillary to
predecessors ("ontogeny reflects phylogeny") is a cataphylls on terminaland lateral shoots (stem and
conclusion we must nowadays treat with caution. branches), in clusters or fascicles of 1-8 leaves.
The cotyledons are succeeded by alternate(helically Single-leaved fascicles occur in P mnonophylla only,
or spirally arranged)primaryleaves on the growing resulting not from abortion of one of its two leaf
stem (and, in some taxa, the initial branchlets)of the primordiain the fascicle (Masters, 1904) but rather
seedling. Primaryleaves are linear-acicular,serrulate, from a single primordium(Gabilo & Mogensen,
amphistomatic, and usually glaucous. In subgen. 1973). Common numbersof leaves per fascicle are
Strobusthey have 1-3 lines of stomataon each side 2 (rare in our region and mostly confined to boreal
of a medial line abaxially and 1-7 lines on each side species), 3, 5, and occasionally 4 and 6-8. Numbers
of the medial line [Link] numberof lines in are variable to some extent in almost all species;

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16 Flora Neotropica

?: ?' X,' t/f

A . B

:
'I1i f j X

'.
H....':i' .
.-.:,:-'l ''j'
'. K '.
FIG. 1. Morphologyof foliage in Pinus. A, Ha. P. montezumae.B. P. [Link]. P [Link]. P. caribaea
var. bahamensis. Hb. P caribaea var. caribaea. He. P. attenuata. J. P strobus var. chiapensis). A. Branchiet typical of
Pinus subgen. Pinus. B. Branchlettypical of P subgen. Strobus.C. "Winterbud"complex, with terminaland subterminal
buds. D. Seedling with cotyledons. E. Seedling with primary leaves. F. Types of cataphylls. G. Secondary leaves in
fascicles. H. Fascicle sheaths, subgen. Pinus. I. Entire leaf margins.J. Remotely serrulateleaf margins. K. Serrulateleaf
margins.(Magnifications X 0.5.)

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Morphologyand Anatomy 17

very constantare the 5-leaved pines of sect. Strobus, deciduous. In most species of subsect. Cembroides,
but even in these taxa exceptions have been found as well as in P rzedowskii, the distal scales are
(Thomson, 1914). Distinctionscan be made between deciduousbut the proximalscales separateand recoil
low-numberand high-numbertaxa in the more vari- to form a basal rosette of scales before falling from
able group. Commonly or prevailinglyheteromerous the fascicle or sometimes with them. In most species
leaf fascicles occur in relativelyfew species, notably of subgen. Pinus (and in P nelsonii) the scales
in several belonging to subgen. Pinus and in some remainimbricate,and while the thinner,often ciliate
species in subsect. Cembroidesof subgen. Strobus. fringes and apices erode away, the remainingparts
Numbers above five are frequent in taxa of the form a tight, persistent, and sometimes resinous
informal "Montezumae" and "Pseudostrobus" sheath around the base of the leaf fascicle. Excep-
groups, most of which show considerable variation tional in this subgenus are P leiophylla and P lum-
indeed in these numbers. There is a north-south holtzii, which have early deciduous sheath scales as
trend or dine in leaf number within the group of in pines of sect. Strobus.
heteromeroustaxa in subgen. Pinus; low numbers The secondaryleaves, while still enveloped by the
are predominantin northerlyspecies, but this dine fascicle scales, are connateand togetherform a terete
is apparentonly on the continent as a whole and fascicle. The numberof leaves per fascicle therefore
much less evident on the subcontinent(Mexico and determinesthe transverseshape of the leaves, being
Central America). Clinal within this more limited a sector of a cylinder: circular (terete) in P mono-
geographical range is, e.g., P leiophylla. Supernu- phylla, semi-circular (plano-convex) in two-leaved
merary leaves, i.e., numbers in a fascicle that are fascicles, transverse-triangularin three-leaved fas-
above what is "normal" in the species, are also cicles, and (broadly) triangular in fascicles with
correlatedwith growth vigour (Thomson, 1914) and leaves in fours and higher (Fig. 2). The abaxial side
are more often found on leading shoots of young is always convex and the adaxial side is straightin
plants. [Link] lateralmarginsof the leaves
Secondary leaves are acicular, highly variable in are serrulatein all species of subgen. Pinus and are
length between and often within species, ranging serrulate,remotely serrulate,or entire in species of
from (2-)2.5 to 50 cm, rigid or lax, varyingin width subgen. Strobus(Fig. II-K).
between 0.5 and 2.5(-3) mm (up to 7 mm in one Stomataon secondaryleaves are arrangedin longi-
extralimitalspecies), and either epistomatic(stomata tudinal (grooved) lines running along the entire
on the adaxial side(s) only) or amphistomatic(sto- length of the leaf, as indicatedabove, either on the
mata on all sides). Hypostomatic secondary leaves adaxial face(s) only or on all faces (Fig. 2). In the
(stomata restrictedto the abaxial face), as in Abies, species of subgen. Strobusthe leaves are commonly
do not occur in Pinus except in P krempfii(as an epistomatic,[Link] abaxial
irregularcondition). Length of leaves is generally face has 1-10 lines of stomata (the terete leaf of P
correlated with both annual mean temperatureand monophyllahas 12-25 or more lines), the two adaxial
aridity,with the shortest leaves occurringin species faces 2-10 lines each. Species of subgen. Pinus have
with boreal or subalpinedistributionsand in species only amphistomaticleaves. The abaxial face of 3-
adapted to semi-arid [Link], there sided leaves has 2-17 lines, the two adaxial faces
is also a correlationbetween leaf width and seasonal have 2-12 lines each (2-sided or plano-convexleaves
aridity, allowing long-leaved species (e.g., P may have more lines, but truly 2-leaved pines do
coulteri, P devoniana, P engelmannii) to occur in not occur in our region). The position and external
areas or environmentswith long periods of intense structureof the stomata and their subsidiary cells
insolationand withoutrain. Long, slender,lax leaves has been the subject of elaborate studies, foremost
with relatively thin epi- and hypodermallayers (see of which is the classic work of Florin (1931), where
below) are confined mostly to warm temperate to a representativenumberof pine species was treated.
subtropicalmesic environments. The stomata are generally longitudinally oriented
The secondaryleaves are subtendedby a number and monocyclic, with 4-10(-14) subsidiary cells.
(usually 6-9) of helically arranged,thin, oblong to The subsidiary cells encircling the stomata form a
linear, chartaceous (papery), and partly hyaline moreor less raisedring termedthe "Florinring"(Fig.
scales, homologous with primary leaves and cata- 3C; see also Yoshie & Sakai, 1985). Investigating51
phylls, which are initially imbricate and are either species, Yoshie and Sakai (1985) observed six types
deciduousor persistent,forming a basal sheath (Fig. of Florinrings in the genus, of which five were found
IH). In the species of sect. Strobus they are early in Mexican pines. Types E and F are characteristicof

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18 Flora Neotropica

S A E

FIG.~~~~~~~~~'
2. Aaoyi rs eto nPnswt
fscnaylae aclrsse
alxl(-)addpoy E
L).A-D Psitonof esidctsexernl.A. .mnohyla.B. . nlsnii C [Link],bfi~mi. DP st~bu vt. hipenis
E - . Poiioeindctsreil
f .P uagni.F .atna .P otravt [Link] oiino
s~~~~~~~~~~~~[Link].L..tpiai.(Mgiiainx 3--40.)

FIG. 2. Anatomy in cross section of secondary leaves in Pinus with vascularsystem haploxyl (A-D) and diploxyl (E-
L). A-D. Position of resin ducts external.A. P monwphylla.B. P nelsonii. C. P strobiformis.D. P stn)bus var. (hiapensis.
E-G. Position of resin ducts medial. E. P durangensis. F. P uttenuata.G. P contorta var. murraiaina.H-J. Position of
resin ducts internal. H. P cubensis. I. PRcaribaea var. hondurensis.J. P praeterniissa. K, L. Position of resin ducts
septal. K. P jualiscana.L. P tropicalis). (Magnifications X 30-40.)

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Morphologyand Anatomy 19

.r;i
~'"~:----------":
'L.
18980).A-C. Outer view. A. Abaxial ";;-..:
~surface
'.. " showing ~T'l
:r------~' ', stomatal plugs C.
stomatal rows. B. Abaxial surface showing

~i~I B .:,,.
~ ':~:.:.}::,.
.
:' ..__._.~i~~ .:!~i~i:A- e ~ii
~ii::~i.i
-::
... ._'.. .. .F.-..
'.

Abaxial
surfae , showing stomatal apparatus
.: and weakl :
..:..:: .d F D.-F. ..
ring. Inner?view.D. A surfac
i~~~~~~~~~~~~~~~~~~~~'~" '~- ... '.

i..: ~ ::,:..... .........,, .-,,~ ....~.~~~~~~~~~~~~~~~~~~

F ,.,.::,
~
i~~~~~~~,.l.~[1Jl~'--"~?'M~
t l'~._..~.~.i:A -
~~~~~~" .,

FIG;. 3. Scanning electronmicrographs

(SEM) of cuticles ofPinus
montezumae Little~~~~~~~~~~~~~~~~~~~~~
(A-C, 10170:
D-F,
Pringle
1880. -C utr ie. .Abxil ufae,shwigstmaalrws B Aaxa srfce hoin somtl lus.C
Abaxial~~
surfae,
shoi
st dvlpdFoi
in ngoaa gpaatsadwal
- F n e r v e . . Aaxaurae
osadlnitui
show~~~~ingsoaa na oienaion.E
daiai
s . tmtlsrfe
edge'
ofsrrl te law ithoaafe

showing stomatal rows and longitudinalstomatalorientation.E. Adaxial surface edge of serrulateleaf with stomata free
zone. F. Adaxial surface, showing stomatal apparatusand six subsidiarycells. (Scale bars = 50 ,um.)

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20 Flora Neotropica

parenchyma _ m~ esophyll
schlerenchyma

bundle "xylem : 1000,,

resin canal
/endo-rd ~ or duct
de r .

guard

esophyllhloem sculr resin

canal
schlerenchymo .iil~r
? duct

epidermis scherenchymo
pore

mesophyll

epidermis schlerenchymo

FIG. 4. Cross section of a two-needled pine leaf (inset) and cellular detail of a section from the epidermis to one of two
vascular bundles.(FromT. E. Weier et al., eds. 1982. Botany: an introductionto plant biology. Ed. 6. John Wiley, New
York.)

taxa of subgen. Strobus and do not occur in other in cross section under magnification of thin slices
pines; all other types are found only in subgen. (microtome-cut and usually stained), has been the
Pinus. Types E and F are characterizedby the ring subject of numerous studies. General reviews are
forming an abrupt, high ridge and a rectangular given by Coulter and Rose (1886), Doi and Mori-
stomatalopening. The other types have more-gradu- kawa (1929), Harlow (1931), Sutherland(1934), Jah-
ally rising rings, grading to virtual absence, and rig (1962), and Kwei and Lee (1963), and summa-
irregularto circular stomatal openings. Distribution rized accounts are given in Shaw (1914), Martinez
of epicuticular wax, formed in the stomatal zones (1948), and Farjon(1984), among others. The trans-
(lines), appears to be correlatedwith the Florin ring verse section of the leaf can be divided into three
types, especially at subgeneric level. Type C, in distinct structuraland functionalparts:(a) the dermal
which the Florin ring overarches the stomatal pore tissues, including the stomata; (b) the green tissue
resultingin a "volcanic"shape, with a small stomatal (mesophyll), containing the resin ducts; and (c)
opening, may be uniqueto the genus Pinus (Yoshie & the stelar tissues, containing the vascular system
Sakai, 1985). Although observed in other species, it (Fig. 4).
is prevalent in subsect. Oocarpae. Generally, more The dermal tissues consist of the cuticula, epider-
wax development was found in stomatal types with mis, and hypodermis, in which the stomata are em-
wider openings, suggesting a correlation with gas bedded, providing openings to the underlying green
diffusion and (alternative) strategies in relation to tissue. The cuticula is a layer of cutin, a highly
waterstress by decreasingwaterloss due to transpira- polymerized mixture of lipids and waxes (Mirov,
tion. 1967), covering the outer surface of the epidermal
The anatomyof the secondaryleaves, as observed cells. It gives most leaves a more or less glossy

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Morphologyand Anatomy 21

surface and appears to exfoliate in a few species, in the mesophyll and the outwardopening is flanked
e.g., P monophylla. The epidermis consists of a by two guardcells, which are sunkenbelow the level
single layer of thick-walledcells. The shape of these of the [Link] guard cells are moveable and
cells is fairly constant but varies among individual control the stomatalopening (Fig. 4).
trees and/or species. Jahrig distinguished four The mesophyllof pines consists of largecells with
classes: square, transverseoblong (rectangular),an thin, undulatedto invaginatedwalls, filling the entire
intermediatebetween the two former, and oblong space between the hypodermis and the endodermis
(either rectangularor elliptic). The hypodermis is and enveloping the resin ducts. They contain organ-
considerably more variable as it consists of I to elles, among which are plastids (chloroplasts),mito-
several rows of sclerenchym cells, often unequally chondria, ribosomes, and the nucleus. They are the
distributed, with concentrations in the lateral and cells from which increasingly taxonomically infor-
axial marginsof the leaf (Fig. 2). In some species, mative molecular data, especially chloroplast-DNA
notably P douglasiana, it forms extensions penetrat- (cpDNA) sequences, are obtained. The resin ducts
ing through the mesophyll and connecting with the are narrowtubes usually runningthe entire length of
endodermis,often enveloping resin ducts. The cells the leaf. They are not connected to cambium or
are also more or less thick-walled but are often xylem resin ducts. They have walls of thick-walled,
variable in this respect as well as in shape within a sclerenchymatouscells of similar appearance and
single leaf. Generally, the thickest cell walls are structureas the hypodermalcells and give a similar
found in the inner rows of multi-layeredhypoderms. chemical reactionto the [Link] cells
Shaw (1914) classified the hypodermalcells as uni- may be laid down in single (most common) or
form (either thin or thick), biform (both thin and multiple rows; in the latter case they decrease in
thick), or multiform(with a gradualtransitiontoward thickness and size inwardly (Fig. 4). Their number
the thicker-walledcells). In some species-e.g., P and position relative to the dermal walls of the leaf
ayacahuite, P strobiformis,and P strobus-epider- are highly variable among species and can be used
mis and hypodermisare of similar single-cell thick- as [Link] numberof ducts within
ness and similar cell shape and size, but their differ- species is generally more variablethantheirposition,
ent nature becomes quite apparent in the staining but some measure of polymorphism is present in
process, whereby under the action of hydrochloric almost all taxa for both [Link] numberof
acid the hypodermis is sharply differentiatedfrom resin ducts in a leaf ranges from 1-12 or more in
the epidermis:the formerbecomes differentlystained Latin American pines, and is even higher in some
(red when safranin is used, against blue or green). otherspecies. Since Engelmann(1880a), the position
The width expressed in uni- or multicellular rows of the resin ducts has been used as a characterto
and the polymorphismof the hypodermalcells have classify the pines; since Coulter and Rose (1886),
been used as diagnostic characters(Harlow, 1931; these positions have been termed external, medial,
Jihrig, 1962). We have not followed this distinction internal,and septal. External(as "peripheral"in En-
for cell thickness in the characters described for gelmann, 1880a) ducts are situatedagainst the hypo-
the species, but have distinguishedbetween uniform dermonly. In some cases the hypodermalcells merge
(homomorph)and polymorphhypodermsin termsof with those surroundingthe resin duct and it is then
their overall structure. Next in complexity to the difficultto distinguishtheirorigin. Medial(parenchy-
unicellularhypodermisof the above-mentionedspe- matous)ducts are free in the mesophyll, touching no
cies is the uniform but double-layeredhypodermis [Link] touch the endodermis
of, e.g., P greggii and P patula. A simple form of only. Septal ducts, additionallyrecognized by Shaw
polymorphism,already mentioned, is the concentra- (1914), touch both hypodermand endoderm, some-
tion of hypodermaltissue along the marginalareas, times interconnectedthroughintrusionsof the hypo-
while the remainderof the hypodermisremains uni- derm, forminga septum. Most species are character-
form with two rows of cells. All other, usually ized by a single position (Fig. 2). In a comprehensive
thicker, hypodermalstructures,including intrusions study of nearlyall species in the genus, Jahrig(1962)
into the mesophyll, are classified as multi-layered never encountereda combinationof more than two
(polymorph)in this work. positions, but considered medial and internalducts
The stomatapass inwardthroughthe dermalcells as occurringwith equal frequencyin severalMexican
to the mesophyllzone and are essentially of the same pines of subgen. Pinus. In several species there is a
structurethroughoutthe family (Florin, 1931; Farjon, combinationin which one positionprevailsbut where
1990a). Commonly there is a substomatalchamber one or a few often smaller ducts occupy one of

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22 Flora Neotropica

the other positions. In many species with variable which produces the phellem (-- periderm) outward
numbers of resin ducts, certain ducts, with a fixed and a thin layer of phelloderm inward. Commonly
position and slightly larger,are always present. We the barkof pines is divided into threezones: phloem,
consider these primaryducts and the others subsid- cortex, and epidermis (Mirov, 1967; Farjon, 1984);
iary ducts. but we have followed the terminology proposed by
The stele of the secondary leaf is a tubularstruc- Junnika (1994) here, as it is more accurate. The
ture, usually terete, but often more or less triangular outer bark is thin on young trees and branches,
or oval in cross section, being correlated to the but increases in thickness by the accumulation of
transverse shape of the leaves. Its wall, the endo- peridermlayers added by the phellogen. Simultane-
dermis, consists of a single row of relatively thin or ously, the outermostlayers of peridermexfoliate, so
partly thick-walled, globose to oval cells. The outer thatthe ultimatethicknessof the outerbarkis defined
cell walls in several species are thickened, and this by the age of the stem and net result of these two
is often used as a diagnostic character,as it appears processes. In some species exfoliation startsearly, in
mostly constant. Only a very few species are poly- others late.
morphic in this respect, e.g., P leiophylla var. chi- Bark patternsin pines result mainly from fissuring
huahuana. Inside this wall is the vascular strand(s) and scaling (Fig. 5). Fissuring is caused by the
or bundle(s) embedded in a matrix of transfusion expansion growth of stem and (larger)branchesand
tissue. Often there are strands of sclerenchym cells results primarilyin lengthwise cracking into fissures
which intermittentlysurroundand often divide the separated by ridges. In most species these fissures
vascular bundles (Fig. 2). The vascular bundles are are irregularlyconnected. Incomplete shedding of
either single or double in pines. This distinctiongave scales results in horizontal cracks that widen by
rise to Koehne's sections Haploxvlon and Diploxy- further wood expansion, forming tessellated or
lon. The double bundle is usually obvious, even polygonal [Link] longitudinalfissuresare deep
when (partly) connate, in at least the majority of or shallow, with "shallow" defined as a depth less
leaves of species of subgen. Pinus. The vascular than one-half the total thickness of the bark. They
bundle consists of two connate strands, an adaxial can be V-shaped, irregular,or [Link]
xylem strand,and an abaxial phloem strand(Fig. 4). ridges are usually flattened,sometimes rounded,and
In diploxyl pines they are either distinctly separate often reticulate. Flakes are usually irregular. On
or connate on the xylem side; in a few instances young trees and the upper stem and branches they
they may be entirely connate and seemingly form a are commonly papery, on the lower trunk they are
single bundle,as in all haploxyl pines. An interesting often thick, harder, and plated or shaggy. Colour
feature has been found in the haploxyl species P patternsresult mostly from scaling, whereby exfoli-
maximartinezii,which has a strandor wall of scleren- ated flakes reveal younger, less weatheredsurfaces,
chyma cells separating the phloem of the single or from fissuring and cracking, which may expose
vascular bundle and occurring in the majority of varicoloured inner bark. Illustrationsof patterns of
leaves observed (Rzedowski, 1964). bark have been publishedby Mirov (1967), Howard
(1971), Farjon (1984), and Perry (1991); the latter
author provides small photographsof boles for all
BARK Mexican taxa. Although some generalizations can
The bark of pines, as commonly mentioned in certainlybe made-such as the long-retainedsmooth
descriptionsof species, is the outerbark(rhytidome), bark of Pinus sect. Strobus, in which the formation
which often forms characteristicbark patternsespe- of periderm is delayed, or the often tessellated or
cially on the (lower) bole of maturetrees. The bark polygonally patternedbark with large fissures and
has a complex structureand consists of a variety of ridges in subsect. Ponderosae-identifying species
tissues, separatedinto two zones outside the vascular on the basis of bark charactersshould be practiced
cambium: the inner bark (phloem) and the outer cautiously. Some species (e.g., P nielsonii in our
bark (rhytidome). From the vascularcambium layer region) have very characteristic barks that make
outward,the barkconsists of the secondaryphloem, identificationat species level possible. In most taxa,
the primarytissues still existing outside the second- however,thereis considerablevariation,and similari-
ary phloem, the periderm, and dead tissue outside ties between taxonomicallywidely separatedspecies
the [Link] vascularcambiumproduceswood are common. The anatomyof bark of 10 species of
(xylem) on the inside and phloem on the [Link] Pinus from the southeasternUnited States (all sub-
first peridermcontains a cambium layer (phellogen) gen. Pinus, one of them, P elliottii var. densa, in-

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Morphologyand Anatomy 23

~
!~;:j~~~~~~~~~~~~~~~~~~~~~~~~~~~a ~
'::
~ ::i;;i;? '
'.'?.?. ' : :: ~
. ~ ~
,:.,~~~~~~~~~~
~~~~~~~~~.-~ ~..:..:..... X.-:.::'.?
~.:"L"....!::-."
...
...
:~~...
..
... ..... ....
...'.'..'...~....:..:
;:... .......-...:

~~~~~~~~~~~~~~~~~~~~~~...
.a?'.'....-.'.. ... ' ':.-..:
!:..:.".:"..'. : ' : :,
. .........~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
.: .";';.~'~.':.
... .. ... .. .
' i'!Si~~~iii~i~i::;:::::.;..
: . . . . . . . . . '"....
. ..... . 'N::.; , . ' ' :..,.
~'?~:i:.::.
.....,;,..!~;;~, ,~,.
,,~,,..,:,:.;.. : : :::::::::::::::::::::::::
:. : : : : :
:Y":.:....:...:>..... . . . ~:E;"

:i'~*.~~ ~ ~ ~~ ~ ~ ~ ~ ~ ~~~~~~~~~
:./':.~ ~ ~~~~~~~~~~~
" ..".,,':'...'.:
..:... ................
;i~~i$
;'..?** '
:"i~i;i"~::'~'":::;'"..'..
.:::.:::::: : . : i .......
:-.:.
i',;i;',,ii:::',:i/,i::;:;i~~~~i~i,::,:,',',i..,.!
..:.:i;;.....
:".
? ::.:i.
!:::" . '.%: ~ ~ .' ill
[Link]~5Ii3:

:i~~~~~~~~~~?:L?~~~~~~~.?
X.:':..: . .

- ?
A~~~~~~~~~~~--
a---- , ~~
?-? ~ ~ i.~~~~~~~~~~~~~~~
.::i:i~

i~. _ ~ . ;~<.....~~~~~~~~~~~~~~~:!,......
7~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~b:
~ ..... ...
.
.... ~ ~ tj ' : :::::::::::::::::::::::::::::~~~~~~~~~~~~~~~~~~~~~~i
i;?;;;i::i::i!
~. ~'"' ,.----~-:;??- i~~~~~~~~~~~~~~~~~~~i,;?:E~~~i::
-"~..."~-,,..';~~~_~...
~ tlW~.~'
~' ":'~~~
~~~~~~;:.

? '
C.
"gi.." :;:::utlos?. b.s. 'D:.::: :..oiina:'Z..:i::
[Link]!::. n.........:'. F.::':::':"...':.:P::.::: ::s.:i. H::. P
nwvi::.;fiez'/?.1.'
'..:;,::~'. .......~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~1
:?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
..!....,
,x~~~~~~~~~~~~~~ ::

': .':''.. :'::':;""::R:

E n~~~~~~~~~~~~~~~~~~~~~:i'
::ii....
....:,:-
:~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~?
"..!.,,i!'
i:,::
;'.- '.' :

FIG.
5. Trunks ine
pten.
A.:
PiQf~iittusih
of Mex~~~icnadCtrAica ilrntbr leiophylt. B. hart
wegi. . Ppeudstrbu. [Link].E.P. axiini. . Pocara. . .. zedwsii.H. ..maxntatiezi. I P

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24 Flora Neotropica

cluded in our region) has been describedby Howard small (1-2 mm) and subpeltate (free only at the
(1971). lower end) and the uppermarginis entire. In species
of subgen. Pinus, the microsporophyllis larger (ca.
2-3 mm diam.), peltate, with an erose-denticulate
REPRODUCTIVEORGANS
[Link] outer surface in the formersubge-
The genus Pinus is monoecious. Male and female nus is smooth and in the [Link]
reproductiveorgans (strobili) are separatefrom each differences are true for all species studied here, but
other, arising from separate axillary buds, though perhapsnot for the entire genus. Furtherdifferences
commonly occurringon the same branchesand occa- are found in the size of the entire pollen cone, which
sionally on the same long shoot. In older literature is smaller in subgen. [Link] species of subsect.
(e.g., Shaw, 1914) and in many papers by foresters Cembroides,pollen cones are not more than 10 mm
on the subject, angiosperm terminology such as long and in P rzedowskii only 5 mm; they also
"flowers,""anthers,""catkins,""staminate,""pistil- extend up to more than one-half of the length of the
late," "flowering,"etc., are commonly used in de- shoot, forming spike-like clusters. In most species
scriptions of reproductiveorgans and phenology of of subgen. Pinus in our region, the pollen cones
conifers. This is to be avoided, as it not only leads form short, "capitate"clusters, but there are some
to confusion but, more importantly,is misleading exceptions and this trait is variable in many species.
since both structure and reproductive biology are The microsporophyllsof some species are initially
very different from those in angiosperms. red or purplish,of others yellowish green or yellow,
and most turn yellowish brown to brown. There is
POLLENCONES again uncertaintyaboutthe fixity of these differences
and, as has been demonstratedfor ovuliferous cones
Pollen cones arise from axillary buds in the proxi- in Pinaceae,the reddishcolorationmay be influenced
mal part of the compound or aggregatebud through by environmentalfactors. Due to the minimal inter-
a modification of dwarf shoot primordia (Mirov, specific differences, or the lack of even these, pollen
1967); their subtendingbractsare homologous to the cones rarely receive attentionin descriptions.
scales of fascicle sheaths. In most species of subgen.
Pinus, buds of incipient pollen cones are slightly
SEED CONES
larger and paler than those of sterile dwarf shoots
above them; in most species of subgen. Strobus The seed cones (ovuliferous cones) are the most
they are inconspicuous(Shaw, 1914). They are also characteristicstructureof the coniferous plant. Much
distributedover a greater length of the shoot in the has been writtenabout their ontogeny, structure,and
lattersubgenus, especially so in the Mexican species homologies. We shall not attemptto fully summarize
P maximartineziiand P rzedowskii. that literature,but we would like to recall the fact
The pollen cone (microsporangiatestrobilus, Fig. that the studies of fossil primitiveconifers by Florin
6A-B) within the Pinaceae is much less diversified (1938-1945, 1951) established beyond doubt that
than the ovuliferous or seed cone (Farjon, 1990a). the seed cone of Pinaceae is a compound structure
It is a relatively small, weak and soft, ephemeral, homologous to a long or branchingshoot. Ontoge-
compoundstructure,consisting of an axis with many netic observations confirm this [Link]-
helically arrangedmicrosporophylls,usually ovoid- oine-Sebastian(1975) found stomataon both apoph-
oblong to cylindrical and up to a few centimetres ysis and umbo of all seven species (in both subgenera
long. Like the leaf fascicle, the pollen cone itself is of pines) studied. This, and a reduction in vascular
surrounded at the base by a number of helically strands from seed scale proper to apophysis, gave
arranged scales, but these are usually broader and her reason to designate the formeras "axial"and the
not [Link] axis is thin and flexible. latteras "foliar,"i.e., homologous to a leaf. Stomata,
Dependent on their length at maturity the pollen however, are not restricted to leaves of vascular
cones of several species become subpendulousafter plants or their homologues. Comprehensivedescrip-
shedding the pollen. The microsporophylls (Fig. tions of cone morphologyare given by Shaw (1914)
6Aa-b, Ba-b) are borne on minute stalks and termi- and Klaus (1980). The cones originate from "winter
nate in a (sub-)peltate structurebearing two pollen buds." The peduncle, varying in length from a few
sacs (microsporangia) on the lower adaxial side, millimetres to 6-7 cm in P nelsonii and up to 12
which dehisce by means of a longitudinal slit. In cm in P. lambertiana,is an axillary shoot positioned
species of subgen. Strobus, the microsporophyllis subterminally,or sometimes laterally,on a branching

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Morphologyand Anatomy 25

2 .. .
I'

'
.. ....
'" i r:

adnate,~~~~~~~~
'tfctv 'igs i. ?hni. ..~k,ruri. I~ i K . :.....h rtcuat, fetv ." .' .-:.:"'ic?lrtui
ig.....
b = seeds. N,~ nlonc~l~lt!lkr
seeds (b)with articulae, vestigial ings remainin on the scale(a)."('.., .

~~crrtunucr~cr.M,I! strr,hif~rrllis seeds with adnate, ineffective ~bortive)'

wings; a = seed scale with seed cavity tone seed

AJ :L. M.b. -. N- b:

FIG. 6. Morphologyof pollen cones, seed scales, and seeds in Pinus. A, B. Pollen cones microsporophylltype. a =
abaxial view. b = lateralview. A. P (subgen. Strn)bus) monoplh U11(.
B. P (subgen. Pinn.s) maximinoi.C. Seed scale type
of P. (subgen. Strobus. sect. Strobs) strn)ifonms. a = abaxial view, b = adaxial view, I = apophysis. 2 = terminal
umbo. D. Seed scale type of P (subgen. Pintos)dlevonni(a. a = abaxial view, b = adaxial view. c = lateral view. I =
apophysis. 2 = dorsal umbo. 3 = transversekeel. E-H. Increaseddevelopment of apophysis and umbo in seed scales
of P. subgen. Pinn.s.E. [Link]-a. F. P cnribaea var. caribiena.G, H. P. [Link]. puk-ensis. 1, J. Seeds with
adnate,effective wings. I. P cubensis. J. P lamberitun. K, L. Seeds with articulate,effective wings.. K. coultei. L.
P attenuila. M. P strohin)bifonns seeds with adnate, ineffective wings; a = seed scale with seed cavity (one seed abortive).
b = seeds. N. P mnoplylla seeds (b) with articulate,vestigial wings remainingon the scale (a). (Magnifications x.5.

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26 Flora Neotropica

shoot. It is initially erect but commonly becomes the growing scales. Growth then stops until the
recurvedor pendulouswhile the cone grows to matu- following growing season, often six months or more
[Link] may be solitarybut more often are placed later. Pinus nelsonii is unique in the genus in not
in pseudo-whorls,and in some species many cones having a definite resting period (Shaw, 1905a, 1909,
appear each growing season. There is considerable 1914); as a consequence, its umbo is rather ill-
variation in the persistence of the cones, especially defined and poorly [Link] conelet at this
within subgen. Pinus, from deciduous shortly after stage (referred to in the descriptions as "immature
seed release to almost indefinitelypersistent(e.g., P cone") exposes the later umbos. The umbo is often
nuricata, P attenuata, P radiata, P greggii). Shaw armed with a spine (crista) or short prickle (mucro)
(1914) assumed an evolutionary trend in the genus in species of subgen. Pinus; it is obtuse (mucro
leading from symmetrical,readily opening, and de- absent) in sect. Strobus. In a few species of sect.
ciduous cones to asymmetrical,serotinous, and per- Parrvanae (e.g., P cembroides and its allies), the
sistent cones (Fig. 7). Cones of pines in subgen. mucro is placed eccentrically,i.e., not on the line of
Strobus are normally deciduous, with the point of the transversekeel (Klaus, 1980). The outer edge of
detachmentat or near the base of the peduncle. In the scale later forms the transverse keel, which is
several species of subgen. Pittus there is a partial continuedon the apophysis(Fig. 6Da-3). Pines show
disarticulationof the cone, whereby the proximal two phases of cone expansion, in the second phase
scales remain with the persistent peduncle; in P the apophysis (the part of the scale exposed when
praetermissa these scales even fall [Link] the cone is still closed) is formed and the seed scale
P nelsonii there is articulationat the apex of the bearing the seeds grows. In this phase, the umbo
peduncle. In this and some other species (e.g., P appears to be terminal in species of sect. Strobus
cembroides and its allies, P pinceana), there is a and dorsal in other pines occurring in our region,
(very) thin attachmentof the scale to a (weak) rachis, belonging to both subgenera (Fig. 6Ca-2, Da-2).
which often leads to early disintegrationbut usually Biennial cone maturationvaries among species-it
not before the cone has fallen. The peduncle is takes mostly from about 17 to 24-25 months-but
covered with helically arrangedcataphylls;normally it is fairly constant within most species. In our
there is no development of vegetative dwarf shoots, region, P oocarpa may have the shortest maturation
though they are known in P nelsonii (M. P. Frankis, period. In a few species (P leiophylla in the Flora),
pers. comm.). The ovuliferous scales are axillary to growth of the cone progresses only slowly in the
small cataphylls, and helically arranged,forming a second growing season, and a thirdseason is required
determinate cone. Shaw (1914) has explained the to reach maturity(triennialcone development).This
phyllotaxis of pine cones. The two subgeneradiffer results in a distinct band between the umbo (first
primarilyin there being a relatively low numberof year of growth) and the lower part of the apophysis
scales per unit length of axis in Strobusand a higher (third year of growth). This concentric band was
number in Pinus. Due to the varying shapes of the termed the "vallum"by Klaus (1980).
cones, their phyllotaxis is variouslycomplicatedand The apophysis is a conspicuous subterminalstruc-
deviates from the theoreticalarrangementin spirals ture of the cone scale, which has attained a high
on a cylinder. Unlike in other genera (e.g., Abies and degree of diversification in the genus separating it
Larix), no proliferation of a vegetative shoot has from other genera in the family (Shaw, 1914; Klaus,
been observed in the genus. The seed scale, with its 1980; Frankis,1989; Farjon,1990a);consequentlyits
subtending bract, is homologous to the vegetative charactershave frequentlybeen used for taxonomic
dwarf shoot (and to the fertile dwarf shoot in several [Link] texture of the apophysis is mark-
fossil conifers). edly different from the anteriorpart of the scale. It
At pollination, the seed scale (ovuliferous scale) is also more extensive on the abaxial side than it is
consists of a flattened structure in the shape of a on the adaxial side, where it consists merely of the
short, apiculate to acuminate spoon. The seed scale outer rim of the scale (Fig. 6D). It is simplest in
in pines is a highly specialized organ, resultingfrom species of sect. Strobus,consisting of the (slightly)
the ontogeny of the cone and the periodical growth thickenedpartof the scale which was exposed in the
phases on the apophysis as concentric differentia- second growing season, terminated by the obtuse
tions. After pollinationthe seed cone (megasporangi- and commonly resinous umbo. It has a terminal
ate strobilus, Fig. 8) closes its scales by expansion. orientationand growth pattern,although outside our
Subtending bracts are still visible at the beginning region some transitionalconditions toward the fol-
of this stage but soon become submerged between lowing type exist in subsect. Cembrae. Its abaxial

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Morphology and Anatomy 27

,j B

A C

FIG. 7. Morphology of seed cone types in Pinus. A-E. Seed dispersal anemochorous.A. P. strobus var. chiapensis.
Pendulouscone type with quickly parting seed scales. B, C. Subsessile cone types with slowly partingseed scales. B. P.
montezumaevar. gordoniana. C. P durangensis. D. P pseudostrobusvar. apulcensis. Semi-persistentand subserotinous
cone type with armored,slowly parting seed scales. E. P attenuata. Persistent and serotinous cone type with armored
and closed seed scales. F. P quadrifolia. Seed dispersal zoochorous. Deciduous cone type with "wingless"seeds. (Magni-
fications x 0.5.)

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28 Flora Neotropica

ovule

"~._ /~. ~-.-" -- '-; ?;" ...polk..

ovufrus megasporo)mcte lh i sca

ni
pltiropylt E

a - wrs....me -~ '
vuitcm . y't="'.
.rs"'r .t:?':" .i I plmiunse

ov ueinl ,
st~~~~~~~robsius~~lom
Ochp ok t ibe_
u
ion
uvn#v->D

. uL R . vegeiaiive mmdci sinu ci

emuinMion

SUSri6flSCf
--^~ [Link]'
- -- - -- c e A

"Staminate" strobilus, pollen, development of the pollen tube; embryo and germination of seed. For further explanation
see text. (From T. E. Weier et al.. eds. 1982. Botany: an introductionto plant biology. Ed. 6. John Wiley, New York.)

surface is often irregularor longitudinally rugose. posed to sun and rain are most developed (see
In some taxa, e.g., P avyacahuitevar. veitchii, the above). Almost all apophyses with dorsal develop-
apophysis is distally contracted and elongated or ment are transversely keeled as a result of pre-
recurved/reflexed. expansive connection and partial overlapping; mu-
The apophysis in other pines of both subgenerais cro, umbo, and transversekeel are (retained)features
more complex and its orientationis dissimilar: bor- of an earlier phase in the cone ontogeny. Klaus
dered in the closed cone on all sides by the margins (1980) distinguishes three ontogenetic phases, each
of other apophyses, its growth is a dorsal extension reflected by its own morphological features of the
or thickening (Shaw, 1914). The outline is rhombic, mature seed cone: I) strobilus at pollination, with
or irregularlypentagonal,or in other cases it has a free ovuliferous scales, transverse keel, and crista
roundedupper marginand an angularlower margin. (mucro);2) firstyear's cone, with expandedovulifer-
It can be nearly flat, conical, or gibbous, or form aous scales with umbo, transverse keel, and mucro;
conspicuous and often curved protuberancein some 3) second or third year's cone, with lignified, en-
cases (Fig. 6G-H). Extreme apophysaldevelopment larged seed scales with keeled apophysis. umbo,
occurs, e.g., in P attenuata, P coulteri, P muricata,transversekeel, and mucro. From the centre to the
and P pseudostrobus var. apulcensis. On such pro- periphery,therefore,one encountersconcentric zones
nounced apophyses the umbo may be extremely of youngerphases. A secondaryontogenetic differen-
developed, too, which merely indicates that this ex- tiation is responsiblefor differences in size and shape
cessive growth started in the earlier stage of cone of mucro, umbo, and apophysis between proximal,
development,interruptedby the restingseason. There central, and apical scales. Whereas such differences
is invariablya longitudinaland one-sided asymmetry may be typical of certain species-e.g., the strongly
in the development of abnormallyshaped apophyses recurved proximal scales of P. avacahuite or the
on a cone, whereby those on the section most ex- straight proximal scales of P strobus var. chia-

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Morphologyand Anatomy29

pensis-comparisons between species are to be made Comparedto other genera in the family, the seeds
on scales of the same zone. In general,differentiation of Pinus are much more variable in size; this is in
decreases toward the apex of the cone. part correlatedto variations in cone size, but Pinrs
The anterior part of the seed scales of a mature cembroides, P culninicola, P m7onophylla, P quadri-
cone ("seed scales," in the species descriptions)con- folia, and to a lesser extent P nelsonii and P
sists of two different structuraltissues and also the pinceanCa have proportionallyvery large seeds. The
thin tissue that forms the seed wings. The abaxial largest seed in the genus is found in P maximarti-
tissue has much greater lengthwise swelling and nezii, which reaches 28 X 12 X 10 mm, in an
shrinkingpropertiesassociatedwith moisturecontent exceptionally large cone. The smallest seeds are ca.
thandoes the adaxialtissue, which consists primarily 4 mm long. Shape is variable,too, but most are (ob-)
of a numberof longitudinalxylem strands(Harlow ovoid and slightly flattened(Fig. 61-N). In the pin-
et al., 1964). This differential provides the opening yon pines (P cembroi(iesand allies) thereis a marked
mechanism of pine cones through parting of the difference in the thickness of the seed coat among
scales by [Link] are considerable differ- the several species. Thick seed coats are a protec-
ences between species, both in the conditions re- tion against predation by mammals and select for
quired for seed scale parting and in the extent to certainbirdsthat play a role in [Link]
which they can [Link] serotinouscones of some in some other genera, there are no resin vesicles in
species usually open only after excessive heating by the seed coat. The outer surface is usually smooth
tire ("closed-cone"pines), whereas in other species and dark grey-brown to blackish, but light brown
the scales part only after considerable time (years) seeds, with or without dark spots, occur in several
but under more normalclimatic conditions of insol- species.
ation and [Link] serotiny is correlated The seed wing is not derived from tissue of or
with persistence. Shaw (1924) further observed a surroundingthe seed coat, as Shaw (1914) under-
correlation with obliqueness ("asymmetry")of the stood it, but from tissue on the adaxial face of the
cone resulting from a lesser or greater inequalityin seed scale. Seed wings are, unlike those of other
the growth of both the seed scale proper and the genera in the family, highly diversified within the
posterior tissues forming the apophysis and the genus Pinus (Frankis, 1989; Farjon, 1990a). This
umbo. The condition is indeed unique in the family diversificationshows adaptationsto different strate-
if (slight) curvatureof the cone without substantial gies of seed dispersal and is evolutionaryand taxo-
differential growth of scales is not considered. nomically significant. Basically there are two types,
Whereasthe symmetrical,cylindriccones of subsect. related to wing attachment:articulate and adnate
Strobi are pendulous on straightor recurvedpedun- (Fig. 6K, J). Even in species with vestigial or "ab-
cles, the cones of many of the closed-cone species sent" wings, remnantsof basal wing tissue are pres-
are reflexeddown the branchon very shortpeduncles ent on the seed scale, on the seed, or on both. The
and become effectively sessile at maturityby thick- first author to observe this was Engelmann(1862),
ening of the branch. Of the two conditions, the in a footnote about P cembroides, but his remarks
formeris consideredthe plesiomorphicstate by most were not acknowledged,nor the fact observedagain,
authors(e.g., Shaw, 1914; Klaus, 1980). The greater by subsequentdescribersof pine morphology(Mas-
development of the apophysis on the side of the ters, 1904; Shaw, 1914; Martinez, 1948; Mirov,
cone persistentlyexposed to sun and weathercan be 1967), who all made a primarydistinction between
interpretedas an adaptationto protectthe developing wingless seeds and winged seeds instead. However,
seeds (Shaw, 1914), thereby enabling the species to Small (1903) and Frankis(1989) correctly observed
occupy environmentswith more extreme daily and the situation. In the articulate seed, the wing of a
seasonal temperaturefluctuations. full-grownseed is loosely attachedto the seed, which
is held either by two narrow,claw-like extensions of
the wing or by these and an extension covering other
SEEDS
partsof the seed on the adaxial side (Fig. 6K, I). It
The family Pinaceae is characterizedby two anat- has been suggested that the outer layer of the integu-
ropous ovules on the proximal end of the adaxial ment plays a part in the formation of this latter
side of each ovuliferous scale (Fig. 8). The seed section of the seed wing membrane(Schnarf, 1933).
containsan embryo,embeddedin the female gameto- These appendages hold to the seed in all species
phyte, which in turn is surroundedby the nucellus with well-developedwings thatdetachfrom the scale
and the seed coat or testa (Fig. 8). during the drying of the cone and the partingof its

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30 Flora Neotropica

seed scales. Upon dispersal they are normally still and theircones are situatedprimarilyon the extremi-
attached(and therebyeffective) but easily separable. ties of the higher (main) branches. Articulate but
Adnate wings remaininseparablefrom the seed upon effective seed wings may be of adaptive value in
dispersal and cover the seed partially,usually more dispersal after fire-they help carry the seed some
so than articulate wings (Fig. 6I-J). All effective distance-but with an (easy) articulation of the
seed wings are thin and membranous,though a few highly flammable wing, chances of the seed being
species (e.g., P coulteri) have very thickened seed burnedare [Link],
wing bases, filled with light, spongy tissue. The dispersal under normalconditions requiresan effec-
outline of effective wings is oblique, usually with tive wing only; the articulatewing is usually as good
one more or less straightside which was adjacentto as an adnate wing in this respect. Perhaps the loss
the other wing on the scale. The colour is usually a of the wings serves to camouflage the seeds against
translucentlight brown with darkerstripesor streaks. [Link] reductionof the wing below effective-
"Wingless"seeds occur in two ways, depending on ness, and/or its loss before dispersal, is connected
whether the wing remnantis adnate, in which case with zoochory (Lanner, 1982). "Wingless" seeds,
it remains attached to the seed, or whether it is occurringin our region in both sect. Strobusand sect.
articulate, in which case it usually remains loosely Parryanae-and also in the only distantly related
attached to the scale and leaves no trace (or a very Mediterraneanspecies, P pinea-evolved in coevo-
faint one) on the seed. lution with birds (jays and nutcrackers;see Tom-
There is evolutionary significance in the various back & Linhart,1990; Mattes, 1994). They are there-
modes of seed wing attachmentand development. fore examples of parallelismthrough selection of a
The more common anemochorous dispersal would morphological potential inherent in the genus as a
be most effective when the seed is small relative to whole: a trend towardcompletion of the articulation
the wing and the wing adnate. Indeed, this situation of the seed wing. Enlargement of the seed and
is found in Pinus subsect. Strobi, in which the cone strengtheningof the seed coat are othertrendsrelated
is pendent and opens readily on maturityto release to this highly selective type of bird dispersal, which
the seeds simply by gravity. Trees of several of its in some cases is connected with a single bird species
species are very tall (P ayacahuite, P lambertiaaa) (Mattes, 1994).

WOOD ANATOMY (by Ian D. Gourlay)

ANATOMICAL DESCRIPTION OF taxodioid pits. Some genera contain xylem

PINACEAE (according to Greguss, 1972) parenchyma,others lack it (Pinus). Some gen-
era exhibit spiral thickening in their late tra-
Cross section-Annual ring boundariesnor- cheids, other genera have it in the axial and
mally conspicuous. Resin ducts presentor ab- transversetracheidsof all their species.
sent. Epithelial cells thin or thick walled. Tangential section-Rays one to 45 cells
Some genera possess axial parenchyma. tall, most of them typically uniseriate,except
Radial section-Ray tracheids present or those including resin ducts. End walls of lon-
absent. Some species have rays without trans- gitudinal parenchymacells nowhere smooth.
verse tracheids or resin ducts. Some species
have both thick and thin walled ray paren-
GENERALINTRODUCTIONTO PINUS
chyma cells. Ray tracheidsmost often charac-
terised by different forms of [Link] The wood of pines, like that of all conifers, is a
of the ray tracheids smooth Pinus, group simple structurecomposed of tracheids and paren-
Haploxylon = subgen. Strobusor dentate [Pi- chyma cells. The tracheidshave lignified walls char-
nus, group Diploxylon = subgen. Pinlus].The acterisedby the presence of borderedpits. The rays
end walls of the ray cells exhibit sieve-like are usually one cell wide and rarely visible without
[Link] fields exhibit one to six (sel- a lens. These vary in depth, the central rows con-
dom eight) pits, large oval pinoid pits filling sisting of parenchymawith simple pits on their radial
out the entire field, one to four small round walls, the upper and lower rows consisting of tra-
piceoid pits with oblique slit-like aperturesor cheids with borderedpits.

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Wood Anatomy 31

The genus Pinus is readily distinguished from and colour, and show a more gradualtransitionfrom
other genera by the presence of resin ducts distrib- early to late wood zones. Resin ducts are not as
uted throughoutthe growth ring and surroundedby numerous. Their ray tracheids have smooth walls
thin-walledepithelialcells. It is in these ducts during and include P cetmbraof Europeand otherAmerican
theiractive life that the productionof oleoresin takes species including P. strobus and P lambertiana. In
place. The ducts are formed as intercellularspaces Mexico, P avacahuite and P strobiformisare com-
between the resin-producingthin-walled epithelial mercially importantmembersof this group.
cells, which are often damaged in the course of In the "hard"pines, the bark is rough, grey, and
sectioning. Resin ducts/canals are often thought to scaly, thickly covering the trunks of even young
be developed as a response to injury; however, in trees. In the "soft" pines, bark formationis delayed,
the pines this is not necessarily so (Mirov, 1967). giving the appearance in young trees of a very
The product of this response to wounding in the smooth [Link] bark formationdevelops at the
pines is often exploited in commercial turpentine base with age, gradually extending up the tree but
extraction. often with the uppertrunkremainingsmooth (Perry,
The wood of pines frequently shows a marked 1991).
transitionbetween early and late wood, togetherwith Jones (1924) subdivided the genus according to
a generally reddish-colouredheart. It is also, on the needle grouping: (a) two-, (b) three-, and (c) five-
whole, light, elastic, strongfor its weight, and widely needle pines:
available. Close-grown and plantationmaterialwill
a. Autumn wood of each ring is well-defined with
normally produce stems of straight form, ideal for numerous resin ducts. Rays are uniseriate and
a wide range of construction uses. This structure,
multiseriate(the latter usually containing a hori-
therefore,can in some species (e.g., Pinus strobus) zontal resin duct). Tracheidcells in the rays usu-
present a fairly uniform wood, easy to work and of
reasonable durability when there is a high resin ally occur only towardthe upperand lower mar-
content. Pine timber can also be used for producing gins, and there is pronouncedirregularthickening
of the walls of the ray [Link]
paper pulp, although it is less suitable than that of cells occur medianly,with large simple pits.
some of the other conifers, such as spruce (Picea),
b. Thoughsimilar in most characteristicsto the two-
because of its resin content.
needle pines, the smooth walls of the raytracheids
are in strong contrast with the jagged walls of
GROUPINGOF SPECIESACCORDING the two-needle pines.
TO ANATOMICALFEATURES c. Though similar to two- and three-needlegroups,
the rays differ in thatthe simple pits are consider-
Several attemptshave been made to furthersubdi-
ably smaller and more [Link],the
vide the genus, most commonly into two groups, the
ray tracheidshave comparativelysmooth walls.
"hard"and "soft" pines (referred to in the United
States as the southern yellow pine group and the It should be noted that our currentunderstandingis
white pine group). that the numberof needles in a cluster is not a good
The "hard"pines (Diploxylon, Fig. 9) possess two phylogenetic marker(see Mabberley,1987).
vascularbundlesper leaf and their wood is generally A furtherclassificationof the genus, on the basis
harder,heavier, and darker in colour, with a much of form of smooth or dentate ray tracheids, was
clearer demarcation between early and late wood presentedby Phillips (1941). He stated that species
zones within a growth ring. The walls of their ray identification is not generally possible unless the
tracheidsreveal thickened,uneven, tooth-likeprojec- geographical origin of the material is known, but
tions (dentation).Resin ducts are [Link] detailed microscopicexaminationwill usually permit
this group are the common commercial northern a specimen being classified underone of the follow-
timber trees, Pinus sylvestris and P. nigra, together ing seven groups (with added Mexican species given
with the majorityof NorthAmericanspecies. Almost in parentheses)recognized by Rol (1932):
all of the economically importanttimber trees of I. [Link] tracheidswith smooth walls; cross-
Mexico and CentralAmerica and all the Caribbean field pits 1-3, large, tangentialwall pits on late-
pines also belong to this group. wood tracheidsnumerous,e.g. P strobus,P cem-
The "soft" pines (Haploxylon, Fig. 10) possess bra, P. koraiensis, [Link]. (P avacahuite
one vascular bundle per leaf, are lighter in weight and P strobifornmis).

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32 Flora Neotropica

.88~'~:: .:~88~88I;::: . .. . ........

":?'-~
BBBBsBg:,:?::
B~B~:.:
?????~ .............
...?.?:::?.?:-'?Bg~H~k~:r:
?????:
?????
??Y
9Bh~'P~n~: ..............
F I i
vertical
res ..anG. dn a cd l eu a rc l 9.P.P.A.... show.
..........t..
[Link]..e..[Link]..[Link].....B..Tangential....ion,
showing
a horizontal
resinducwithi.. Radia l si

tracheids and larger

. (3-6)......cross-field
:'::i~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
.....os0.
........ ... . ...... ..
pits...........agifc ... ...............~
....... ...... ~~~~~~~~~~~~~~~~~~~~~~~~~~
t

~~~~~~~~~.....
,,...... ..p~
~ ~
Ei~~~~~~~~~~~~~~~~~~~~~~i IAi
?II'?lli
..... ....

i:lrii :I:I~~~~~~~~~~~~~~~~..
''''''''''
... .
...
.
...

:~~~~~~~:?:? I?.-.?.-.-~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~....
i::i~i~i.Z~i~~~P::ii' ...

ij~4'3
?

~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~'..-:.7...
.
.. .... .. . .

:?:?:?:?: :::~~~~~~~~~~~~~~~~.. ...... ...

::: ... iii . ii:::i!'i:

:?:~~~~~~~~~~~~~~~~~~~.
'~i .

?I: S8~~~~~~~~~~~D
j~~~~B~~~-i~~il
8~...
l,l,l'l II .... .. .............

i:::::::g~~~~~~~~~~~~~~~~~~~~~~~~~~~...
B3I:::~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~..... .....
.........

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

.... .. ...... Ix ::::::::.::::::

.... . ....

..........~~~~~~~~~.
...... . .....

FIG. 9. Piiiiis ooc(irpi, a~~~~~~~~~~~~~~~~~~~~

of the "hard"pines (Diplox'i-loit subgen. Piiiiis). A. Transverse section, showing
verticalresinduct and a typical
demarcatedzone between early and late wood. B. Tangentialsection. showing a
horizontal:::::
resi dut wihina ry. clearly
C Raialsecton,showng hicenedtooh-lke pojetion (dntae) i th wals o th ra
trachidsad lager (-6) coss-ield its. Magnficatons x100.

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Wood Anatomy 33

g l ~~~~~~~~~~
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~l

?..........

~~~~~~~~~...
~~~~ ~~~~~~~~~~~~~~~~
. :'~'"
.
.
....0
" :... ... . ..... ......

wE _z ?`: -

_i~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

?. :i:?'-i ? "::i:i::.:
::!':.~:~ ':::. ....
.... :~i:~:.!ii"':i"1i~?:;::!i :iiiiiii~::~'i::~
'::.-':!:iiiiiiii~?:i: : ?:i:::i." ::::
'
i::i ii ii:ii- : !ii : ' ii

4 crs-ied . MgA. I0 C. 25._ _

FIC,.[Link]~tltsceotbroidles.
typicalof the"soft"pines;(Halplo.-v~lo.t
= [Link]).
A. Tralnsiverse
[Link]
two vertically
resinductsand a less [Link] transition, fromearlyto late woodl.B. Tangentiall
section.
showinga horizontalresinductwithinaI ray.C. Radialsection. showingraytracheidsi withsmoothwallsandsmall(2-
4) [Link]-field A. B. x 10():C. x2:50.)
pits.(Malgnificaltions;:

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34 Flora Neotropica

2. [Link] tracheidswith smooth walls; small ray tracheids, e.g., P ponderosa, P taeda, P
cross-field pits 2-4, piceoid to pinoid; tangential palustris.
wall pits on latewood tracheidsnumerous,e.g. P
Frey-Wysslinget al. (1955, 1956) suggested that
gerardiana (P cembroides).
3. Sula. Ray tracheids inconspicuously dentate; pines with dentate ray tracheidshave "warts"on the
inner wall of the borderedpits of the axial tracheids
cross-field pits 1-4, pinoid; tangential wall pits
(observablewith electronmicroscope),and thatpines
absent,e.g. P halepensis, P canariensis, P leuco-
with smooth-walled ray tracheids have no "warts."
dermis, P roxburghii.
Furtheranatomicaldescriptionsof individualspecies
4. Ponderosa. ray tracheidsdentate; cross-field pits
were given by Greguss (1972).
1-4, pinoid; tangential wall pits absent, e.g. P
ponderosa, P. contorta, P patula, P pittaster, P
radiata. CENTRAL AMERICAN PINES
5. [Link] tracheidsdentateand reticulate;cross-
field pits 3-6, pinoid; tangentialwall pits sparse, Although few of the above charactersare observ-
able in the field, the terms"hardpine"and "soft pine"
e.g. P palustris, P banksiana, P. caribaea, P have significance in Mexico and Central America
echinata, P taeda.
because species in the soft pine group are much
6. Khasya. Ray tracheids inconspicuously dentate;
cross-field pits 1-2, large; tangentialwall absent, sought after, as they are more easily worked by
hand and less resinous (Perry, 1991). The turpentine
e.g. P kesiya.
7. Sylvestris. Ray tracheids dentate; cross-field pits industry in Mexico and Central America is quite
large and is based on the collection of resin from
1-2, large; tangential wall pits rare, e.g. P. sylv-
species in the hard pine group.
estris, P densiflora, P nigra, P resinosa.
Wood anatomicaldescriptionsof 12 Mexican tim-
A more detailed breakdowncomparingboth gross ber trees including eight pines-P ayacahuite, P
morphologicaland wood anatomicalfeaturescan be cemnbroides, P leiophylla, P monophylla,PFnelsonii,
found in the work of Hudson (1960), who further P pinceana, P quadrifolia, and P strobus var. chia-
attempted to classify the pines by the degree of pensis-can be found in Crespo (1963). A summary
dentation in the ray tracheids. He stated that slight of the charactersstudied in each species is included
swelling and low dentationwere seen to be common in the condensed tables. The form, frequency, and
in those species that have smooth-walled ray tra- dimensions of the tracheids, rays, parenchyma,and
cheids. It is to be appreciated, however, that the resin canals are reported,together with photomicro-
classes described are arbitraryand that some diffi- graphs.
culty is involved in deciding into which class some Adopting the system proposedby Martinez(1948,
specimens should be placed. The classes attemptto 1953) Crespo placed P cembroides, P monophylla,
show the gradual development of the ray tracheid P nelsonii, P pinceana, and P quadrifolia in his
dentationfrom slight swellings to reticulations. pinyon section. These species were characterizedas
Classificationof the ray tracheidsaccordingto the having in common 1-4 piceoid pits (with the excep-
degree of dentation (Hudson, 1960; some classes tion of P nelsonii, 1-6), small areolate pits in the
with minor differences have been taken together summer wood tracheids, smooth-walled ray tra-
here): cheids, and ray cells with pitted horizontalwalls and
nodulartransversewalls. Interspecificdifferences in-
1. Slight thickening of the wall adjacent to the
cluded the fact that P monophylla possesses very
bordered pits of the ray tracheids, e.g., P.
wide rays, whereas splits in the tracheid walls of P
cembra.
pinceana and P nelsonii are visible.
2, 3, 4. Very slight dentations most pronounced
Martinezplaced P. ayacahuite and P strobus var.
near the borderedpits of the ray tracheids,e.g.,
P koraiensis, P strobus, P wallichiana. chiapensis in his Ayacahuite section, and Crespo
describes them as differing from the pinyon group
5. Moderatelydentateray tracheids,e.g., P pinea.
6-10. Gradualdevelopmentfromdentateto heav- by having pinoid or fenestroid pits, 1-2 per field,
with slightly toothed walls in the ray [Link]
ily dentate ray tracheids, e.g., P merkusii, P
leiophylla is also placed in this group (wood anat-
roxburghii,P cubensis, P pintaster,P resinosa.
I I. Heavily dentate, with some reticulate ray omy, but not Martinez'sgrouping) but differs in that
the ray tracheidsare more clearly toothed.
tracheids,e.g., P banksiana.
12-14. Partiallyreticulateto completely reticulate

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Karyology 35

The above-mentionedMexican species and others appearto correspondwith the general morphological/
examined at the University of Oxford (i.e., P cacri- anatomical features that attempt to separate pines
baea, P ntontezumae,P oocarpa, and P ponderosa) into hardand soft groups.

KARYOLOGY

All species of Pinus have the same haploidnumber Morphologicaldifferences between Pinus leiophy-
of chromosomes: n = 12; 2n = 24 (Mehra & lla and P lumiholtzii,two species forming subsect.
Khoshoo, 1956; Saylor, 1972, 1983; Mehra, 1988). Leiophyllae,include resinousand non-resinousbuds,
This is also the predominantbasic number in the 3-5 and 2-3 leaves per fascicle, medium size and
remainderof the Pinaceae, where only one species long leaves, low and high numberof stomatallines,
in Pseudotsuga (P menziesii, n = 13; 2n = 26) and uniform and pluriform hypodermis in the leaves,
the monotypic genus Pseudolarix (n = 22; 2i = and three- and two-year developmentof ovuliferous
44) are known to be [Link], or even cones, [Link] only conspicuous similarity
the additionor loss of a single chromosome,does not is the deciduous fascicle sheath, which appearsas a
exist in the genus except for an occasional individual uniquely shared characterin subgen. Pinus but as a
(Saylor, 1983). Mirov (1967) reportedthat attempts parallelismin the genus as a whole. Yet it is the only
to artificially induce polyploidy in pines met with reason for their classification in a subsection or
difficulty and abnormal development in almost all "group"(Shaw, 1914; Little & Critchfield, 1969).
cases. Chromosomestructureis equally conservative. PinusI leioph!lla and P lumholtzii have different
Karyotypes,based primarilyon centromerelocation karyotypes.A noticeable difference was found be-
and arm lengths, are constanteven within very wide- tween the karyotypes of P elliottii (including var.
spreadspecies such as Pintussv,lestris (Mirov, 1967), densa) and P caribaea (including its varieties) in
and differ but slightly among species. The most chromosome No. 11, which links the latter to P
comprehensiveanalysis has been providedby Saylor occidentailis with a similar pattern. This evidence
(1964, 1972, 1983), covering 87 species in [Link] supports the classification proposed in this mono-
I I largerchromosomes of all species are similar in graph. Another interesting observation is that the
length and have median centromeres,i.e., two arms karyotypeof P [Link] closely resembles
of equal length. Only the smallest chromosome [no. P. ponderosa and its relatives than it does the mor-
12, also No. 11 in species of subsect. Sylvestres phologically similar P pseudostrobus,with which it
(= subsect. Pinus)] is consistently heterobrachial is also largely sympatric. Pinus lawsonii and P
(centromeresubmedian),with a ratio of ca. 0.6 be- teocote have similarkaryotypeswhich set them apart
tween the a and b arms. from the other species classified in subsect. Pondero-
Nevertheless, more detailed observations, using sae (Little & Critchfield, 1969). The species in the
charactersequences as summarizedby Saylor (1972, rather heterogeneous subsections Ponderosae and
1983), allowed species comparisonsand yielded oc- Oocarpae as delimitedby Little and Critchfieldhave
casional differences of karyotypeespecially at taxo- been rearrangedby van der Burgh (1973), who ac-
nomic levels above species. Some of these, such as commodatedboth species in the lattersubsection,an
the additionof No. I I to the heterobrachialchromo- arrangementthat is supported by the classification
somes in subsect. Pinus, seem to delimit such group- proposedhere.A similarbreakupseparatingthe "Cal-
ings of lower [Link],however,appearto group ifornia closed-cone pines" from Little and Critch-
species across the two subgenera. There are also field's subsect. Oocarpae(van der Burgh, 1973) finds
some differences between species karyotypeswhich supportin the karyotypes,which are similarfor these
seem helpful in elucidating their relationships. In pines but differ from the Mexican species originally
general,Saylor (1983) concluded,that karyologyhas grouped in that subsection.
"not provided evidence to contradict major taxo- In subgen. Strobus only a few Mexican species
nomic relationshipsestablishedpreviously,"referring were studied (Saylor, 1983), some of which also are
to the classification given by Little and Critchfield widespread in the United States. It was found that
(1969). A few of his observations relevant to Latin the karyotypesfor Pinus flexilis and P strobiformis
American pines will be mentioned and discussed are rathersimilar but that P ayacahuite differs no-
here. ticeably from these two taxa. Since there appearsto

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36 Flora Neotropica

be a geographical dine in several morphological is a serious omission. Unfortunatelyalso, only three

characters of the first two species, with P. strobi- species of Little and Critchfield's subsect. Cem-
formis approachingP. ayacahuite in its more south- broidces were studied-P cembroides, P edullis, and
ern populations(see discussion with the species ac- P monophylla-thereby leaving out all the more
counts in this monograph),it would be of interestto distinct ones which could have yielded interesting
have known the localities from which the samples [Link] surprisingly,the three taxa investi-
for Saylor's studies [Link] informationabout gated appearedto share similar karyotypes.
the origin of the samples (seedlings) is given, which

POLLENMORPHOLOGY(by Marie H. Kurmann)

Pollen grains of Pinus are bilateral,characterised to date. For each of the species, she listed the size
by a centralcorpus with two sacci placed symmetri- and shape of the corpus, shape and ornamentationof
cally, one on each side (Fig. IA-C). They range in the distal area, structureand shape of the saccus,
size from 70 to >100 /jm. Surface sculpturingand ratio of saccus height to width, and the proximal
wall stratificationvary around the pollen grain. In attachmentof the saccus to the corpus. She con-
the distal area, the sculpturing ranges from psilate cluded that it is difficult to separatepine species on
(Fig. IIA) to scabrate (Fig. IIB). The distal wall the basis of pollen morphology and considered it
(leptoma)consists mainly of a thick, lamellatedend- impossible to draw phylogenetic conclusions. Her
exine (Fig. I IF). The proximal surface is more or results do not contradict the classification into the
less rugulateand the proximal wall (cappa) is thick, two subgenera,but neitherdo they allow their recog-
consisting of an outer, alveolate ektexine subtended nition on the basis of pollen morphologicalfeatures
by a narrow lamellated endexine (Fig. IG). In the alone. Erdtman(1965) described the pollen of 25
lateralregions, the ektexine is split to form the sacci. Piins species. In most instances the characters he
The saccus floor (Fig. II E) consists of a lamellated used concern size and size ratios. Klaus (1975, 1978)
endexine and the ektexinous foot layer, and the also investigated the pollen morphology in Piiins
saccus roof is made up of the tectum subtendedby species, using not only light microscopy but also
endoreticulations(Fig. 1D). scanning and transmission electron microscopy.
Pine pollen is morphologicallyrelatively uniform Electronmicroscopy appearsto enhance species dis-
in most species, and the range of intraspecificvaria- tinctions, not only by providing new charactersbut
tion overlaps with the interspecificone. Furthermore, also by clarifyingcharactersused in light-microscop-
preparationmethods may affect morphologicaland ical studies. Morphologicallyuseful charactersin his
size charactersdifferently. It is therefore important account included saccus structureand variation in
to take preparationtechniques into account when corpus structuresas revealed by transmissionelec-
describing the pollen grains (Kurmann& Zavada, tron microscopy. He concluded that in individual
1994). There seems to be some evidence that pollen cases it might be possible to differentiategroups or
morphologicalcharactersdiffer between the two sub- even species if a large number of characters were
genera Strobus(Haploxylontype) and Pinus (Diplox- used. He also consideredornamentationof the distal
ylon type), but there is also a group of species in area to be the most useful characterin differentiating
which these are mixed. In the Haploxylon type, the the two subgenera.
sacci are usually broadly attached,the distal area is Pinus pollen is often found in great abundancein
sculpturedand is often borderedlaterally by thick- fossil deposits. Since differentspecies have different
ened ridges (e.g., Pinus rzedowskii,Fig. II B). In the climatic requirements,identification of these fossil
Diploxvlon type the sacci are contractedalong their grains to the species level would greatly enhance
roots. The distal area is usually smooth and is not [Link] studies
borderedby a thickenedridge (e.g., Pinus leiophylla, have been undertakento identify modern Pinaceae
Fig. IIA). pollen grainsto species level in restrictedgeographic
In 1950, van Campo-Duplan published a study areas. In an attemptto distinguishmodernspecies of
of the Pinaceae based on pollen morphology. She Piins in the Pacific Northwest, Hansen (1947) used
investigated ca. 60 species of Pinus pollen with size frequency. However, this approach has been
the light microscope, and this remains the most questioned by Mack (1971). Hansen and Cushing
comprehensivepalynological treatmentof the genus (1973) proposeda method for distinguishingspecies

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Pollen Morphology 37

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ekeiean nein aela E.(Sae..s arw) -.10Cm -G r.

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38 Flora Neotropica

which combines quantitative and morphological considerationof preparationtechniquesand a combi-

[Link] (1985) also demonstratedthat for nation of structuraland quantitativecharactersmay
some geographicgroups, pollen identificationat low prove to be useful in distinguishing species in re-
taxonomic levels is possible if a combination of stricted geographic areas. Electron microscopy pro-
structuraland linear charactersis used. vides additionalcharactersand should be employed
The relative uniformityof pollen in Pinus makes in attemptsto identify pine species based on pollen
it difficult to classify it at the species level. Careful morphology.

THE ROLEOF MONOTERPENESIN THE TAXONOMYOF THE MEXICAN

AND MESOAMERICAN PINES (by JacquelineS. Birks)

INTRODUCTION [Link] are the subset of terpenoids

which have exact multiples of the isoprene units
Plant isoprenoids are compounds derived from
(hemiterpenes, C5; monoterpenes, C)o; sesquiter-
[Link] are divided into primaryand second- penes, C15;
diterpenes,C20).There are thousandsof
ary [Link] secondary metabolitesof plant known terpenes and
many have been found in the
cells, chemicals of low molecular weight that are resins of
pines. They have a role in disease resistance
not directly involved in essential metabolic action, and defence
against insect attack.
include the monoterpenes, the sesquiterpenes, and The biosyntheticpathways,by which the terpenoid
the diterpenes. The secondary metabolites of the constituents of resin
are formed, are difficult to
genus Pinus include a wide range of chemical prod- determine
directly (White, 1983) and were little un-
ucts with industrial or pharmaceuticaluses. As a derstooduntil
significantadvances were made in the
result, the chemistry of many of these productshas last decade (Gershenzon& Croteau,
1990). A review
been thoroughly investigated and this in turn has
by Chappell(1995) emphasises the progressthat has
often been the precursorof more fundamentalscien- been made in
understandingthe enzymes involved
tific [Link] results of this extensive research in the
biosynthetic pathways but reminds us that
generated interest in measuring and describing the there is still much that is not understood.
chemical components and using this informationas Resin samples may be obtained from steam dis-
a purely scientific tool to investigate relationships tillation of leaf oils
(Zavarin, 1970) or from cortical
between the different species. tissue (Squillace, 1977; Meier & Goggans, 1978) or
Mirov (1967) reviewed the full range of chemical
xylem tissue (Franklin, 1976; Strauss & Critchfield,
products of pines, which are numerous. Although 1982). The cortical resins are easier to collect
(Squil-
attempts at classification and delimitation of taxa lace, 1977).
have been made with all secondary compounds for
pines, the components of the oleoresin, and particu-
larly the monoterpenes,have attractedthe most atten- Gas Chromatographic Analysis and the
tion and yielded the most contributionsto the taxo-
nomic debate. Quantitative Results
Gas chromatography(GC) is an efficient and pre-
cise method for determinationof resin composition
The Pine Oleoresin
(von Rudloff, 1975). Results appear as a plot of a
The mixtureof rosin and essential oils that can be series of peaks, each of which can be identified as
obtained by cutting the resin canals, of leaves or of belonging to a particularcompound, using previous
the wood, is known as the oleoresin but is usually results from known [Link] area undereach
referredto simply as the resin. Resins are mixtures peak measures the relative proportionof each com-
of compounds including terpenoids,flavonoids, and pound presentin the sample. Results are reportedon
fatty acids. Essential oils are volatile oils that are a number of different bases (Squillace, 1976). For
recovered from plant tissue by steam distillation or monoterpenes three bases are commonly used: 1)
solvent extraction. The terpenoids, comprising C5 relative to all other monoterpenes (total monoter-
isoprene units, have been the most widely studied pene), 2) relative to all other constituents of the
class of compounds extracted from pines because oleoresin (oleoresin basis), and 3) per unit weight or
they are easily identifiedand quantifiedby gas chro- volume of tissue (tissue basis). The data defined by

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The Role of Monoterpenesin the Taxonomyof the Mexican and MesoamericanPines 39

the latter are usually described as absolute amounts production of certain monoterpenes is often con-
(White, 1983), althoughthereis nothinguniqueabout trolledby single genes with majoreffects. The belief
this base as other absolute amountscould be defined. that the non-genetic effects are minimal or easily
The proportionsof compounds are independentof controlled is what made resin composition such an
the total yield of oleoresin, but data reportedon a interestingtrait for taxonomists.
tissue basis are not. White and Nilsson (1984) found
that the yield of leaf oil monoterpenes in Pinus
Monoterpenes Found in Mexican and
contorta was relatedto the frequencyof resin canals
Mesoamerican Pines
in the leaves, which was itself variable and subject
to genetic control. They concluded that the propor- The oleoresinof Mexicanand Mesoamericanpines
tions of compounds should be used to study differ- is clearly differentiatedfrom that of other pines, the
ences between populations. Shaw et al. (1982) main constituents being a-pinene and /3-phellan-
reachedthe same conclusion but cited the confound- drene./-phellandrene is almost uniqueto the species
ing effects of environment and genotype on yield. of Mexico and Mesoamerica(Mirov et al., 1965). A-
Consequently, most studies of resin composition 3-carene and longifolene are quite common constit-
have been based on the interpretationof data com- uents of the oleoresins, again rarelyfound elsewhere,
prising the proportionsof individual terpenes de- their only other occurrencebeing in the pine species
tected in resin samples. Such data are referredto as of western North America. In the north of Mexico
compositionaldata. there is evidence of a different oleoresin. The oleo-
resin of Pinus jeffrevi, a species whose southern
extent is the north of Mexico, is dominated by
Variation in the Composition of Resin
heptane, with minute proportions of a-pinene,
The resin is synthesised in the resin ducts of the whereas P coulteri, which also just extends into the
different tissues. Each of these samples an indepen- north of Mexico, shows a more typical oleoresin
dent system (von Rudloff, 1975), and the composi- dominatedby a-pinene, plus /3-phellandrene.
tions of the oleoresin obtainedfromdifferentsources,
from a single tree, are not the same. The composi-
THE USE OF MONOTERPENE DATA
tions of foliar oils and cortical resins are often simi-
lar. The composition of xylem resin is often less IN TAXONOMIC STUDIES
complex than that of cortical resin (Squillace, 1976). Monoterpenedatahave usually played a role com-
The composition varies within the same system at plementaryto other taxonomic charactersand there-
differentpositions in the tree. Forexample, the xylem fore have been introducedto solve the more intracta-
resin has been found to vary with the height of ble problems concerned with species delimitation
sampling (Rockwood, 1973; Roberts, 1970). There in groups of closely related species alliances or
is variation with the developmental stage of a tree widespreadpolymorphicspecies complexes. Initially,
and with the season of the year (von Rudloff, 1975). qualitative or discrete characters (the presence or
Moisture stress has been shown to affect the resin absence of compounds) were used to examine rela-
composition in some species (Langenheim et al., tionships or delimit taxa, but when the group of
1979), and insect infestation has been shown to species being investigated displayed the same set
increase monoterpeneand diterpeneproductionin P of compounds, quantitative traits were used. The
contorta (Raffa & Berryman,1983). possibilityof single-gene controlof some compounds
Because there are so many factors affecting resin is often mentioned but not seriously addressed in
composition, apparentlycontradictoryresults are re- taxonomic studies.
[Link] of P monticola,
growing in diverse environments,showed negligible
differences in terpenecomposition (Hanover, 1966), Qualitative Characters
but genotypically identical trees of P sylvestris Occasionally the monoterpenedata have shown
grownon threevery differentsites in Sweden showed discrete, qualitativedifferences between species, but
moderatedifferences in terpene composition (Bara- these results have often reflected inadequate sam-
dat & Yazdani, 1988). pling at one or more levels. A compound that is
Hanover (1966) and Squillace and Fisher (1966) declared missing in a population in one study may
believed thattheirresearchshowed thatthe composi- be detected in small proportions,or at low frequen-
tion of resin is not only strongly inherited but the cies, in [Link] early comparisonof resincompo-

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40 Flora Neotropica

sition of Piins monophylla and P quadrifolia (An- raised to species rankfrom that of variety within the
derson et al., 1970) found that B/-phellandrene, previous two decades on the basis of morphological
terpinolene, and camphene were missing from P evidence. Zavarin and Snajberk, with others, have
qucadrifoliabut were presentin P. monophylla:but a conducteda series of investigationson the monoter-
laterstudy of these two species (Zavarin& Snajberk, pene traits of subsets of the pinyon pines, together
1980) shows these threecompoundspresentin small with morphologicaltraits, in order to evaluate these
proportionsin both species. An investigationof the taxonomic revisions. Zavarinet al. (1980) compared
closely relatedspecies P estevezii, P. pseudostrobus, P quadrifolia with P monophylla: Bailey et al.
and P montezunae (Perry, 1982) showed heptaneto (1982) comparedP discolor with P cembroides:and
be present in the oleoresin in all samples of P Zavarin and Snajberk(1985, 1986, 1987) looked at
estevezii but absent in all samples of the other two differences between populations within P cem-
species. However,only 10 trees from one population broides, compared P discolor with P johannis, and
of each species were examined, and this interesting investigated differences among P culnirnicola, P
observationrequiresfurtherinvestigation. nelson ii, P pinceana, and P mraxilartilnezii. The
first two papers deal with the question of (putative)
hybridizationbetween these species. In both studies
Quantitative Traits there is an apparentlack of understandingof what
taxonomic characters are: "deviating values" (e.g.,
The terpene traits are used to investigate species
when these fall within the range of the wrong spe-
relationshipsas a basis for classifications and, more
cies) are left out of the [Link] same problems
widely, for the delimitation of species (Zavarin & are apparent in Zavarin and Snajberk (1987: table
Snajberk, 1973). Most studies employ monoterpene
data as though the proportionof each compound 3), where, for instance, P maximartine-iiis present-
ed with a mean of 88.6% limonene, but a sample
were a continuous variable measuredindependently
of the other compounds. The means and variances measuring only 16.8% from the same population
is left out as a "deviating value." We are not in-
of the proportionsof each compound are analysed
formed about the percentages of the other com-
between and within populationsusing univariateand
multivariatestatistical methods. pounds, which must of necessity be higher than in
the "mean." Since this highly distinctive species
It is this wide use of quantitativeterpene traits,
does not hybridize with any other (to explain the
nearly always in the form of proportions,that has
deviation),the figuresgiven in the table are meaning-
been questioned, and discussed in depth (Birks &
less as taxonomic characters.
Kanowski, 1988, 1993) in relation to the known
The separationof Pinus patula and P oocarpa is
problems inherentin the analysis and interpretation not usually disputed, but the status of other closely
of data describing compositions.
related taxa has caused considerable controversy.
Many interpretationsof resin compositional data
have acknowledged but not acted upon the concep- Monoterpenedata were used to investigate the status
of P patula var. longipeduncultata,P tectunumanlii,
tual difference between compositional data and a
and the possible hybridizationbetween P oocarpa
multivariatedata set, where the same constraintsdo
and P caribaea var. hondurensis(Squillace & Perry,
not apply. These proportionsare not metric traits in
the usual sense. They are not measuredindependently 1992). The statistical procedures ("renormalizing"
of each other,and methodsof analysingmetrictraits, proportions to sum to 100% and assuming 10%
limonene even where this constituent was not as-
such as analysis of varianceor t-tests, are invalid if
sessed) as well as the presentationof data in the
applied to single componentsof these data. Multivar-
iate methods of analysis, such as principal compo- analysis makean assessmentof taxonomiccharacters
from this paper [Link] appears to be
nents analysis, are invalid if appliedto compositional
little support for the taxonomies based on morpho-
data.
logical data from the monoterpenesanalysed in this
study.
Some Regional Examples of Taxonomic
Studies Using Monoterpene Data CONCLUSION
The Mexican pinyon pines, Pinus subsect. Cem- Absolutedeterminationof resincompositiondefies
broides Engelmanns.l., comprise seven species and definition and measurement; the only assessment
three varieties or subspecies. Several taxa have been possible is of the proportionsof differentconstituents

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ReproductiveBiology 41

in a sample, but these cannot be defined uniquely. ways that do not constitute meaningful taxonomic
The analyses of the data are all affected by some or characters.
all of the problemsdescribedby Birksand Kanowski It is possible that the presence or absence of
(1988, 1993). Inferentialstatistical methods applied a constituent may provide taxonomic charactersas
to single components of a composition, as has been defined by Davis and Nixon (1992), but there has
done in the studies described here, are not correct, been little evidence so far that these exist at the
and the results will be misleading. Descriptivemeth- species level. The conclusion must be that,consider-
ods of data analysis applied to a subcomposition ing the combined problems of the analysis of the
may help in clustering populations, but these are data of compositions together with the unsuitability
superficialmethods for taxonomic studies. The stud- of the traits derived from the monoterpenedata as
ies described here not only fail to address these taxonomic traits, we are not in a position to review
problemsbut also attemptto use quantitativedata in the taxonomy of the pines.

REPRODUCTIVE BIOLOGY

Generalbutdetailedaccountsdescribingthe repro- opening with a closing mechanism("neck")and two

ductive biology-including pollination,the ontogeny appendages("arms").The sticky fluid on these parts
of the reproductiveorgans, and embryology-in co- (Fig. 8) causes pollen grains to adhere to their sur-
nifers are given by Strasburger(1869, 1872), Pilger faces. Reabsorptionof the pollinationdrop, possibly
(1926), Chamberlain(1935), and Hirmer (1936); a by surface tension or partial drying, brings some
synopsis is presentedby Weieret al. (1982), exempli- pollen grains into the micropylarchamber, where
fied by PinLus. one of them germinatesand grows a pollen tube into
the nucellus. During this process, the germinated
POLLINATION pollen grain slowly develops into a tubular male
gametophyte. Meanwhile, the micropylarcanal has
The pines, like almost all gymnosperms, are en- been closed by a swelling of the neck cells. With
tirely wind [Link] produceenor- this, pollinationis completed.
mous quantitiesof pollen, which can be blown great
distances from its source. Pollen dispersal(anthesis)
FERTILIZATION AND EMBRYOGENY
times vary among species in our region from some-
time in November/Decemberto June/July,coinciding In Pimus,a single megasporocytelies in the centre
with the dry season in most regions of Mexico and of each ovule; it is surroundedby the nucellus,
[Link] late springdispersalsare found containing nutritive tissue. The pollen tube grows
mostly in taxa with northerlydistributions and of through the nucellus and reaches an archegonium.
high altitudes (e.g., Pinus hartwegii). Pines, being This development requires about one year to be
monoecious, are frequentlyself-pollinated, although completed, when the pollen tube releases its contents
cross-pollinationis stimulatedby the relativeposition into the archegonium. Meanwhile, the megasporo-
of male and female strobilion the branchesof a tree. cyte in the ovule has been divided by meiosis into
At the time of pollen dispersal, the female strobili four haploid megaspores;one of these furtherdevel-
have emergedfrom the buds and have slightly parted ops into a female gametophyte,containing archego-
their scales, permittingwind-blown pollen grains to nia surroundingan egg cell (Fig. 8). When the male
sift down to the axis of the cone, where they come gametophyte reaches the archegonium, the female
into contactwith a sticky substance(pollinationdrop) gametophyteis fully developed and ready for fertil-
secreted by the ovule. This fluid was first observed ization (Fig. 8). One of the two sperm nuclei then
by Strasburger(1872), and its role in transporting unites with the egg nucleus. Theoretically,since the
the pollen up to the receptive surfaceof the nucellus female gametophytecontains two or more archego-
was definitely established by Doyle and O'Leary nia, each with an egg cell, more than one egg may
(1935). The role of this micropylar fluid has been be fertilized and develop into an embryo in a single
studied in detail by McWilliam (1958). The micro- seed. However, although seeds with two embryos
pyle of the anatropousovule consists of a chamber occur regularly,normallyonly a single embryodevel-
formedby the integumentextendingbeyond the apex ops (Fig. 8). At the time of fertilization, the seed
of the nucellus, a canal terminating in a narrow cone will have reached nearly full size but is still

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42 Flora Neotropica

"green"and the scales are tightly closed. The events which one succeeds and develops further into the
following fertilisationhave been described in detail final embryo. Cellular diversification leads to the
by Maheshwariand Konar (1971) and only a short formation of a rootcap followed by root initials
summaryof embryogenywill be given here. Mitotic (radicle)at the proximalend of the suspensor,cotyle-
divisions (4 X) result in the formationof a proem- dons at the distal end of the embryo and the hypo-
bryo with four tiers of four cells each (Fig. 8). Any cotyl in between these structures(Fig. 8). The fully
remaining nuclei (paranuclei)disappearduring this developed embryo is embedded in the enlarged fe-
stage. Of the four tiers of the proembryo,the two male gametophyte (often referredto as endosperm,
nearest the micropyle degenerate,while the suspen- but its ontogeny is entirelydifferentfrom angiosperm
sor elongates, thrustingthe terminalembryoniccells seed development). Both embryo and gametophyte
deeper into the gametophytic tissue. The proem- are enclosed in a thin chartaceous shell (all that
bryonalcells then divide into several proembryos,of remainsof the nucellus) and a hard seed coat.

PHYLOGENYAND EVOLUTION

GENERICRELATIONSHIPS popinus) has been acknowledged by most workers,

but few have had access to materialof this rare and
Pinus (Pinaceae) is considered monophyletic
endangered Vietnamese pine. Apart from its two
(Flous, 1936; Niemann& van Genderen, 1980; Price
peculiar, flat, sometimes hypostomaticleaves, it fits
et al., 1987; Farjon, 1990a); it is also the largest and
well in subgen. Strobus.
most diverse genus. Its sister group is probablythe
genus Picea, from the evidence of two types of
characters(chemicaland morphological)investigated
Further Divisions
in these studies. Intergeneric relationships among
the II (sometimes 10, 12, or 13) genera currently The subdivision of the two subgenerais consider-
recognizedare not well [Link] phyloge- ably morecontroversial,and a largenumberof classi-
netic studies involving genera in the Pinaceae reject fications has been proposed, dealing either with the
the traditionalclassificationbased on foliage charac- entire genus or with parts of it (see History of
ters as artificial (Niemann & van Genderen, 1980; Classification). Some, because they are based on a
Price et al., 1987; Frankis, 1989; Farjon, 1990a), but very limited numberof characterschosen for identi-
only the latter two, based primarilyon morphology, fication purposes (e.g., Pilger, 1926), are definitely
give Pinus a more basal position on the tree in a artificialand do not reflectphylogeneticrelationships
distinct [Link] using or evolution of [Link], though intention-
DNA and comparingthe genera are yet to be under- ally phylogenetic, seem to classify species in groups
taken. defined by unlikely combinations of homoplasies,
e.g., the reversal to deciduous fascicle sheaths in
some and the reversal to adnate seed wings in other
INTERGENERICRELATIONSHIPS
species of the same (sub)section (Shaw, 1914; Lit-
Subgenera tle & Critchfield, 1969). Phylogeneticanalysis based
on the requirementof monophyly and considering
It is widely accepted that there are two natural
only sharedderived (synapomorphous)charactersas
groups within the genus Pinus, which are often called indicative of common
ancestry (Hennig, 1966) has
Haploxylon and Diploxylon but which, under the called such classifications into
rules of nomenclature,must be named subgen. Stro- question (Strauss &
Doerksen, 1990; Malusa, 1992; Wang & Szmidt,
bus and subgen. Pinus, respectively (see History of
1993).
Classification). Morphological and anatomical
(Koehne, 1893; Shaw, 1914; Little & Critchfield,
1969; Farjon, 1984), chemical (Lindstedt, 1951), and
molecular characters (Strauss & Doerksen, 1990; Subgen. Pinus
Wang & Szmidt, 1993) have been analyzed,support- Of the 60 taxa covered by this treatment, 42
ing this basic division. Since Little and Critchfield belong to subgen. Pinus. Speciationin Mexican pines
(1969), the subgeneric distinction of P krempfii(or is consideredto be concentratedmainly in this subge-
even its recognition as a monotypic genus, Ducam- nus, with various "species complexes" being com-

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Phylogeny and Evolution 43

monly recognized (Martinez, 1948; Stead & Styles, paucity of discrete characters,which in the genus
1984; Squillace & Perry, 1992). Subsections (ac- Pitus are scarce when compared to other plant
cording to Little & Critchfield, 1969) with major groups (Styles, 1993a). A cladistic analysis of all
representationin Mexico and Central America in- taxa in this revision resulted in a strict consensus
clude Leiophyllae, Ponderosae, and Oocarpae; in tree with both unresolvedand resolved clades within
the Caribbean the taxa belong mainly to subsect. subgen. Pinus. With successive approximations
Australes. A refinement of Little and Critchfield's weighting, which uses the rescaledconsistency index
classification [which retained much of Shaw's to calculate weights, nearly all clades (trees not
scheme despite claims by others that it was based shown) were resolved. Due to the preliminarynature
largelyon Duffield's (1952) hybridizationtrials] was of the cladistic analysis, the incomplete sample of
given by van der Burgh(1973) and slightly amended taxa, and the high levels of homoplasy in most
by Farjon(1984). This classificationespecially subdi- characters,the analysis is used sparinglyto influence
vided Little and Critchfield'ssubsect. Ponderosaeby the classification. We therefore propose to present
splittingoff subsect. Pseudostrobi,and Oocarpae by the species of this subgenus tentativelyaccordingto
splitting off [Link] Leiophyllae was Little and Critchfield (1969) and van der Burgh
elevated in rank to a section, with two subsections (1973), with such emendations as are substantiated
each accommodating one or two species. Styles by the cladistic analysis of our data. The list of
(1993a) slightly modified Little and Critchfield's charactersand a data matrix used in the analysis
scheme as applied to Mexican taxa, with additions (using PAUP version 3.1) are presentedin Tables I
of species and varieties not recognized or known in and II, [Link] one cladogram(Fig. 12),
the originalclassification.A comprehensivephyloge- representinga phylogeny of species in subgen. Stro-
netic analysis of subgen. Pinus is hamperedby the bus, is presentedhere.

Table I
Character numbers and states used in the cladistic analysis of taxa in the genus Pinus. Characters 4, 5, 11
and 15 are represented by the first state only in species of subgen. Strobus; character 6 does not apply for
that subgenus (see Table II).

1. Winterbuds with resin, 0; without resin, I.

2. Sheaths of matureleaf fascicles early deciduous, 0; late deciduous, forming a rosette, 1; persistent,2.
3. Leaves epistomatic,0; amphistomatic,I.
4. Resin ducts (primarily)external, I; medial, 2; internal,3; septal, 4.
5. Vascularsystem in leaves haploxyl, 0; diploxyl, 1.
6. Vascularbundles in diploxyl pines connate, 0; separate, I.
7. Hypodermisuniform,0; of variablethickness, I.
8. Outercell walls of endodermisthin, 0; thickened, I.
9. Seed cones opening soon, deciduous, 0; serotinous,persistent, 1; opening slightly, disintegrating,deciduous, 2.
10. Peduncledeciduous with cone, 0; persisting with some basal scales, 1.
11. Apophyses in mid-cone nearly monomorph,0; strongly heteromorph,1.
12. Umbo terminal,0; dorsal, 1.
13. Umbo with deciduous spine, or spine absent, 0; spine persistent, 1.
14. Seed wing articulate,much larger than seed, I; seed wing adnate, vestigial, 2; seed wing absent from matureseed,
3; seed wing adnate, much largerthan seed, 4.
15. Long shoots uninodal,0; multinodal, 1.
16. Fascicle bract (cataphyll)bases virtuallynon-decurrent,0; decurrent,I.
17. Numberof leaves per fascicle 1(-2), 1; 2(-3), 2; (2-)3, 3; 3(-5), 4; 3-5, 5; 5, 6; 5-8, 7.
18. Sheaths of leaf fascicles initially long (>20mm), 1; of medium length (15-20mm), 2; short ( 15mm). 3.
19. Sheaths of leaf fascicles reduced in length at maturity(?30%), 0; persistentlylong, I.
20. Length of leaves -5cm, 1; 5-10cm, 2; 10-15cm, 3; 15-25cm, 4; 25-50cm, 5.
21. Leaves -I mm wide, rigid, erect, 1; 1-1.2 mm wide, lax, drooping or erect, 2; - I mm wide, soft, drooping or
pendent,3.
22. Peduncles of seed cones long (>2.5 cm), slender, 1; 1-2 cm long, stout, 2; very short (-I cm), 3 (for very large
cones multiply by 2).
23. Length of matureseed cones -5 cm, 1; 5-8(-10) cm, 2; (8-)10-15 cm, 3; (15-)18-25 cm. 4; -25 cm, 5.
24. Numberof seed scales per cone (average) -60, 0; 60-110, 1; 110-170, 2; >170, 3.
25. Numberof stomatal lines on abaxial face of leaves 2-5, 1; 6-10, 2; -10, 3.

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44 Flora Neotropica

Table II ayacahuite
Characterstate matrix for taxa in Pinus subgen. veitchii
Strobus,comparing20 characters(excluding lambertiana
characters4, 5, 6, 11 and 15, which are
reflexa
autapomorphiesof Pinus subgen. Pinus).
strobifonnis
Polymorphismand absence of charactersare
indicated by a question mark(?). chiapensis
pinceana
11111111222222
12378902346789012345 nelsonii
rzedowskii
P. strobus var. chiapensis 1000000004063?32131?
P laihbertiana 1010000004063?221521 maximartinezii
P flexilis var. reflera 00?0000002063?22231? lagunae
P strobiformis 10?0000002061?232???
P. vacahitte 1000000004062?332?2?
cembroides
P avtacahuite var. veitchii 1000000004062?33152? bicolor
P rzedowskii 1200000111053?23131? orizabensis
P pincelana 1010020113143'22121?
P nelsoniii 01100201131431221211 culminicola
P maximartinezii 0200020103063?23341? remola
P culhninicola 0200000113063? 11310?
P remote 1010100103023?113101 monophylla
P cembroides var. bicolor 1200000103043?123101 quadrifolia
P cembroides subsp. orizahensis 02 10000)103043?123101
Picea
P. cembroides subsp. lagunae 1210000113033?223101
P cemtroides subsp. cembroides 1210000113033?123101
P mnonophylla 0011000103013'?113103 Fig. 12. Strictconsensustreeof 24 equallyparsimonious
P. /uadlrifoiia trees(Cl = 0.586, numberof steps = 58), analyzing20
0000000 103053?113101
Picea chihualmana 07?0100??7?10'?? 11?322 morphological characters(see TablesI andII),usinga heu-
risticsearch(randomaddition;subtreepruning/regrafting)
withFitchparsimony (Swofford,1993).

[Link]
(LOUDON)VANDERBURGH: pringlei, classified in subsect. Oocarpae [Link]. by van
[Link] BURGH der Burgh, but in two different subsections by Little
Little and Critchfield(1969) classified Pinus leio- and Critchfield,appearin the cladistic analysis (strict
phylla (including [Link])and P lumiholtzii consensus tree) in a clade with P. leiophylla. When
as a subsection within sect. Pinea Endl., which con- charactersare weighted against homoplasy, this ar-
tained two other subsections of Mediterraneanpines. rangementis refuted (strict consensus of nine trees,
Van der Burgh (1973) recognized the distinctness of not shown). The fossil recordappearsto supportthe
these Mexican pines and accorded each of them the occurrence of pines related to P leiophyllacin the
rank of section. Morphologically, these pines are Late Paleogene of Colorado. In the fossils described
unique in subgen. Pinus in having deciduous fascicle by Axelrod (1986), no mention is made of the diag-
sheath scales. Recent isozyme analysis (Karalaman- nostic marksof three-seasonalgrowthon the apophy-
gala & Nickrent, 1989) and phylogenetic analysis sis of the seed scale, but if cones and leaf fascicles
using rDNA restriction fragments (Govindarajuet with lost sheaths indeed came from the same pine,
al., 1992) confirm the genetic distinctiveness of P P coloradensis is likely to be closely related.
leiophylla and show that it is ill-placed within sect.
Pinea as circumscribed by Little and Critchfield [Link]
(1969). This species is also distinguishedby its three- Van der Burgh (1973) restrictedsect. Pittus sensu
year development of the seed cones, which is shared Little and Critchfield (1969) to a group of pines
by the MediterraneanP pinea. Both charactersap- with a primarilyEurasiandistributionand only two
pear to be examples of parallelismratherthan syna- American representatives,one of them P. tropicalis.
pomorphies. Pinus lumholtzii appears in our clado- This species is anomalous within that group for
gram (not shown) near P pseudostrobus. Its having septal resin ducts in the leaves (Farjon, 1984).
relationshipdoes not seem to be with P leiophylla, The section also contains the Southeast Asian P
but unlike that species, its place in a distinct section nerkusii,which has an anomalouscone morphology
is not confirmed. Pinus teocote, P. lawsonii, and P and in a cladistic analysis based on cpDNA charac-

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Phylogeny and Evolution 45

ters emerges as a sister group to sect. Pinus (Wang& sifted species of western NorthAmerica and Mexico
Szmidt, 1993). Phylogenetic analysis comparing P to CentralAmericaof Shaw's (1914) groupAustrales
tropicalis with its putative relatives as well as with in an amendedsubsect. [Link] includes three
species with nearbyCaribbeanand SE NorthAmeri- commonly recognized "species complexes" around
can distributionsmay change its classification, but P ponderosa, P mnontezumae,and P pseudostrobus
such work could not be undertakenwithin the scope (Mirov, 1967). Vander Burgh(1973) segregatedtaxa
of this [Link] differs from other Caribbean considered to belong to the latter two groups in his
pines in its septal resin ducts (versus internal), its subsect. Pseudostrobi. Species delimitation within
uniform hypodermis, its low leaf number and its these groups is often difficult and, in part as a
very short cone peduncle. consequence of this, there is little agreement con-
cerning relationshipsof the constituenttaxa. Cladis-
[Link] tic analyses of all taxa in the region, as well as of a
This section as recognized by van der Burghcon- limited numberwithin subgen. Pinus, permit a few
tains the remaining species in subgen. Piiiius. Al- observationsabout their possible relationships:
though it represents fewer species than Little and I. Pinus montezutmaeand P pseudostrobusare spe-
Critchfield'ssect. [Link], which included the group
cies in a monophyleticgroup in most consensus
of pines mentioned in the previous section, it has
still some 40% of all the species in the genus. This trees, commonlyjoined by P devoniana, P doug-
lasiana, P lumiholtzii,and P miaximinoi.
reflects the difficulty in separatingthem into distinct
2. Pinus harvtegii, placed in subsect. Pseudostrobi
taxonomic units. Both Little and Critchfield(1969)
and van der Burgh (1973) have divided their largest by van der Burghand consideredto be a species
in the "montezumae complex" (Martinez, 1948:
section into 6-8 subsections, some of which seem to
Mirov, 1967) does not appearto be closely related
representnaturalgroups. to it.
Subsect. ContortaeLittle & Critc/ifield.- This is 3. Pinus engelinanniimay be relatedto P arizonica,
a subsection with a northerndistribution,restricted P ponderosa, and P jeffrevi, but also to a clade
by van der Burgh (1973) to Pintus contorta and formed by species mentionedunder 1 above.
P [Link] is representedby P contorta var.
There are a few Tertiaryfossils known that resem-
imurrayana,which extends along the Cascades-Si-
erra Nevada axis and has a southern disjunction in ble cones and seeds of different extant species as-
to these subsections, most of them from the
our region (SierraSan PedroMartir,BajaCalifornia). signed
Rocky Mountains northof the U.S.-Mexican border
Pilius herrerae,placed in subsect. Oocarpae [Link]
van der Burgh,does not seem to fit therecomfortably. and dating from the Eocene to the Miocene (Axelrod,
1986). Large cones resemblingthose of P montezu-
Pirus contorta var. mulrravanac could be related to
this pine with very small cones, as the cladistic inae from the fossil Creede flora (Late Oligocene,
26.5 Ma) of Colorado, and others from Miocene
analysis [Link] fossils are scarce, P alvor-
densis Axelrod from the Late Eocene Bull Run flora (12.5 Ma) deposits in Western Nevada resembling
of northernNevada is believed to belong here (Axel- those of P pseudostrobus, indicate that these or
relatedspecies occurredconsiderablyfarther
rod, 1986). We tentatively group these two species closely
the need for furtherresearch into norththan they do at present.
together,stressing
their relationships. [Link]& Critchfield,Attenuatae
-
Subsect. Australes Loudon. In both classifica- van der Burgh, and SabinianaeLoudon.- As pro-
Little and Critchfield(1969), subsect. Ooc-
tions, this subsectioncomprisesspecies rangingfrom posed by
the eastern and southeasternUnited States through arpae includes the Mexican and Californian"closed-
the Caribbean to Central America. In our region, cone" pines. Van der Burgh (1973) divided this
these are P elliottii var. densa, P caribaea with its subsection into two subsections, restrictingsubsect.
threevarieties,P cubensis, and P [Link] Oocarpae to the Mexican-Central American pines
and adding species to both. His subsect. Attenuatae
synapomorphyis a multinodallong shoot (not in P
The is at consists of P muricata, P radiata, P attenuata, P
occidentalis). group poorly, if all, repre-
sented in the fossil record. greggii, and P. patula. Cladistic analysis revealed
that Little and Critchfield's subsection is paraphy-
Subsects. Ponderosae Lotuldon and Pseudostrobi letic. The first three species of van der Burgh's
van der Burgh.- Little and Critchfield(1969) clas- subsect. Attenuatae form a clade with P greggii

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46 Flora Neotropica

and/or P coulteri (trees not shown). Millar (1986a) either a pair of closely related species (Perry, 1991;
considered the only truly naturalgroup in subsect. Perez de la Rosa, 1993) or a single species (Carva-
Oocarpae sensu Little and Critchfieldto be the one jal & McVaugh, 1992). Pit2uslambertiana is more
that comprises the three Californian closed-cone closely related to P ayacahuite in this cladogram.
pines. Pinus coulteri is classified in both systems in The homogeneity within subsect. Strobi relative to
subsect. Sabinianae,togetherwith two (or one) other the subgenus as a whole seems to indicate that these
Californian pines with large cones and prolonged species have radiatedrelatively recently, yet fossils
apophyses. It shares the characters Millar (1986a) very similarto P lambertianahave been found in the
mentionsas distinctfor the closed-cone pines: medial Middle Eocene nearSan Diego, California(Axelrod,
resin ducts, multinodal long shoots, and armed, 1986). Interestingly,species of this subsection are
asymmetrical,serotinouscones. The cladistic analy- rare in Mexico and Central America. Pinus strobi-
sis does indicate,contraryto Millar's (1986a) objec- formis representsthe northernelement, P ayacahuite
tions, the existence of two groups similar to van the southernelement in highland pine forests, while
der Burgh's classification, with P coulteri possibly each is accompaniedby numerousspecies in subgen.
related to the Californianclosed-cone pines. Pinus Pinus (see also Mirov, 1967). In all, six taxa of
greggii shares clustered, asymmetrical, serotinous subsect. Strobioccur in our region, two of them only
cones, multinodallong shoots, and medial resin ducts just south of the U.S.-Mexican border. It seems
with the Californianclosed-cone pines. The position that this subsection originated elsewhere, either in
of P patula is less clear. Pinus durangensis, P northernNorth America or, perhaps more likely, in
jaliscana, P. praetermissa, and P tecunumanii,and Asia. No species of subsect. Cembraeoccurs in our
more distantly, P oocarpa and P pringlei, seem to region, and only one (P albicaulis) occurs in the
be related to it. This outcome gives some support United States and Canada and is strikingly similar
for an amended subsect. Oocarpae. Pinus lawsonii, in its seed cones to the Eurasian species-pair P
P pringlei, and P teocote are provisionally retained cembra and P sibirica. For both subsections, an
in it. The closed-cone pines ([Link])are early Tertiaryorigin is supportedby palaeobotanical
abundantlyrepresentedin the fossil record(Axelrod, evidence (Axelrod, 1986).
1980, 1986). Most are from the Pleistocene of Cali-
fornia and resemble or belong to extant taxa of the SECT. PARRYANAE MAYR
region, but some are older and date as far back as Nearly all recentcommentatorson pine phylogeny
the Miocene. and classification agree that Pinus sect. Parryanae
Mayr,as interpretedby Little and Critchfield(1969),
is extremely heterogeneous,more so than any other
Subgen. Strobus (Sweet ex Spach) Lemmon section commonly recognized in the genus (van der
Burgh, 1973; Klaus, 1980; Farjon, 1984; Strauss &
[Link] SPACH,[Link] Doerksen, 1990; Malusa, 1992; Styles, 1993a). Only
LOUDON one of its threesubsections,Cembroidess.l., is repre-
Recent cladistic analyses of morphologicalcharac- sented in our region,but the heterogeneitywithin this
ters (Malusa, 1992; Farjon & Styles, this volume) subgroupis nearly as great. Phylogenetic analysis of
and of molecular characters (Strauss & Doerksen, both morphological and molecular characters(Ma-
1990; Govindarajuet al., 1992; Wang & Szmidt, lusa, 1992; Strauss & Doerksen, 1990; Wang &
1993; Perez de la Rosa et al., 1995) agree on the Szmidt, 1993; P6rezde la Rosa et al., 1995; Farjon&
monophyly of sect. Strobus and the paraphyly of Styles, this volume) gives evidence for paraphylyin
sect. Parryanaewithin the [Link] sect. Parryanae,indicatingthat it is unlikely to be a
the section Strobus, the subsections Cembrae and naturalgroup. Malusa (1992), using Wagner parsi-
Strobi are very similar (Strauss & Doerksen, 1990) mony analysis of mainly morphological characters
and either monophyletic (Govindarajuet al., 1992; for all species in the section, found a clade consisting
Wang & Szmidt, 1993, who looked at Asiatic spe- of P. maximartinezii,P rzedowskii, P nelsonii, P
cies) or unresolved/paraphyletic(Strauss & Doerk- pinceana, and the Asiatic pair P gerardiana and P
sen, 1990). The strict consensus tree using Fitch bungeanato form a sister group to the other "pinyon
parsimonyanalysis of morphologicalcharacters(Fig. pines" of the traditionalsubsect. [Link]
12), thoughcomparingonly a limited numberof taxa de la Rosa et al. (1995) found P. nelsonii to be very
in subsect. Strobi, separates Pinus ayacahuite from distinct in that it possessed neitherof the two length
P strobiformis,two taxa commonly consideredto be mutationsfound in the cpDNA of 11 other Mexican

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Phylogeny and Evolution 47

species of Pinus representingboth [Link] lb. P leiophylla var. chihuahuana

pinceana and P. maximartineziishare one synapo- Sect. Pinus
morphy in their cpDNA, but none with P cem- 2. P tropicalis
[Link] strictconsensus tree found in our analy- Sect. Pinaster
sis (Fig. 12) seems to confirm Malusa's findings Subsect. Contortae
which place these pines clearly outside a group with 3. P. contorta var. murravana
P cembroides(subsect. [Link].).This tree 4. P. herrerae
supportsthe theory that the four Mexican species P Subsect. Australes
maximartinezii, P rzedowskii, P nelsonii, and P 5a. P caribaea var. caribaea
pinceana are relict taxa, today with (very) limited 5b. P caribaea var. bahamensis
and probably naturallydecreasing distributionsand Sc. P caribaea var. hondurensis
populations. They very likely had relatives in the 6. P elliottii var. densa
geologic past which became extinct. Their treatment 7. P cubensis
in the same section or subsection with P cembroides 8. P occidentalis
and its allies has no phylogenetic basis. Pinus nel- Subsect. Ponderosae
sonii has a numberof autapomorphies(see System- 9. P ponderosa var. scopulorum
atic Treatment)that set it apart;a separatesubsection O1a.P arizonica var. arizonica
has been proposed for it (Carvajal, 1986). Pinus lOb. P arizonica var. cooperi
nelsonii and P pinceana appear to be united on a IOc. P arizonica var. stormiae
clade in the strictconsensus tree (Malusa, 1992; Fig. I 1. P jeffreyi
12). As these two species also form a pair of related 12. P engelmannii
terminal taxa in all trees generated from our data, (?) 13. P hartwegii
we proposeto include P pinceana in that subsection. Subsect. Pseudostrobi
Pinus rzedowskii has a seed cone quite unlike any 14a. P pseudostrobus var. pseudostrobus
"pinyon pine"; it could representa more primitive 14b. P pseudostrobus var. apulcensis
cone type. It has also seeds with large, effective 15a. P. montezumae var. montezumae
wings, which almost certainly is the plesiomorphic 15b. P mnontezumaevar. gordoniana
state in pines. Fossil evidence is as yet of little help: 16. P devoniana
almost all of the materialthat is known from fossils 17. P douglasiana
with resemblanceto subsect. Cembroidess.l. seems 18. P maximinoi
closest to P cembroidesor P edulis. Axelrod (1986) (?) 19. P lumholtzii
discusses the possibility that the fossil P lindgrenii Subsect. Oocarpae
Knowlton,from the Late Miocene in southernIdaho, 20a. P. oocarpa var. oocarpa
is intermediatebetween these and P pinceana. Pend- 20b. P. oocarpa var. trifoliata
ing a comprehensive study of morphological and 21. P praetermissa
molecular characters,it seems most appropriateto 22a. P patula var. patula
recognize subsect. Cembroidesin a strictsense in this 22b. P patula var. longipedunculata
monograph,to recognize subsect. Nelsoniae Carvajal 23. P jaliscana
with two species, and to list the two remaining 24. P tecunumaniii
aberrantspecies as unclassifiedwithin subgen. Stro- (?) 25. P durangensis
bus. Section Parryanae is marked with "?" in the (?) 26. P lawsonii
classificationbelow, to indicate its paraphyly. (?) 27. P pringlei
Based on the considerations outlined above, the (?) 28. P teocote
taxa of the genus Pinus accepted in this monograph Subsect. Attenuatae
are provisionallyclassified; it must be stressed that 29. P muricata var. muricata
the ranking presented below does not necessarily 30. P radiata var. binata
follow from the [Link] subsec- 31. P attenuata
tions neverthelessformed lower-orderclades within 32. P. greggii
the consensus trees. (?) 33. P coulteri
Pinus subgen. Strobus
Pinus subgen. Pinus Sect. Strobus
Sect. Leiophyllae Subsect. Strobi
la. P leiophylla var. leiophylla 34a. P avacahuite var. avacahuite

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48 Flora Neotropica

34b. P ayacahuite var. veitchii and resin ducts in the bract-scalecomplexes. Despite
35. P lIambertiana these peculiarities of their anatomy (how accurate
36. P flexilis var. reflexa are these observationsin each case?), several of these
37. P strobiformis cones probably represent fossil species of Piiis.
38. P strobus var. chiapensis These fossils date from the Lower Cretaceous(Weal-
(?) Sect. Parrvanae den, ca. 125 Ma) as well as from youngerCretaceous
(?) 39. P rzedowskii to Palaeocene (Axelrod, 1986) formations;they sug-
(?) 40. P maximartinezuii gest that Pinus may have a central place in the
Subsect. Nelsoniae phylogeny of the family (Miller, 1976, 1977). Older
41. P nelsonii pinaceous cones (e.g., ComisostrobusDelevoryas &
42. P piniceana Hope, from the Late Triassic of North Carolina)
Subsect. Cembroides are as yet of more doubtful affinity, due mainly to
43al. P cembroides subsp. cembroides imperfectpreservationof these remains.
var. cemlbroides Unambiguousevidence of macrofossilsdates back
43a2. P cembroides subsp. cembroides to the Lower Cretaceous(Miller, 1976, 1977; Millar,
var. bicolor 1993). Structurallypreserved ovulate cones of the
43b. P. cembroidessubsp. lagunae genus (Pilnus belgica Alvin) are known from the
43c. P cembroides subsp. orizabensis Wealden (Neocomian) of Belgium; several other
44. P culminicola early species date from the Lower Cretaceousto the
45. P remota Eocene/[Link] all appearto be members
46. P monophyllla of subgen. Pinu.s,having small, conical cones with
47. P quadrifolia dorsal (abaxial) umbos (Miller, 1976, 1977), which
are comparable to the modern cones of subsects.
Contortae, Oocarpae, and Pinus (syn. Svlvestres).
EVOLUTION
Cretaceousfossils of anatomicallypreservedsecond-
The Mesozoic, lasting from ca. 225 Ma to ca. 65 ary leaves reveal diploxyl vascularizationin accor-
Ma, was the era of conifers; there were many more dance with subgen. Pinus (Miller, 1977). The only
forms than there are at the presenttime, and most of evidence for subgen. Strobusto have occurredin the
these became extinct. The evolution of the modern Cretaceous is a cone described as P. magothensis
conifer families-e.g., Araucariaceae,Cupressaceae, Penny from Delaware,of which the original material
Pinaceae, and Podocarpaceae-has been interpreted has been lost (Miller, 1977). Similarly described
as a radiation from the Permian-MesozoicVoltzia- cones have more recently been reassigned to the
ceae (Voltziales),which in turnevolved from Palaeo- organ genus Pityostrobus.
zoic Lebachiaceae(Florin, 1951, 1954; Miller, 1982). The origin and early radiation of pines, being
However, there is cladistic evidence that different xerophytes, is likely to have occurred in relation to
genera in Voltziaceaewere possibly ancestralto dif- increased seasonality, especially aridity, of which
ferent modernconifer families, one clade leading to there is evidence in the Early Cretaceous (Mirov,
Cupressaceae,anotherto the other families, and that 1967; Axelrod, 1986). By the Middle Eocene, a
some genera were not ancestralto any extantconifers majorshift to the spreadingof dry climate in south-
(Miller, 1988). The Pinaceae must have evolved westernNorthAmericacreatedconditions favourable
from a common pinaceous ancestorsometime in the to furtherspeciationin Piinus(Axelrod, 1986; Millar,
Jurassic, as several distinct lineages, among them 1993).
Pinus, were already present at the beginning of the Shaw (1914) based his classification of the pines
[Link] organ genus Pityostrobus(Nathorst) on "the gradual evolution of the fruit from a cone
Dutt possibly representsseveral genera of pinaceous symmetricalin form, parenchymatousin tissue, inde-
structurallypreserved ovuliferous cones belonging hiscent and deciduousat maturity,releasing its wing-
to Pinaceae. More than 20 species have been de- less seed by disintegration[subsect. Cembrae]-to a
scribed so far. Almost all of these fossils have some cone oblique in form, very strong and durable in
characters assigned to Pinus but also have other tissue, persistenton the tree, intermittentlydehiscent,
features not known in modern pines (Miller, 1976). releasing its winged seeds partly at maturity,partly
The thickened scale apex (apophysis?) is the only at indefiniteintervalsduring several years." He con-
morphological character;the others are anatomical cluded: "Thisevolution embracestwo extreme forms
and relate to the position of vascularstrandsor traces of fruit, one the most primitive, the other the most

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Distributionand Ecology 49

elaborate,among conifers."When the cone morphol- in a part of subgen. [Link] "typical"Strobus

ogy of Pinus is compared with the other genera in cone with a terminalumbo is then the derived state.
Pinaceae, neitherof these types occur outside Piinis. The Cembrae cone, with nearly indehiscent scales
The "plesiomorphic"type would seem to be the and vestigial seed wings, is a furtherspecialization
pendulous, deciduous, and more or less cylindrical from this lineage, in coevolution with seed dispersal
cone with relatively thin, freely parting seed scales by birds. Serotinouscones with asymmetricaldevel-
releasing seeds with adnate wings. Secondly, phylo- opment of apophyses are derived from the ancestral
genetic analysis of many charactersclearly indicates type. Their evolution is probably linked with more
that the species of subsects. Cembrae and Strobi extreme conditions of climate, but defense against
have evolved ratherlaterthanthose with "less primi- predationmay also have played a part. Most species
tive" cone types (Fig. 12; see also Strauss& Doerk- with such cones occur at present in regions with
sen, 1990; Wang & Szmidt, 1993). We now know climatic extremes, especially prolongeddrought.
that Shaw's "most primitive"cone type in reality is Evolutionarytrendsin foliage charactersare diffi-
the outcome of coevolution involving seed dispersal cult to [Link] number is highly variable
by corvid birds, e.g., the relationship between P in most groups; it is most consistent in subsects.
cembra and Niucifraga(carvocatactes in the Alps and Pinus ([Link].)and [Link] is a very general trend
similarmutualismsbetween nutcrackersandjays and toward increased numbers from boreal to tropical
P albicaulis and P edulis in the United States North America, apparentlyrelated to climate. Pinus
(Mattes, 1994;Tomback& Linhart,1990). The serot- nelsonii is exceptional within subgen. Strobus in
inous cones (in several species of subgen. Pinuss)are having fascicles enclosed in persistentsheaths. This
indeed a resultof "elaborateevolution,"but no more condition and a variable (low?) number of leaves
so than those of subsect. Cem7brae. The admittedly may representthe primitive state. The fossil record
scant fossil recordfrom the Mesozoic gives evidence from the Cretaceous is too fragmentary to draw
contrary to Shaw's ideas on pine cone evolution conclusions, yet the earliest pines known were of
(Miller, 1977). From the Tertiaryof NorthAmerica, the few-leaved, diploxyl type (Miller, 1976, 1977).
no fossils have been ascribed to subsect. Cembrae Convergence or parallelism, such as the deciduous
(Axelrod, 1986); from Eurasia there is fossil evi- fascicle sheaths of P. leiophylla and P. [Link]. or
dence from the Palaeogene only (Gaussen, 1960). the "wingless"seeds in the MediterraneanP pinea,
Given the probability of monophyly of the two seems to have occurredoccasionally. However, until
subgenera, the ancestral cone form of Piinuswould we know in more detail how these characters
have had apophyses with a dorsal umbo, a condition evolved, we cannot be entirely certain of their ho-
which now prevails in subgen. Piinusand is present mology.

DISTRIBUTION AND ECOLOGY

DISTRIBUTION (for P ctiribaea var. bahllmensis in the Bahamas) to

4000 m (for P hairtwegiiin Mexico). Other genera
The genus Pinus is the most widespreadgenus of in the fanily (Abies, Picea) cover similar or even
Pinaceae,extending across the NorthernHemisphere greater altitudinal ranges, but their low-elevation
roughly between latitudes 15?N(-02?S) and 66? species occur only in northernlatitudes,as compared
(-70?)N in NorthAmericaand Eurasia(Fig. 13). The to several low-elevation pines in the tropics. Even
most southerly occurrence in the Western Hemi- though the altitudinalranges of these genera reflect
sphere is at 12?N in Nicaragua, where Caribbean their latitudinalranges, as both are correlatedwith
coastal populationsof Pinus caribaea var. hondur- climate, the pines can be said to have attained the
ensis reach as far south as Bluefelds (LUckhoff, [Link] ad-
1964). As with other genera in the family, species aptations(see Ecology, below), resulting in species
distributionsare very large in the boreal zone but radiation,have allowed the pines to spread into the
species diversity is low; the distributionsare reduced warmerand drierenvironmentsof southernlatitudes
towardthe oceanic coasts and in the southernmon- and lowlands.
tane regions, and species diversity increases(Farjon, Unlike the other medium-sizedto large genera in
1990a).Altitudinaldistributionranges from sea level Pinaceae, species diversity in Piins is greater in

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50 Flora Neotropica

FIG. 13. General distributionof the genus Pinus in North America and Eurasia, showing gaps in midwestern North
America and SW and CentralAsia.

North America than in [Link],this the 47 species and 20 infraspecific taxa recognized
coincides with a more complete occupation of the in this monograph,42 species and 18 infraspecific
North American continent as compared to Eurasia. taxa occur in Mexico (all taxa in Central America
In North America, the genus is totally absent only are also in the country of Mexico, including P
in the Arctic, on the Great Plains, and in scattered caribaea var. hondurensis; see, e.g., Eguiluz, 1982;
deserts. In Eurasia,it is absent in the greaterpart of Styles & Hughes, 1983). Five species and two varie-
Central,Southwest, and South Asia, with only nota- ties occur in the Caribbean,one of them, P. elliottii
ble exceptions in the Turco-Caucasianregion and the var. densa, on the Florida Keys and mainlandcoast.
Himalayas. While pines occur in many areas within Nine taxa are Californian;one of these, P radiata
the deciduous forest zone of the eastern United var. binata, is an island endemic that does not occur
States, they are virtually absent in the equivalent, in Californiabut has strong affinities with P radiata
but impoverished,zone in Europe. In the subtropics var. radiata, which is confined to that state. The
and tropics of the Western Hemisphere, pines like- coastal and insular Californian pines are relicts of
wise occupy an almost continuous belt and more a Madro-Tertiaryflora confined to fog-influenced
groundthan in SoutheastAsia, where they are mostly maritime and disjunct distributions resulting from
scattered. Three North American regions of high the northwestwardspreadof the Sonoranand Mojave
species diversity in the genus Pinus can be recog- Deserts (Axelrod, 1976, 1980).
nized: I) Mexico, 2) California,and 3) southeastern Rzedowski (1993) mentions the genus Pitius
United States. These are here defined in a phytogeo- among the genera for which (parts of) Mexico pro-
graphical sense: Mexico extends into the United vided centresof [Link] other possible source
States north to the Mogollon Rim in Arizona and of species diversity in a given region is migration.
New Mexico, and into CentralAmerica as far as the In order to evaluate the relative importanceof each
distributions of pines are concerned to Nicaragua; of these factors,it is necessary to look at the distribu-
California includes parts of Oregon, Nevada, and tion of pines on the North American continent and
northernBaja California. Rzedowski (1993) recog- its environs as a whole. The species that extend into
nises an extended Mexican region (Mega-Mexico 3) Mega-Mexico 3 [as defined by Rzedowski (1993)]
in a similar sense to define Mexican endemism. Of have distributionsthat can be grouped into six re-

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Distribution and Ecology 51

lo,

. :/..F` j -t ( I ^-/^ p<'^ z 'Ji ..

. 1?. -o

--.-
? -1"- . s
_.../
-_

...../ . V

FIG. 14. Regions of North American pine species extending into Mexico and the SE United States-Caribbean-Central
American connection. A. Californianregion. B. Great Basin region. C. Rocky Mountainregion. D. N Mexican region.
E. Mexican
E. Mexican Gulf-Atlantic
Gulf-Atlantic coastal
coastal plain
plain region. F. Caribbean
region. F. Caribbeanregion.
region.

gions (Fig. 14). Two of these-the Californianregion From there, a third variety (P caribaea var. hondur-
(A) and the Mexican Gulf-Atlantic coastal plain ensis) has somehow made it across the Cayman
region (E)-are North American centres of pine di- Trough-perhaps its seeds were carried by hurri-
versity, as is Mexico itself. From neither of them canes.
have species migratedvery far northinto Mexico. In The much less diverse Rocky Mountain pine re-
BajaCalifornia,the only truly migratoryspecies may gion (C) has been of more significance in the spread
well be the high-elevationones, i.e., P contorta var. of pines into greaterMexico. Two taxa with a north-
murrayana,[Link], and P lambertiana,which are erly distribution(P flexilis var. reflexa and P pon-
distributedon the higher portions of the block-fault derosa var. scopulorum) reach into the country of
zone that runs from here far into [Link] uplift Mexico, and several of the pines restricted to the
started in the Late Tertiaryand still continues. The northernMexican region (D) are closely related to
pinyon pines of Baja California (P monophylla, P species occurring in C and are derived either from
quadrifolia), as well as P coulteri, occur at lower these or from a common ancestor. These are P
elevations and may have spread from their centres arizonica (with 3 varieties), P durangensis,P engel-
of origin following the desertificationof the south- mannii, P flexilis var. reflexa, and P strobiformis.
western partof the continent. Pinus monophyllais a The pinyon pines common to these two adjacent
Great Basin element, and the other two are Califor- regions-P. cembroides var. cembroides. P cern-
nian. In the southeastern United States there is an broides var. bicolor, P edulis, and P remota-are
abundanceof pine species, six of which occur in the constituentsof the widespreadpinyon-juniperwood-
lowlands of Florida. Only P elliottii var. densa ex- land ecosystem, in which the minor diversification
tends a short way into the Caribbean;the closely of the species as well as the coevolution with corvid
related P caribaea occurs with two varieties in the birds related to seed dispersal strategies (see Ecol-
lowland pine savannas of the Caribbean islands. ogy) point to a relatively recent radiation related

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52 Flora Neotropica

CARIBBEAN AMERICA

II? ??)~?? ?~ , ,' . . ' ~:_; ...i

FIG 15 Nmesotaa(pcs, suseis n aite)i hisi tesae [Link]

counries
an Caiba islnds

FIG. 15. Numbers of taxa (species, subspecies, and varieties) in [Link] the states of Mexico. Central American
countries, and Caribbeanislands.

to climatic change. The common ancestor of this Mountainregion. Anotherquite different component
monophyletic group of pines (see Phylogeny and is local endemism, with species such as P culmini-
Evolution) probably originated somewhere in the cola, P nelsonii, and P pi,nceana. As noted above,
Mexico-United States borderarea. this region is also the most likely centre of origin of
Eguiluz (1985) recognized six principalregions of the species of Pinus subsect. Cembroides. Toward
distributionof Mexican pines. These regions with the southeast,diversity decreases, with the numberof
majordiversity are I) the Sierra Madre Occidental, taxa in Chiapas (along with Oaxaca. the botanically
2) the Sierra Madre Oriental, 3) the Eje Volcdnico richest state of Mexico; see, e.g.. Graham, 1993)
Transversaland Mesa Central, 4) the Sierra Juarez alreadysignificantlyless, until the distributionof the
and Sierra San Pedro Martir in Baja California, 5) pines reaches its terminationwith only four species
the Sierra Madre del Sur and Macizo de Oaxaca, in Nicaragua. Many of these southern pines are
and 6) the highlands of Chiapas, which extend into widespreadspecies, pointingto migrationas the most
[Link] were termed "centres of species likely cause of pine diversity in [Link]
diversity of pines in Mexico" by Styles (1993a). The of the Eje VolcanicoTransversal,local endemism of
Sierra Judrez and Sierra San Pedro Martir in Baja pines is absent, in line with the scarcity of local
California, as mentioned above, have only Califor- endemics in most other plant groups (Rzedowski,
nian pines. Mappingthe numberof taxa in each state 1993). This, too, is an indication that speciation
of Mexico roughly confirms Eguiluz's regions, but has not occurred in southern Mexico and Central
the Mesa Central, being a distinct morphotectonic America, possibly with the exception of P tecunu-
province (Ferrusqufa-Villafranca,1993) is conspicu- manii, which may be a relatively recently derived
ously less diverse than the Eje VolcanicoTransversal taxon.
(Fig. 15). High diversity in northeasternMexico is From this evidence, two centres of pine diversity
partly explained by past migrationsfrom the Rocky anldevolutionemerge: the Eje VolcdnicoTransversal,

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Uses and Conservation 53

with extensions to the northwestand southeastalong 1. lowland [1-300(-700) ml

the Sierra Madre Occidental and Sierra Madre del 2. foothills [(100-)300-1200 m]
Sur, and northeasternMexico. Here we find 14-18 3. montane [(700-)1000-2600(-2800) ml
taxa in most states (only the very small states around 4. high montane [(2000-)2500-4000(-4300) ml
Mexico City have fewer taxa, as expected). The
species of the first centre are almost all Mexican In Table III all taxa have been tabulatedagainst
taxa sensu stricto. This centre is also the major these two [Link] region I, the Californian
area for the three polymorphicspecies complexes P taxa are predominantlymontane,but two taxa occur
devoniana, P monntezumae, and P pseudostroblts(for at lower elevations. Their altitudes have been given
the first two see also Styles, 1993), which may have only for Mexico here, but these do not greatly differ
radiated from here over much of montane Mexico from the distributionsfound in California.
and CentralAmerica. The geological history of two In northwesternMexico (region II), most distribu-
of its main mountainchains is similar (Ferrusquia- tions are confined to the Sierra Madre Occidental.
Villafranca,1993). The SierraMadreOccidentaland Nine taxa are montane and four are high montane.
the Eje VolcanicoTransversalare both volcanic com- Most of these populationsoccur some distance from
plexes-the first with a long history of volcanism the coast, and the lower and middle elevations tend
beginning in the Upper Cretaceous and encom- to be much drier than similar altitudes nearer the
passing the Tertiary,the latterof more recent origin. coast, especially on the continentalside of the Sierra
The SierraMadredel Sur has a much more complex Madre. Here, the drought-resistantP cembroides
geology, but (high) mountainshave existed here at formsan intermittentbelt below the mainpine forests
least since the early [Link] second centre of (Perry, 1991). Pinus cembroides var. bicolor occurs
pine diversity and evolution is characterizedby 1) at higher altitudes but in dry regions with cold
adaptationto aridity, 2) disjunctionon high "island winters.
mountainranges" surroundedby (semi-)desert, and WesternMexican species (region III) have distri-
3) local endemism. Speciation as well as migration butions predominantlyin the southernSierra Madre
from the Rocky Mountainregion are the main causes Occidentaland westernmostpartof the Eje Volcanico
of pine diversity here. A more ancient geological Transversal,partly overlappingwith those of north-
historyof this partof Mexico-with Late Cretaceous western Mexican taxa. Most taxa in this region are
to Early Tertiaryuplift of mostly Mesozoic marine montane; the higher precipitationlevels allow pine
sediments, forming mountain ranges separated by forest to grow at lower [Link] is particularly
intermontanebasins (Ferrusquia-Villafranca,1993), evident on the oceanic slopes of the mountainranges.
and climatic change involving desertificationin the Here we find the greatest diversity of pines in
Miocene (Millar, 1993)-mostly explains these pat- Mexico.
terns. In northeasternand eastern Mexico (region IV),
A somewhat different approachis to analyze the distributionsare concentrated in the Sierra Madre
distributions, both geographical and altitudinal, of Oriental,with extensions to the northwestand in the
all taxa and classify these. Altitudinal ranges have easternmost part of the Eje VolcainicoTransversal.
been computed from the specimen database for this Most taxa are montane;several are scatteredalong
monograph; for altitudinal ranges of species oc- an intermittentbelt below the main pine forests on
curring both in Mexico and the United States, see the continental side of the Sierras, e.g., P c-em-
also Yeaton (1982). The geographical distributions broides, P nelsonii. and P pinceana.
(see maps in SystematicTreatment)can be classified A few species are restrictedto southern Mexico
into seven regions: (region V): P lawsonii, [Link], and P strobusvar.
1. California chiapensis, the latterextending into the highlandsof
[Link] occupy montaneregions in a sec-
II. northwesternMexico
tion of Mexico where the distance to the coasts is
III. western Mexico
much reduced, with the ameliorating influence on
IV. northeastern& eastern Mexico
climate as mentionedabove.
V. central & southernMexico
Othertaxa in centraland southernMexico (region
VI. Mesoamerica
V) have very wide [Link] of them have
VII. Caribbean
very wide altitudinalranges, too. These are due in
The altitudinalranges can be classified into four part to an upward shift of the lower (and upper)
categories: altitudinallimits from southeast to northwestgoing

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Table III
Distribution of Latin American pine species and infraspecific taxa according to geographical regions (I-

I (California) II (NW Mexico) III (W Mexico) IV (NE & E Mexico) V (C & S M

I (lowland: P. muricata
1-300[-700] m)

2 (foothills: P. attenuata P. devonian

[100-]300-1200 m) P. maximino
P. oocarpa

3 (montane: P. contorta var. P. arizonica + var. P. cembroides P. arizonica var. P. cembroid

[700- 1000-2600 nurravana cooperi subsp. lagunae* stormiae P. devonian
[-2800] m) P. coulteri P. cembroides P. devoniana P. cembroides + P. douglasia
P. jeffreyi P. devoniana P. douglasiana subsp. oriza- P. herrerae
P. lambertiana P. engelmannii P. durangensis bensis* P. lawsonii
P. monophylla P. leiophvlla var. chi- P. herrerae P. devoniana P. leiophylla
P. quadrifolia huahuana P. jaliscana* P. engelmannii leiophylla
P. radiata var. P. leiophylla var. lei- P. Ieiophylla var. P. greggii P. maximino
binata* ophylla chihuahuana P. leiophylla var. lei- P. montezum
P. oocarpa P. leiophylla var. ophylla var. gor-
P. teocote leiophylla P. montezumae doniana
P. lumholtzii P. nelsonii* P. oocarpa +
P. maximartinezii* P. patula + variety P. patula + v
P. maximinoi P. pinceana P. pringlei
P. montezumae P. pseudostrobus P. pseudostr
P. oocarpa P. remota + var. apu
P. praetermissa P. teocote censis
P. pseudostrobus + P. strobus va
var. apulcensis chiapensis
P. rzedowskii* P. tecunuma
P. teocote P. teocote

4 (high montane: P. cembroides var. P. hartwegii P. avacahuite + var. P. avacahuit

[2000-]2500-4000 bicolor P. montezumae veitchii P. hartwegii
[-43001 m) P. flexilis var. reflexa P. pseudostrobus + P. culminicola* P. montezum
P. strobiformnis var. apulcensis P. flexilis var. re- P. pseudostr
P. teocote P. strobiformis flexa + var. apu
P. teocore P. hartwegii censis
P. montezumnae P. teocote
P. pseudostrobus
P. teocote
P. strobiformis
* Narrow endemic taxa; for furtherexplanation, see text.

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Uses and Conservation 55

from Mesoamerica further into Mexico. With the they form the dominant forest types northwardas
widening of the Mexican landmass, interior forests well as at higher altitudes in mid-latitudemountain
are blocked from oceanic weather and only receive ranges. The duration of the growing season (the
sufficient precipitation at higher altitudes. Longer periodwith meandaily temperature> 10?Cfor conif-
summer day length away from the equator enables erous trees; Schmidt-Vogt, 1977) shortens progres-
trees to reach higher altitudes as growing-season sively with increasing latitude and altitude, until it
temperaturesare higher, and they can become dor- reaches the 10?C July-isotherm (Northern Hemi-
mant in the colder season. However, even within sphere) which is the climatic tree line. At the other
a relatively restricted area the range can be very end of the temperaturedine, two factors limit suc-
considerable,e.g., P nmontezumae is found in Oaxaca cessful growth of conifers: droughtand competition
between 1200 and 3000 m. Most of these widespread by broad-leaved trees. Moisture stress is usually
species are morphologically highly variable, with related to lack of precipitationat critical periods of
several varieties being recognized here. Pinus hart- the life cycle, e.g., the seedling and sapling stages.
wegii is found at the tree line, to an upper limit of Competitionby broad-leavedtrees occurs undercon-
ca. 4000 m, on the volcanos of the Eje Volcanico ditions where limitationsto their success are absent
Transversalwhich rise above 5000 m. Yet this spe- and one or more potentially successful species are
cies occurs down to 2300-2500 m in the highlands [Link] means that an undisturbedsuccession
of Oaxaca and elsewhere. of the vegetation under these favourableconditions
With the one exception of P. culminicola, all will lead to dominanceof broad-leavedtrees.
narrowly restrictedtaxa (indicated with an asterisk As can be surmisedfrom its greatergeographical
in Table III) have a montane distribution with a and altitudinalrange,the genus Pinus has a consider-
narrowaltitudinalrangenot exceeding 1000 m. Pinus ably greater ecological amplitude than any other
culminicola is a high-montanespecies occurring at genus in the family Pinaceae (Farjon, 1990a). This
3000-3700 m. The term "endemic"has been vari- is especially true toward the warmerand drier eco-
ously interpretedin the literaturebut is here restricted systems of scrubland/woodland bordering semi-
to taxa with a localized distribution (narrow en- deserts and in pine savannas with prolonged dry
demic). Rare but scatteredspecies which occur over seasons and naturalfire [Link] these
a large area-e.g., P muricataand P. pinceana-are conditions, involving specializationof vegetative as
thus excluded from this category; their distributions well as reproductiveorgans, have led to radiations
are more aptly described as disjunct or interrupted. of taxa and colonization of environmentswhere few
No narrowendemics occur in the southernregions; other trees can survive. Such specialization is espe-
most are western and northeasternand eastern Mex- cially true for the pines of California and Mexico.
ican. Both regions have experienced great geological and
The Caribbeantaxa are mostly lowland pines, with climatic changes in relatively recent (Miocene
P cubensis ascending to a maximum of 1200 m in throughPleistocene) geological time; it has therefore
the mountainsof eastern Cuba. Several occur only been suggested that much of this diversity reflects
just above sea level, but this could be due in part to relatively recent evolution (Little, 1962; Axelrod,
a limited topography,such as on the Bahamas and 1980).
the Florida Keys. One notable exception is the only Mexico is a predominantlyupland country, with
pine on mountainous Hispaniola, P occidentailis, more than one-half of its area above 1000 m. The
which, with an altitudinal range of (200-)900- Central American countries where pines occur are
2700(-3175) m, exceeds thatof all other LatinAmer- also [Link] lowlands occur on the
ican pines (Darrow& Zanoni, 1991). Yucatan Peninsula, the Atlantic coastal plains of
Central America, and, on a smaller scale, in the
Isthmus of [Link] P oocarpa de-
ECOLOGY scends into the lowlands of the isthmus, only the
The geographical distributionof the Pinaceae in Atlantic coastal plain from Nicaraguato Belize has
the Northern Hemisphere reflects the climatically significantpine forests with P caribaea [Link]-
dominatedecological conditions under which these ensis. "Pine savannas"also occur on several Carib-
conifers thrive and compete successfully. Relatively bean Islands,but here, too, most islands with natural
low temperaturesduring the growing season are a pine forest are mountainous. Because of Mexico's
limiting factor to deciduous broad-leavedtrees, and extraordinarilyvariedtopographyand climate, which
evergreenconifers take advantageof this; as a result range from wet, lowland tropical rainforests and

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56 Flora Neotropica

montanetropicalcloud forests to hot scrublandsand are species of Juniperusand often of [Link]

deserts and snow-capped mountains, often in close type is the equivalentof the pinyon-juniperwoodland
proximity to each other, virtually all ecotopes in of the United States, especially of the variants in
which pines are found worldwide are also present in southernArizona and New Mexico. This woodland
this region. Due to the great elevation of some of forms the lower limit of the upland forests in much
the mountainsand their situation between 12?N and of the interiorpartof northernMexico, especially on
33?N latitude,climatic zonationis pronouncedwithin the semi-arid slopes and mesas bordering the two
short distances and isolation of populationsand taxa Sierra Madres. Studies (papers in Everett, 1987) of
is commonplace. Several authorshave tabulatedcli- succession after disturbance, as well as of stand
matic zones for Mexican pines (e.g., Loock, 1950; structuresof mature woodland types, indicate that
Eguiluz, 1982, 1988), using altitudinal,temperature, Juniperus would dominate the vegetation, perhaps
and precipitationgradientsas criteria. even to the exclusion of Pinus, in the prolonged
absence of majordisturbancessuch as fires or (over-
Plant Communities )grazing. In all pinyon pines and some white pines,
seed is [Link] researchcarriedout in the
The vegetation of Mexico has been classified by United States, the pinyon jay (Gymnorhinuscvano-
Mirandaand Hernndez (1963). Pines occur in chap- cephalus) and Clark's nutcracker(Nucifraga colum-
arral,juniper woodland, pine forest, oak forest, de- biana) are the bird species involved in pine seed
ciduous forest, and fir forest, and in transitional dispersal (Tomback & Linhart, 1990). No detailed
vegetationas well as mixed types of these. Chaparral, studies on these mutualisms have been carried out
best developed in northwesternBaja California and for Mexican pinyon pine species, but it has been
north into Californiaon the lower Pacific slopes of suggested that two species in the genus Apheloconma
the mountainranges, is a semi-evergreenvegetation and the raven (Corvus corax) perform a similar
dominatedby [Link] is a coun- function (Peterson& Vargas-Barajas,1993). The ex-
terpartin parts of the foothills and lower slopes of treme similarity and homology of the cones and
the Sierra Madre Oriental, referred to as montane "wingless" seeds of P cenbroides and its (close or
chaparral. Pinus attenuata, P muricata, and, to a more distant) allies-including P edulis, for which
lesser extent, P monoplhyla and P quadrifolia are mutualism with Clark's nutcrackerand the pinyon
associated with the Pacific variant;P cembroides,P jay has been demonstrated(VanderWall & Balda,
culmniicola, P nelsonii, and P pinceana with the 1977; Ligon, 1978)-make it very likely that such a
interiortype. There is a climatic difference between bird connection exists in Mexico as well. The scrub
the two types: the Pacific coast type (chaparralin jay (Apheloconmacoerulescens) is largely dependent
the strict sense) has winter rains (and summerfog in on pinyon seeds where it lives in this type of wood-
its lower reaches) and is largely frost-free;the inte- land and stores them, yet this bird also occurs in a
rior type has summer rains (thunderstorms),occa- variety of habitatswhere no pines exist (Peterson &
sional frost, and, at high elevations, snow. Both are Vargas-Barajas,1993); furtherstudies on the bird's
subject to natural fires by lightning. Adaptations ecology seem to be necessary to determinethe rela-
are diverse and specialized. Prolonged drought and tionship between tree and bird. On the other hand,
intensive radiation have selected for short, rigid similar but larger seeds occur in P mnaximartinezii,
leaves. Along the Pacific coast, where fires are both which has developed an extremely large, hard-scaled
relatively frequent, with 20-50-year intervals in cone from which the seeds are not easily removed
closed-cone pine forests, and intense,the closed-cone as they are in P cembroides and its allies. It seems
type of seed dispersal, requiringextreme heating of unlikely thatjays can handlethese cones as they can
the outer partsof the persistentand serotinouscones the small, soft-scaled cones of the other pinyon
to open, has evolved in a number of species. The pines. Is the raven responsible for dispersal here?
seeds have articulatebut effective wings, which en- Differences in thicknessof the seed coat and content
able the seeds to be transportedsome distance and of starch and fatty oils in the seed among various
be scattered sufficiently to land in places where fire species of pinyon pines may also be of as yet un-
has eliminated the chaparralundergrowth. known ecological significance.
Juniper woodland is usually an open arborescent Pine forests ("Pinares")are a heterogeneous as-
vegetation type with well-interspersedsmall trees, in sembly of forest types dominatedby species of Pinus
which P. cembroides and/or its close relatives may (Miranda & Hernandez, 1963). They extend from
occur in various proportions;the other components the lowlands to the tree line and from xeric pinyon

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Distributionand Ecology 57

pine woodland to mesic montane mixed pine forest conditions in northernMexico. In the southernparts,
with a tall, closed canopy. Foothilland lowlandtypes seasonality is dominated by precipitationlevels; in
phase into "pine savannas"on the Atlantic coastal the north, low winter temperaturesare more im-
plains in Central America, where trees are widely [Link] these differences are primar-
spaced and the groundcover is dominatedby grasses, ily found in the foliage, with long, slender, often
with scattered fire-resistant(resprouting)shrubs. In pendulouslax leaves occurringespecially in pines of
the Caribbean,P. caribaea, P. elliottii [Link], and subgen. Pinus in the warm temperate regions and
P tropicalis occur in pine savannas, but in Central shorter, thick and rigid leaves predominantin the
America only P. caribaea var. hondurensisoccupies winter-cold Sierra Madres of North Mexico. The
this type of vegetation. Seedlings have an elongated predominantcone type is thatof Pinus [Link],
stem (P caribaea var. hondurensis) or develop a showing moderateto extreme diversificationin de-
"grassstage";both aredefences againstvery frequent velopment of the apophyses (see Figs. 6-7), pro-
grass fires which usually occur every few years. tecting the inner parts of the scales with the seeds
Annual precipitationcan be high, but it is unevenly from intenseradiationby thicker,protrudingapophy-
distributed and there is a prolonged dry season. ses on the sun-exposed side. Cones open gradually
Edaphic conditions prevent moisture retention on and are persistentuntil the seeds with articulatebut
most pine savanna sites, and in the tropical climate fully effective wings are dispersed. Several species
total evaporationis extremely high. In the adjacent range from the foothills to high mountainsbut are
upland areas, precipitationincreases rapidly, espe- nevertheless frequent constituents of the montane
cially on sites exposed to moisture-ladenwinds from pine forest belt. Their heterogeneity(see above) may
the Gulf of Mexico. Here P. cubensis and P. occiden- reflect genetic variation as much as environmental
talis are the pines of the Caribbeanuplands,but both adaptationto the variousclimatic and edaphiccondi-
occur as the only species of the genus in their tions they occupy. Pinus devoniana, with large,
[Link] the mainland,it is mainly woody cones and long, thick leaves, remains a me-
P oocarpa that occupies this zone above the pine dium-sizedtree and seems to occupy open secondary
savannas,and more locally also P [Link] woodland more often than its close relative, P mo,n-
canopy is more often nearly closed and there is a tezurnae,in which these featuresare less pronounced.
predominanceof shrubs in this type of pine forest, Intermediateforms between both species are often
among which are many legumes, but grasses are still difficult to classify, and variationsof the cones have
abundantin most places. Fires (many of them man- often been interpretedas representingdistinct taxa
made) are still frequentbut occur at longer intervals. or hybrids. Investigation of the possible effects of
Especially P. oocarpa has developed persistentand environmental conditions on the morphology of
semi-serotinouscones, which enables the species to leaves and cones in this complex could help delimit
be opportunisticin its dispersalof seeds after fire. genetically based, fixed characterstates more clearly.
The more mesic pine forests of the montanezone, There is a very gradual transition between pine
generallybetween (XX)and 2600 m above sea level, forest and oak forest at montane altitudes: most
are more diverse. In much of Mexico and Central forests classified as pine have a mixtureof Quercus
America,more thanone species, and not infrequently and most oak forests harbour some Pinus. Many
as many as 5-8, may occur in a single forest, with oaks are deciduous, shedding leaves during the dry
usually some altitudinal zonation involving some season, i.e., roughlyfrom Novemberto June. Natural
species but with others ranging throughoutthe pine disturbances,such as fire, now increasinglybrought
forest. On one hand, sympatry is common among about by man, tend to promote Pitus, but selective
pines in these forests, to a greaterextent than any- logging of pines in many areas has given Quercusa
where else. On the other hand, there are few if any chance to close the canopy, preventing pines from
other conifer genera present, except at the higher regeneratingin the absence of calamitousevents.
altitudes. In this they contrast with the equally di- A few species of Piins are commonly associated
verse conifer forests of Californiaand Oregon,which with broad-leavedtreesof warmtemperatedeciduous
are "mixed,"i.e., composed of membersof different forests. For instance, P. strobus var. chiapensis and
families and genera but with only a few representa- P tecunumaniiare almost always growing with Liq-
tive species of each. Within the montanepine forest uiMamnbarstyracifua and various other associated
belt, there is a climatic zonation from south to north, broad-leaved trees characteristic of the moisture-
with warmtemperateconditionsprevailingin Central catching slopes of the mountainsborderingthe Gulf
America and southern Mexico and cool temperate of Mexico in Mesoamericafrom Chiapasto Hondu-

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58 Flora Neotropica

ras. Precipitationis very high, often exceeding 3000 cola. This shrub-likepine may cover extensive areas
mm annually, and fog occurs year round. Other exclusively, as on Cerro Potosi, or in association
species of Pinus thatcommonlyoccur in these forests with other conifer species (Abies, Picea, Pinus) in
are P. patula and P. teocote. The pines often occupy "parklandcommunities" or open woodland. Apart
the steepest slopes and crags at higher elevations from the populationon Cerro Potosi (3713 m), it is
within the temperatedeciduous forest belt. A gradual not a treeline species (Riskind & Patterson, 1975)
transitiontowardoak forest occurs especially toward but, rather,a species associated with an open, mon-
the drier eastern slopes, where pine species more tane chaparral,merging with P hartwegii forest at
tolerant to drought appear, such as P montezumae more favourablesites, where the calcareous rock is
and P pseudostrobus. covered with deeper soils and the slopes are less
On cool, cloudy, wet mountain summits up to exposed to wintry,snow-laden gales from the north.
3000 m, or in belts on largersierras or taller moun-
tains, species of Abies-mainly A. guatemnalensis
in Mesoamerica and A. religiosa further north in Mycorrhizae
Mexico-dominate the coniferous forest. Locally, Most plants form a symbiotic association of fungi
stands of Cupressuslusitanica may replace the firs, with their roots creating [Link] species of
or both genera may occur in mixed stands. In this Pinus form ectotrophicmycorrhizaewith the hyphae
mixed coniferous forest Pinus avacahuite, P max- of basidiomycetousand ascomycetous fungi; in this
iminoi, P pseudostrobus, and, often at the upper type the hyphae form a mantle aroundthe fine roots
limit, P hartwegii are common; in northwestern and root hairs and rarely penetratethe cortex cells.
Mexico, P strobiformis replaces P ayacahuite. In Roots affected by mycorrhizaecease to grow longitu-
Baja California on the Sierra San Pedro Martir,a dinally; instead the mycelium to which the mycor-
Californianmixed coniferous forest occurs with Ab- rhiza is attachedextends the root system, increasing
ies concolor. Pinus lambertiana, P jeffreyi, and P the absorbingcapacityof the roots. The fungus dislo-
contorta var. murrayana. This northern type has cates the outer cortex cells of the root and forms a
warm summers and snowy winters, while the south- network of hyphae called the "Hartig net." The
ern equivalents have a more equitableclimate domi- symbionts (plant and fungus) cooperate in growth
nated by fog and rain. In these high-montaneforests, and function. The role of the fungus is primarily
where competitionamong tall-growingconifers (and absorptionof soluble nutrients;that of the host, to
fewer broad-leaved trees, mainly oaks) is severe, provide the fungus with its assimilation products
effective seed dispersal is of utmost importanceto (organic carbon compounds). The increased intake
reach distant gaps. The species all have cones that of nutrients-e.g., phosphates-enables the pine tree
open readily upon drying at maturityto release the to subsist on soil types otherwise too poor for tree
winged seeds (P avacahuite,P lambertiana,P strob- growth. Many differentspecies of fungi are involved
iformlis);especially those of P strobiformis, with in mycorrhizal symbiosis, both in temperate and
reduced wings, are also [Link] cones of tropicalclimatic zones (Barnes et al., 1996).
these conifers are placed high in the treetops, as in
Abies, or toward the end of higher branches in
Diseases and Predators
the pines, to facilitate long-distancedispersal of the
seeds. Under naturalconditions in pine or pine-oak for-
Finally, at the highest elevations in Mexico and ests with mixed species and/or mixed age classes,
Guatemala, some pines grow at or near treeline. both diseases and phytophagous predators have a
Foremost among these is P hartwegii, which is more or less chronic but low-level negative effect
scattered on the highest mountain summits from on tree [Link], such as Arceuthobiumspp.
Honduras to northeasternMexico. In Mexico and (Hawksworth& Wiens, 1972), can be abundantin
Guatemala it often forms pure stands, in Mexico some partsof the forest but rarelypreventtrees from
encircling the snowy summitsof the tallest volcanos long-termsurvival and [Link] attacks,
such as Pico de Orizaba,Popocatepetl,and Iztaccihu- if not limited to physically damaged trees, rarely
atl, generally to about 4000 m altitude. In northeast- spread far and wide. Unlike these pathogens, some
ern Mexico, the summits of several mountains, all insects-e.g., the pine bark beetle (Dendroctonus
of which are considerablylower than 4000 m, have mexicanus)-are capable of large-scale destruction,
only scattered stands of P hartwegii, and of some especially in single-species pine forests. Enormous
more or less extensive populations of a dwarfed tracts of Pinus oocarpa forest in Honduras were
species closely related to P cembroides, P culmini- defoliated in 1963-1966, and thousandsof hectares

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Uses and Conservation 59

of pine forest in Mexico were similarly affected However,if regenerationis somehow prevented(e.g.,
in 1949-1950 (Perry, 1991). Many areas are soon by livestock grazing), a more permanent shift to
naturallyreseeded,and the overall effect undernatu- other types of vegetation may be the ultimate result
ral conditions may be rathersimilar to a forest fire. of insect epidemics.

USES AND CONSERVATION

to the attentionof foresters, with the shift to multi-
PINES AS RENEWABLE RESOURCES
purposeforestrygoals, especially in the UnitedStates
The pines collectively constitute one of the most (Everett, 1987) but also in Mexico, where three
valuable natural resources in Mexico and Central recentsymposia were held on the [Link] yield
America (Carvajal& McVaugh, 1992); in Mexico of edible seeds representsa type of sustainableuse
they produced nearly 7.5 million cubic meters of that was already practised by prehistoric tribes in
pine logs in 1986. The most importantproducers pre-Columbiantimes (Passini et al., 1988). It is
were the states of Durango,Chihuahua,and Jalisco, probablyan importantsource of cash locally. Yet the
in that order. Hondurasproduces more pine timber collecting of the seeds, mostly in September, is
than any other Central American country (Perry, tedious labour carried out mostly by an itinerant
1991), and a substantialportion of its economy is labourforce of women and [Link] crops vary
based on it. Exploitationof the extensive pine forests considerablyfrom year to year (Styles, 1993b), with
on Hispaniola in the Caribbeanonly began in 1920 bumperharvestsoccurringapproximatelyevery five
(Darrow & Zanoni, 1991). Wood production for years, and the methodof harvestingdiminishescone
lumberis most important,followed by pulped wood production(Morales & Camacho, 1988). Although a
for kraft paper and cardboardand local uses for considerablevolume of seed is marketedas far away
firewood, fence posts, etc. Pine's importancelies in as Mexico City, most of it is sold locally, and this
the fact that the xylem of most "hard"pine species type of use can at best be considered a cottage
produces long fibres, giving the final productextra industry(Styles, 1993b). Largerseeds are produced
strength."Soft" pines, less abundant,are sought for by P maximartinezii,a species with a very limited
their less resinous, evenly grainedwood. In Mexico, distribution;it was discovered by the botanist J.
currentlythe most heavily exploited species are Pi- Rzedowski by way of the exceptionally large seeds
ntuspatula, P oocarpa, P pseudostrobus,P herrerae, he saw being sold in the market at Juchipila in
P leiophylla, and P arizonica (Perry, 1991; Styles, southernZacatecas.
1993a).However,frequentlymixturesof pine species A commerciallymore importantsecondaryproduct
are harvested without regard to species. In Cen- of pines is resin, mainly tapped from "hard"pines
tral America, P caribaea var. hondurensis and P (Pinus subgen. Pinus). Some 200,000 ha of pine
oocarpa arecommerciallythe most importantspecies forest in Mexico are exploited in this way, according
at the lower and middle altitudes,where they are the to the CamaraNacional de las IndustriasDerivadas
dominantcomponentsof the forest, and are similarly de la Silvicultura(CNIDS). In CentralAmerica,it is
exploited. In the Guatemalanhighlands, mixed pine also a major secondary product, derived from the
forests are the rule and exploitation largely does not extensive forests of P oocarpa, covering large parts
discriminateon the basis of species. In that country, of Hondurasand Guatemala,and of P pseudostrobus
but also in other densely populatedruralregions of and other species in the [Link] resin forms
Central America and Mexico, pine trees with the basis for the turpentine industry, which is a
branches lopped off from low on the bole upwards majorsource of revenue in several Mexican states-
are a common sight. Often very little crown is left- particularlyChiapas, Oaxaca, Michoacan, Jalisco,
insufficient to sustain the tree for long, as most and Nuevo Le6n-and in Hondurasand Nicaragua.
species of pines do not coppice. The branches are If the process of resin tappingis carriedout carefully,
used for fuel. the tree does not appearto suffer any ill effects for
In northernMexico, P cembroides is widely ex- many years. Frequently,however,large stripsof bark
ploited for firewood and charcoal burning,and to a are crudely removed from the bole, allowing entry
limited extent locally for [Link] trees are too of pathogensthat graduallyweaken and kill the tree.
small and heavily branchedto yield long, straight The preferred species for resin tapping appear to
lumber, and they are also less desirable for pulp be P oocarpa, P montezumae, P teocote, and P
wood. The pinyon pines have recently been drawn [Link].s.

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60 Flora Neotropica

Several other countrieshave or have had an inter- of growth, form, and wood properties,especially in
est in Latin American pines, especially the (sub-) P caribaea, are preferentiallysought out. However,
tropical species, introducingthem for plantationfor- such selections may also turn out to be the more
estry schemes in their own territoriesor in other "aggressive" invaders in the event of (unwanted)
(sub-)tropicalcountries. Pines are virtuallyabsent in naturalization(Hughes, 1994). After Pinus radiata, P.
the Southern [Link] indigenous conifers patula has now become one of the most troublesome
there do not have an equivalentproductionpotential invasive species of pine threateningnaturalvegeta-
or adaptabilityfor fast-growing,even-aged monocul- tion and biodiversity in the highlands of southern
tures. South Africa was one of the first countries to Africa. In many cases, it will be only a matter of
researchthese potentialsand implementintroduction time for the newer introductionsto follow suit, the
programs(Loock, 1950; Liickhoff, 1964). Pinus pa- apparenttime lag between introductionand invasion
tula was introducedin South Africa from Mexico as being a thresholdrecently overcome by a numberof
early as 1903. Loock, in 1947, was the first to species (Richardsonet al., 1994).
conduct a systematic search for suitable species,
followed in 1957 by Liickhoff, who expanded the
CONSERVATION
efforts to P caribaea [Link], to a lesser
extent, P [Link] Food and Agricultural The importance of pines to the forestry-based
Organization(FAO) of the United Nations has recog- industriesin the region means that timber exploita-
nized the potentialof the CentralAmericanpines for tion is increasing,and in many areas it is indiscrimi-
afforestation in tropical countries since the early nate. Areas formerlycovered with pine forests have
1950s and has cooperatedwith governmentalforestry now been depleted, and the resultingeroded hillsides
departmentsin Mexico and severalCentralAmerican and mountainsidesare a common featurein all coun-
countries (Styles, 1993a). Concernsabout deforesta- tries of the region. Exploitationis almost exclusively
tion in tropical countries (e.g., at the Eighth Com- of naturalstands, where regenerationis wholly de-
monwealth ForestryConference, held in Nairobi in pendenton the remainingseed trees in the area and
1962) focused on the loss of timber(not on biodiver- the conditions of soil, grazing, and fire that prevail
sity) and led also to suggestions of afforestationwith after clear-cutting. Hundreds of small to medium-
(neo-)tropicalpines. This in turngeneratedresearch, sized sawmills have penetrated most forested re-
funded by the British Government,at the Common- gions. With the building of roads, logging trails
wealth Forestry Institute,now the Forestry Institute furtheropened up the mountainsto broaderuse by
(OFI) of the University of Oxford, beginning in man. Sawmills require and use only trees with the
1963, focusing on explorationand seed collection of best form, so forests are frequently"creamed"of the
P caribaea var. hondurensis. Later, other species straightest,small-crownedindividuals,a practicethat
were added, e.g., P oocarpa, P patula, P tecunu- will ultimately lead to the genetic impoverishment
manii (discovered by F. Schwerdtfeger,investigating of a species (Vela, 1976; Styles, 1993a). Plantation
insect attacks on pines on behalf of FAO in Guate- forestry, although begun, is still practised on too
mala), P maximinoi (then known as P tenuifolia), small a scale to be a substitute for exploitation of
and P pseudostrobus(Styles, 1993a). All these spe- naturalforests. Populationpressuresare increasingly
cies were introducedto several tropical countries in high, especially in alreadydensely populatedregions
South America, Africa, and Asia. More recently,the such as Hispaniola, El Salvador, Guatemala, and
CentralAmerica and Mexico Coniferous Resources many locations in Hondurasand Mexico. Indiscrimi-
Cooperative(CAMCORE),operatingfrom the North natedestructionof the forestenvironmentfor agricul-
Carolina State University and financed largely by tural production,especially on richer volcanic soils,
government as well as private-sectorsubscribers,is is widespreadin these regions.
continuing these programs,expanding them to still Species with restricteddistributionsare most vul-
more species (Dvorak & Donahue, 1992). Seed is nerable, but the actual threatof extinction of these
being collected from many locations and established taxa depends on the situation locally. Some of them
in two types of field plantings: gene conservation grow on sites that are under little exploitation pres-
banks and genetic tests especially in Central and sure, as is the case with Pinus nelsonii, and unless
South America. Many provenanceshave been tested grazing and burning were to increase dramatically
and the most productive identified, and these are on these marginal lands, they seem relatively safe.
eventually to be planted in tropicalcountriesaround Farjon et al. (1993) have listed all conifer taxa
the globe on a commercialscale. Genetic differences presentlygiven a conservationcategoryby the IUCN

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Uses and Conservation 61

Species Survival Commission's Conifer Specialist conservation management (regeneration schemes

Group. Ithis list, as a continuously updatedworking foremost) is in place. Although P. pinceana is more
document,has recently been evaluatedin connection widespreadthan P nelsonii, the latteroccurs on soils
with the draftingof an IUCN Action Plan for coni- (often almost bare calcareous bedrock) that support
fers; see Farjon,[Link] pines in our region neitherforest nor forage for [Link]
are currentlylisted; the 10 listed below are of most of the vegetation cover related to livestock grazing
urgentconcern. appears to be a major threat to P. pinceana, at least
in areas we have seen (see also Perry, 1991). Both
Pitnusculminicola: critically endangered yield edible pinyon nuts ("pinones"), but in low
Pinus rzedowskii:critically endangered quantitiesas comparedto the largerand more abun-
Pilnusnmaximartinezii:endangered dant P cembroides,and exploitationis [Link]
Pinus pinceana: endangered island populationsof P radiata, recognized as var.
Pinus radiata var. binata: (critically) endangered binata, are under threatfrom grazing of feral goats.
Pinus cembroidessubsp. lagunae: vulnerable The problemis especially acuteon GuadalupeIsland,
Pinus cubensis: vulnerable the farthestout in the Pacific (Libby et al., 1968).
Pilnusjaliscana: vulnerable Eradicationof goats would restore regeneration,but
Pinus nelsonii: vulnerable this is moreeasily proposedthanpracticallyeffected.
Pinlus strobus var. chiapensis: vulnerable and is not likely to become a priorityfor a relatively
poor state like Baja California Norte, unless aided
A furtherseven taxa are categorizedas "low risk," by external funding.
meaning that although present trends are leading to The vulnerabletaxa are all being depletedmore or
a decrease in population sizes in most areas, they less rapidlyand are likely to fall into the endangered
will not lead to foreseeable genetic depletion and categories in the near future if present trends con-
extinction of the taxon. They are nevertheless listed [Link] exemplaryaccount is given by Zamoraand
because the trendsobserved may lead them into one Velasco (1977) for P strobus var. chiapensis in the
of the higher categories when current conditions state of Chiapas, Mexico. A preferentialpine for
persist for a long time. lumbermenwith the same wood quality as its near
Garcia and Gonzales (1991) have demonstrateda relative (P strobus var. strobus) of eastern North
dramaticdecrease in area covered by P. culminicola America, it has been excessively exploited over
on Cerro Potosi since Beaman and Andresen (1966) much of the state. Regenerationhere, as elsewhere,
investigatedthe same [Link] principalthreat is hampered by livestock grazing. At the time of
is fire, to which the species is not resistant(see also Zamoraand Velasco's study, some small exclosures
Perry, 1991). There is as yet no official protection had been establishedto observe regenerationwithout
for any of the areas (mountainsummits) where this grazing pressure, and plantation forestry with the
species occurs. Pinuts rzedowskii occurs in three species had been started up. However, as Perry
small populationssurroundedby a largebutexploited (1991) pointed out, with the currentgrowth of the
pine forest. Two are on inaccessible talus slopes, populationin Mexico and CentralAmerica,the need
and although one is situated right underneatha fire for food productionincreasingly becomes the over-
lookout, they can potentiallybe wiped out by a single whelming factorand nullifiesmost efforts to preserve
wildfire. The third and largest population is in part the pine forests. The situation on the Caribbean
mixed with other species. Although foresters and Islands is similar. The best strategy for preserving
conservationists are aware of these pines, none of the biodiversity of Pinus in this part of the world
the populations enjoys conservation status at the where it is still the greatest is probablyto establish
currenttime. A similar situationapplies to the single a numberof nationalparksand [Link]
but larger population of P maximartinezii(Farjon, like Honduras and Mexico already have a good
1994). The woodland is privately owned by a few number of these, but without financial support for
villagers who use the area for cattle grazing and pine protectionand managementfrom the wealthierparts
nut collecting at the same time. Both may have of the world where conservation of biodiversity is
adverse effects on natural regeneration. Fire is a professed to be valued so greatly, we predict that
hazard,but awareness is growing and effective land most of these areas, too, will succumb sooner or
managementin cooperation with local people may later to the more basic needs of a growing human
prevent disastrous calamities until adequate in situ population.

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62 Flora Neotropica

SYSTEMATIC TREATMENT

Pinus Linnaeus,Sp. PI. 2: 1000. 1753; Gen. PI., ed. bud scales or their remnants,persisting 2-30 years
5, 434. 1754; Miller, Gard. Dict., abr. ed. 4, 3. and falling as fascicles; length 2.5-50 cm, width
1754; Jussieu, Gen. PI. 414. 1789; L. C. Richard, 0.5-2.5(-7) mm, acicular (one species lanceolate),
Comm. bot. Conif. Cycad. 145, 147. 1826; Zuccar- plano-convex or triangular(rarely terete or flat) in
ini in Endlicher,Gen. PI. suppl. 2: 26. 1842; End- cross section, entire or serrulate;epistomatic or am-
licher, Syn. Conif. 81. 1847; Carriere,Tr. Gen. phistomatic(one species occasionally hypostomatic);
Conif. 291. 1855; Parlatorein Alph. de Candolle, resin ducts (1-)2 to several, variously positioned in
Prodr. 16(2): 378. 1868; Shaw. Gen. Pinus 24. the mesophyll; vascular bundles single or double,
1914; Pilger in Engler & Prantl, Nat. Pflanzen- enclosed in a stele. Pollen cones spirally arranged
fam.,ed. 2, 13: 331. 1926; Little & Critchfield,U.S. near the proximal end of new long shoots, ovoid-
Dept. Agric. Forest Service Misc. Publ. 1144: 7. oblong to cylindrical;consisting of a thin axis with
1969; Price, J. ArnoldArbor.70: 262. 1989; Carva- numerousspirally arranged,(sub-)peltatemicrospor-
jal & McVaugh in McVaugh, Fl. Novo-Galiciana ophylls, each bearing two longitudinally dehiscent
17: 32. 1992; Kral in Flora of North America sporangia;pollen bisaccate. Seed cones (ovuliferous
Committee(ed.), Fl. N. Amer. 2: 373. 1993. Lecto- cones) subterminal or sometimes appearing to be
type species (vide Britton,N. [Link] 5. 1908). lateral, borne singly or more commonly clustered,
Pintts sv',estris Linnaeus, Sp. PI. 2: 1000. 1753. pedunculate,maturingin the second year or rarely
in the third year, shed early or variously persistent,
Pinus sect. Petce Grisebach, Spicil. Fl. Rumel. 2: 347.
1844. initially erect; maturecones pendulousor spreading,
StrobusOpiz, Lotos (Prague)4: 94. 1854. opening soon or variously serotinous, (obliquely)
Cairyopitys Small, Fl. s.e. U.S. 29. 1326. 1903. ovoid to cylindrical, 2-60 cm long. Seed scales
ApinusNeckerex Rydberg,Bull. TorreyBot. Club32: (ovuliferous scales) persistent, obovate to oblong,
597. 1905. thin or thick woody, attached to a slender to very
[Link].3: 69. 1913.
Nieuwland,
Leucopitys thick axis; the exposed portion(apoplhysis)variously
Ducanlpopinus Chevalier, Rev. Bot. appl. & Agr. trop.
24: 30. 1944. thickenedand/orelongated; bearing in a terminalor
dorsal (abaxial) position the remnantof the exposed
Derivation:classical Latin name pinus. English pine, portion (iimbo) from the first year's development,
Spanish pino. which may be terminatedin a (deciduous) spine or
Trees,or less commonly shrubs, evergreen, resin- prickle. Seeds obovoid, slightly flattened;seed coat
ous in wood, bark,leaves, and often cones, aromatic; thin or thick, without resin vesicles; seed with an
1-75 m tall, diameter at breast height (dbh) 5-320 adnate or articulatemembranouswing derived from
cm. Trunkmonopodial, or shrubs in some species the adaxialpartof the seed scale; several times larger
multistemmed,branchingin [Link] of than the seed and effective, or (greatly) reduced
older stems variously (deeply) furrowed or plated, and in articulate wings remaining with the scale.
to thin and scaly or thin and smooth, ridges and/or Cotvledons3-24, denticulateor entire. Chromosome
plates layered or scaly, more or less conspicuously number 2ii = 24 (x = 12).
exfoliating in often intricate patterns. Wood with
normally conspicuous annual ring boundaries;resin
ducts present or absent; ray tracheids present or Glossary of Botanical Terms Used in the Keys
absent, xylem [Link] dimorphic, adnate seed wing: wing not easily separable from
with long shoots and dwarf shoots; long shoots the seed, usually tearing instead (Fig. 61-J).
uninodalor less commonly multinodal;dwarf shoots amphistomatic: with stomataoccurringon all faces
axillary to spirally arranged primary leaves (cata- around the leaf; there are usually more lines or
phylls). Catapllh!ls on long or short decurrentleaf rows of stomataon the convex abaxial (outer) face
bases (pulvini) on long shoots, non-chlorophyllous, than on each of the two adaxial faces.
enclosing "winter buds" (primordiallong shoots or apophysis: part of the seed scale that is exposed in
ovuliferous strobili), subtendingdwarf shoot buds or a mature, closed cone, representing the second
pollen strobili,early deciduousor [Link]- phase of growth (Fig. 6Cb-1, Dc-i).
ary leaves (needles) borne in fascicles of (1-)2-5 articulate seed wing: wing easily separable from
(-8) on dwarf shoots; the fascicles surroundedat the seed, it is held by two claw-like appendages
base by an early deciduous or persistent sheath of (Fig. 6K).

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Systematic Treatment 63

attenuate: widest near the base, gradually tapering noting the interruptedarrangementof leaf fascicles
to the apex; here referringto the shape of ovoid- and some weaker lateralshoots between the main
oblong closed cones. nodes of branching.
cataphyll: scale leaf or primaryleaf, which initially pulvinus (pi. pulvini): small, raised section on a
covers the vegetative buds (Fig. 1C); later they long shoot at the base of a cataphyll,either form-
are more or less persistenton the shoot (Fig. 1A, ing a ridge downward (decurrent, Fig. IA) or
F). ending abruptly(not decurrent,Fig. IB).
endodermis: layer of large cells surroundingthe resin ducts: tubulartractscontaining resin and situ-
stele with the vascularbundle(s) in a leaf, in cross ated in the wood as well as in the leaves of pines;
section seen as a ring of cells, of which the outer diagnostic of several species is the position of
walls are thin (Fig. 4D) or thickened(Fig. 4E). these resin ducts in the leaf in cross section:
epistomatic: with stomata occurring on the 1-2 external (Fig. 2A-D), medial (Fig. 2E-G), inter-
adaxial (inner) leaf faces only. nal (Fig. 2H-J), septal (Fig. 2K-L).
fascicle sheath: Aggregationof scales enveloping a serotinous cones: seed cones that open their scales
leaf fascicle; it can be deciduous or persistent;in only afterextreme heatingor many years of dehy-
some species the scales recoil before becoming dration; normally found to be closed or nearly
deciduous (Fig. lHa-c). closed on the tree.
leaf: secondary,chlorophyllousleaf; commonly re- umbo: part of the seed scale that is exposed in the
ferredto as "pine needle." first phase of growth, distal to the apophysis (Fig.
leaf fascicle: A group of leaves (needles) growing 6Ca-2, Da-2).
from a dwarf shoot, initially or persistentlyenve- vegetative bud: the terminaland subterminalcom-
loped at its base by a sheath of scales (Fig. IGa- plex of shoot primordiumandbudsof dwarfshoots
b). with young leaves, covered by cataphylls (Fig.
multinodal shoot: a long shoot elongating in more IC).
thanone flushof growth,this detectedprimarilyby

Key to the subgeneraof Pinus

1. Pulvini decurrent(Fig. IA); leaves with 2 vascularbundles and variously positioned resin ducts (Fig. 2A-
L), with persistent(rarely deciduous) fascicle sheath; seeds with an articulate(Fig. 6K, L). rarely adnate,
wing..............................................................................................................................................subgen. Pinus (spp. 1-33)
1. Pulvini not decurrent(Fig. IB); leaves with a single vascularbundle and external resin ducts (Fig. 2A-D),
with deciduous (rarely persistent)fascicle sheath; seeds with an adnate (Fig. 61, J), rarely articulate,wing
or wingless when free from the seed scale (Fig. 6Nb).........................................................subgen. Strohus(spp. 34-47)

Key to the sections and subsectionsof subgen. Pinus

I. A conspicuous narrowband visible aroundumbo on seed scale; sheath of leaf fascicle deciduous...............
........................................................................................................................................sect. Leiophyllae(I. P leioph!lla)
1. No band visible around the umbo, differing from other parts of the apophysis; sheath of leaf fascicle
persistent.
2. Seed cones serotinous (remaining closed), sessile in tenacious whorls, or in pairs but then large and
with elongated, hooked spines ......................................................................................................... subsect. Atte
2. Seed cones opening (relatively) soon after maturity,(short) pedunculate,solitary or in whorls (open
cones may seem to be sessile).
3. Long shoots multinodal;vegetative buds often resinous;resin ducts internal.......................................
....................................................................................................................subsect. A str les (excl. P occide utlis)
3. Long shoots uninodal;vegetative buds usually not resinous;resin ducts variously positioned.
4. Seed cones asymmetrical,oblique at base or curved. 4.5-35 cm long; seed scales spreadingto 90?
or more; apophyses nearly flat to prominentlyraised (Fig. 6E-H).
5. Leaves relatively short, erect or spreading,or if up to 25 cm long. 1.5-2 mm wide, in fascicles
of (2-)3-5(-6) (count 15-20 fascicles) .............................................................. subsect. Polerosae
5. Leaves 20-50 cm long, drooping to pendulous,in fascicles of 5 or more ................ subsect. Pseudostrobi
4. Seed cones symmetricalor slightly oblique, ovoid or ovoid-oblong, (2-)3-10(-12) cm long; seed
scales spreading<90?; apophysis flat or slightly raised (Fig. 6E-F).
6. Seed cones up to 5.5 cm long; leaves in fascicles of 2-3 (count 15-20 fascicles)......... subsect. Contortae
6. Seed cones (4-)5-10(-12) cm long; leaves in fascicles of 2-5.
7. Leaves in fascicles of 2(-3) .......................................................................subsect. Pi s (2. P tropicalis)

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64 Flora Neotropica

7. Leaves in fascicles of 3-5.

8. Vegetativebuds resinous;umbos of seed cone scales with a persistentprickle......8. P occidentalis
8. Vegetative buds not resinous; umbos of seed cone scales without a prickle in mature
sct. Oocarpae
cones .........................................................................................................................subse

Key to the species of subsect. Attenuatae

1. Leaves in fascicles of 2 (rarely 3) (count 15-20 fascicles).
2. Umbo of seed scales with a persistentspine or prickle..................................................29. P. uricata var. Imuricata
2. Umbo of seed scales without a spine or prickle................................................................ . . P radita var. bi t
I. Leaves in fascicles of 3 (rarely 2).
3. Seed cones very large, 20-35 cm long; apophyses and umbos of seed scales very strongly developed;
leaves 1.9-2.2 mm wide .......................................................................................................................... 33. P coulteri
3. Seed cones smaller; leaves <1.6 mm wide.
4. Vegetative buds resinous;apophyses of seed scales strongly developed on one side of the cone (Fig.
7E).......................................................................................................................................................3 1. P. a ttenu
at a
4. Vegetative buds not resinous;apophyses flat or only slightly raised...................................................32. P. greggii

Key to the species of subsect. Australes

1. Leaves (6-)10-15(-18) cm long; seed cones (3.5-)4-7 cm long; seeds with adnate wing (Fig. 61) ......7. P cubensis
i. Leaves (I -)14-28(-30) cm long; seed cones 5-13 cm long; seeds with articulatewing (Fig. 6K, L).
2. Vegetative buds not resinous; leaves in fascicles of 2(-3) (count 15-20 fascicles) .............. 6. P elliottii var. dcensa
2. Vegetative buds resinous; leaves in fascicles of (2-)3-5.
3. Long shoots multinodal;fascicle sheath of latest leaves 15-20 mm long: leaves in fascicles of (2-)
3-4, very rarely 5 .............................................................5. P. (carihbae
3. Long shoots uninodal; fascicle sheaths of latest leaves (8-)l0-15 mm long; leaves in fascicles of
3-5 .....................................................................................................................................................8. P occilentt lis

Key to the species of subsect. Contortae

1. Leaves in fascicles of 2; vegetative buds resinous; umbos on seed scales with a persistentprickle...............
u1lt i ana
................................................................................................................................................3. P contorta var. m
1. Leaves in fascicles of 3; vegetative buds not resinous; umbos on seed scales with a minute, deciduous
prickle................................................................ 4. P iherrerue

Key to the species of subsect. Oocarpae

I. Cones broadly ovoid to subglobose when closed.
2. Cones semi-serotinous (only the distal seed scales parting), remainingon the tree; leaves 0.8-1.6 mm
wide; leaf resin ducts septal...............................................................20. P. oocarpa
2. Cones opening completely when mature,falling after 1-3 years; leaves 0.5-1 mm wide; leaf resin ducts
medial or internal.
3. Cones falling with the proximal seed scales missing; fascicle sheaths 10-14 mm long; leaves (8-)
10-16 cm long, 0.5-0.8 mm wide; leaf resin ducts internal......................................................21. P. praetermissa
3. Cones falling intact: fascicle sheaths 12-18(-25) mm long; leaves (14-)16-18(-25) cm long, 0.7-1
(-1.3) mm wide; leaf resin ducts medial..................................................................................... 24.. tectunumoatii
1. Cones ovoid-attenuatewhen closed.
4. Leaves in fascicles of (4-)5-7, rarely 8 ............................................................................................ 25. P. durangensis
4. Leaves in fascicles of 3-5, rarely 2.
5. Leaves with 9 or more lines of stomataon the abaxial face; leaf resin ducts internal.
6. Cones semi-serotinous,persistent,leaving some basal scales on the branchwhen falling; umbo of
seed scales flat or depressed ..........................................................................................................27. P pringlei
6. Cones opening at maturity,falling soon with peduncles attached;umbo of seed scales pyramidal.
.........................................................................................................................................................26. P tawsonii
5. Leaves with 3-7(-10) lines of stomataon the abaxial face; leaf resin ducts medial or septal.
7. Leaves in fascicles of (4-)5, rarely3; fascicle sheaths 8-10(-15) mm long; leaf resin ducts mostly
septal..............................................................................................................................................23. P jaliscan
7. Leaves in fascicles of (2-)3-4(-5); fascicle sheaths initially long (ca. 20 mm or longer); reduced
in matureleaves to 10-15 mm; leaf resin ducts medial.

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Systematic Treatment 65

8. Leaves (7-)10-15(-18) cm long, 1-1.4 mm wide..................................................................... 28. P. teocote

8. Leaves (11-)15-25(-30) cm long, 0.7-0.9(-1) mm wide ..........................................................22. P patul

Key to the species of subsect. Ponderosae

I. Leaves in fascicles of 3-5(-6). 5 usually present,often predominating.
2. Seed scales of cones relatively thin, flexible; leaf endodermiswith thin-walledcells; leaf vascularbun-
dles connate................................................................... 13. P hartwegii
2. Seed scales of cones relatively thick, rigid; outer walls of leaf endodermalcells thickened;leaf vascular
bundles mostly separate......................................................................................................................... 10. P ri onic
I. Leaves in fascicles of (2-)3(-4), rarely 5.
3. Leaves in fascicles of 3, rarely 2.......................................................................................................... . jeffre i
3. Leaves in fascicles of 2-3 or more.
4. Seed cones 5-10 cm long; fascicle sheaths of leaves 10-15(-20) mm long; leaves in fascicles of 2-
3, 1.3-1.6 m
mm mw wide
ide............. ;........ 9.. P
................................................................................................
P. pon(lerosa
po,derosa var. scoplrlorul
scoptdorum
4. Seed cones 8-15 cm long; fascicle sheaths of leaves (15-)25-35(-40) mm long; leaves in fascicles
of (2-)3(-4), rarely 5, 1.5-2 mm wide ....................................................................................... 12. P el nnii

Key to the species of subsect. Pseudostrobi

I. Fascicle sheaths deciduous at maturity;leaves pendulous ....................................................................... 19. P luholtii
I. Fascicle sheaths persistent;leaves drooping or spreading.
2. Cones 5-10(-12) cm long; falling intact (peduncle deciduous with the cone).
3. Seed scales of cones thin, flexible, in open cones usually strongly recurved;leaves often very slender,
drooping to pendulous,0.6-1 mm wide.......................................................................................... 18. P a i i oi
3. Seed scales of cones not flexible, not strongly recurvedin open cones; leaves droopingor spreading.
0.7-1.2 mm wide...........................................................................................................................17. P gl si
2. Cones (7-)10-35 cm long; leaving a few proximalscales on the branchwhen falling.
4. Fascicle sheaths (20-)25-35(-40) mm long, resinous; leaves 1.1-1.6 mm wide; outer walls of leaf
endodermalcells thin; seed cones 15-35 cm long. ....................................................................... 16. P. levoia
4. Fascicle sheaths (15-)20-30(-35) mm long, usually not resinous; leaves 0.8-1.3 mm wide; outer
walls of leaf endodermalcells thickened;seed cones 7-20 cm long.
5. Leaves with connate vascular bundles;seed cones usually obliquely ovoid when closed. 14. P pseudostrohus
5. Leaves with separatevascular bundles; seed cones usually ovoid-oblong to ovoid-attenuatewhen
closed.. ......................................................................................................................................15. P. onlte e

Key to the sections and subsections of subgen. Strobus

1. Umbo of seed scale terminal(Fig. 6Ca); leaves in fascicles of 5 ..............................................................subsect. Strohi
1. Umbo of seed scale dorsal (Fig. 6Da, c); leaves in fascicles of 1-4(-5).
2. Seed cones with -60, very widely spreading, very flexible scales; cones not longer than wide; seeds
relatively large, wingless when detached from seed scale ...........................................................subsect. Cembroiles
2. Seed cones with >60 scales spreading-70?; cones longer than broad;seeds large or small, wingless or
winged.
3. Leaves in fascicles of 3, rarely 4 (count 15-20 fascicles); seed cones oblong, irregular,pendulous.
up to 10-12 cm long; seed scales easily flexible......................................................................... subsect. Nelsoi e
3. Leaves in fascicles of (3-4-)5; seed cones regularly shaped, usually >10 cm; seed scales rigid to
inflexible.
4. Seeds small (ca. 8 mm long), winged; seed cones 10-15 cm long, with thin, rigid scales....39. P rz.e(dowskii
4. Seeds large (20-28 mm long), wingless; seed cones (15-)17-25(-27) cm long, with thick, inflexi-
ble scales...............................................................................................................................40 . P . i rt inezii

Key to the species of subsect. Cembroides

. Leaves in fascicles of 1, rarely 2.............................................................................................................46. P ono lla
1. Leaves in fascicles of 2 or more.
2. Leaves in fascicles of 5 (very rarely4 or 6); a low, spreadingshrub..............................................44. P cull,lilicola
2. Leaves in fascicles of 2-4(-5); a shrubor (small) tree.
3. Scales of fascicle sheaths recoiling strongly before falling, forming a rosette; leaves usually <1 mm
wide..................................................................................................................................................43. P ce broides
3. Scales of fascicle sheaths not forming a rosette; leaves usually 1-1.5 mm wide.

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66 Flora Neotropica

4. Leaves in fascicles of 2(-3), 0.8-1.1 mm wide........................................45. P. remota

4. Leaves in fascicles of (3-)4(-5), 1-1.5 mm wide ...................................................................47. P. quadrfolia

Key to the species of subsect. Nelsoniae

I. Leaves in fascicles of 3, rarely 4, but connate, appearingas one (Fig. 60B); seed cones on thick, 2.5-6.5
cm long and curved, persistentpeduncles..................................................................................................... 41. P. nelsonii
i. Leaves in fascicles of 3, rarely4, separate(Fig. 61C); seed cones on slender, easily breakingpeduncles....
......................................... ..............................................................................................................................4 2 . P pinceanu

Key to the species of subsect. Strobi

I. Leaves amphistomatic,with 2-4(-5) lines on the abaxial face; seed scales with broad, straight and flat,
smooth apophyses....................................................................................................................................35.P. a bertian
I. Leaves epistomatic (on adaxial face), or rarely with a few stomata on the abaxial face; seed scales with
tapering,recurvedor concave, often grooved apophyses.
2. Seed wing shorter than seed or vestigial (Fig. 6Mb); leaves (5-)6-11 cm long, 0.8-1.2 mm wide (on
sterile shoots).
3. Vegetative buds resinous; fascicle sheaths up to ca. 15 mm long; seed cones 10-15 cm long.............
..............................................................................................................................................36. P. flexilis var. reflex
3. Vegetativebuds not resinous;fascicle sheaths20-25 mm long; seed cones 12-30 cm long (or longer).
. strobiformis
.........................................................................................................................................................37..
2. Seed wings (much) longer than seed (Fig. 6J); leaves (5-)6-15(-18) cm long, 0.6-1 mm wide (on sterile
shoots).
4. Seed cones large, (10-)15-40(-50) X 7-15 cm when open; seed scales with recurved or reflexed
apophyses (at least the proximal scales recurved)....................................................................... 34. P avacahuite
4. Seed cones (6-)8-16(-25) X 4-8 cm when open; seed scales with straight, ? concave apophyses..
.......................................................................................................................................38. P strobus var. chiapensis

Additional Keys by Regions I-VII

Key to the species in Baja CaliforniaNorte (I)
1. Leaves in fascicles of 5; fascicle sheaths deciduous; seed cones >25 cm long, pendulous; umbo of seed
scale terminal (Fig. 6Ca)..................................................................................................................... 35. P lambertiana
1. Leaves in fascicles of 1-4, rarely5; fascicle sheaths persistentor recoiling and deciduous;seed cones small
or large, not pendulous;umbo of seed scale dorsal (Fig. 6Da, c).
2. Fascicle sheath persistent.
3. Leaves in fascicles of 2 (count 15-20 fascicles, especially on leading shoot).
4. Leaves 4-7 cm long, persisting 5-8 years; seed cones opening gradually,(3-)4-5.5 cm long........
.................................................................................................................................3. P contorta var. murravana
4. Leaves (7-)10-14(-16) cm long, persisting 2-3 years; seed cones serotinous,5-7(-8) cm long....
.................................................................................................................................29. P muricata var. nm uricata
3. Leaves in fascicles of 2-3 (count 15-20 fascicles, especially on leading shoot).
5. Seed cones usually in whorls of 2-5, serotinous,(5-)8-15 cm long.
6. Vegetativebuds resinous; leaves in fascicles of 3, rarely 2; seed cones ovoid-oblong to ovoid-
attenuate............................................ ...................................................... P..........31. a ttenuata
6. Vegetative buds not resinous; leaves in fascicles of 2, sometimes 3 on leading shoots; seed
cones ovoid or obliquely ovoid. .............................................................................. 30. P. radiat var. binata
5. Seed cones solitary or in pairs, rarely in whorls of more than 2, opening graduallyor soon after
maturity,10-35 cm long.
7. Seed cones very large and heavy, 20-35 cm long, very resinous;apophyses and umbos of seed
scales very strongly developed ..................................................................................................33. P. coulteri
7. Seed cones smaller, lighter, 10-17 cm long, not resinous;apophyses and umbos of seed scales
slightly or moderately raised. .......................................................................................................1 . P. effreyi
3. Fascicle sheath deciduous.
8. Leaves in fascicles of , rarely 2..............................................................................................46. P. monoph la
8. Leaves in fascicles of 3-5 usually 4 .......................................................................... 47. P quadrifoia

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Systematic Treatment 67

Key to the species in northwesternMexico (I1)

(Sonora, Chihuahua,Sinaloa, Durango,Zacatecas)
1. Bases of cataphylls (pulvini) not decurrent(Fig. IB); leaves with a single vascular bundle and external
resin ducts (Fig. 2A-D); sheath of leaf fascicle deciduous.
2. Leaves in fascicles of 2-3, rarely 4 or 5; seed cones small (3-5 cm), not longer than wide when open;
seeds wingless when detached from the scale ...................................................................................43. P. ce roies
2. Leaves in fascicles of 5, rarely 6; seed cones 10 cm long or longer; seeds with a short (sometimes
vestigial) wing.
3. Vegetativebuds resinous;fascicle sheaths up to ca. 15 mm long; seed cones 10-15 cm long.............
..............................................................................................................................................36. P fl exilis var. reflex
3. Vegetativebuds not resinous;fascicle sheaths20-25 mm long; seed cones 12-30 cm long (or longer).
.........................................................................................................................................................37. P strol iformis
I. Bases of cataphylls (pulvini) decurrent(Fig. IA); leaves with 2 vascularbundles and variously positioned
resin ducts (Fig. 2A-L); sheath of leaf fascicle persistentor deciduous.
4. Sheath of leaf ascicle deciduou ........................................................................................................... . P eioph ll
4. Sheath of leaf fascicle persistent.
5. Seed cones persistent,semi-serotinous,ovoid to globose when closed; leaf resin ducts mostly septal.
................................................................................................................................................................20.P. ooc rpa
5. Seed cones falling, at least after a few years, opening when mature,obliquely ovoid to oblong when
closed; leaf resin ducts medial.
6. Seed cones obliquely ovoid-oblong, often curved, 15-35 cm long; leaves in fascicles of 5, rarely
4 or 6 (count 15-20 fascicles) ................................................................................................. 16. P. devo ia
6. Seed cones ovoid or obliquely ovoid, less than 15 cm long; leaves in fascicles of 3-5, rarely 2.
7. Peduncledeciduous with the seed cone, which falls intact....................................................... 28. P teocote
7. Peduncle persistent;the cone leaves some of the basal scales on the branchwhen it falls.
8. Seed cones (4.5-)5-10(-14) cm long; apophyses of seed scales moderatelyraised (<'/2 as
high as wide); leaves rarely longer than 20 cm................................................................. 10. P. arizonica
8. Seed cones 8-15 cm long; apophyses strongly raised (>/2 as high as wide); leaves (18-)
20-35 cm long............................................................................................................... 12. P. engel annii

Key to the species in western Mexico (III)

(Baja CaliforniaSur, Nayarit,southernZacatecas,Aguascalientes,Jalisco, Colima, Michoacan)
1. Leaf fascicle sheaths deciduous.
2. Seed cones with '60, very widely spreading,very flexible scales; cones not longer than wide....43. P. cemnbroides
2. Seed cones with '60, inflexible or at least rigid scales; cones longer than wide.
3. Leaves epistomatic, rarely a few stomataon the abaxial face; one vascularbundle in the leaf.
4. Fascicles sheaths with distinct, individuallyfalling scales; seed cones cylindrical, 12-30(-60) cm
long; apophyses of seed scales, at least the proximalones, recurvedor reflexed ................37. P. strobiformis
4. Fascicles sheaths with basally connate scales recoiling to form a rosette before they fall; seed
cones not cylindrical;apophyses not recurvedor reflexed.
5. Leaves in fascicles of (3-)4-5; seed cones 10-15 cm long, with thin, fixed but flexible scales;
seeds small (ca. 8 mm long), winged................................................................................. 39. re o skii
5. Leaves in fascicles of 5, rarely3-4; seed cones (15-)17-25(-27) cm long, with thick, inflexible
scales; seeds 20-28 mm long, wingless ........................................................................40. P. inartineii
3. Leaves amphistomatic;2 vascularbundles in the leaf.
6. Leaves (4-)6-15(-17) cm long, spreading:seed scales with a conspicuous narrow band visible
aroundthe umbo........................................................................................................................... . P. eiophll
6. Leaves (15-)20-30(-40+) cm long, pendulous;seed scales lacking a band aroundthe umbo.........
.......................................................................................................................................................19. P lui holtzii
I. Leaf fascicle sheaths persistent.
7. Leaves in fascicles of 3, sometimes a few 2-5 (count 15-20 fascicles); seed cones (2-)3-6(-7) cm long.
8. Seed cones ovoid to subglobose when closed, shorterthan wide when opened......20b. P oocarpa var. trifblicata
8. Seed cones ovoid or obliquely ovoid when closed, longer than wide when opened.
9. Leaves slender, lax, (10-)15-20 cm long, 0.7-0.9 mm wide; seed cones (2-)3-3.(-4) cm long.4. P. herrerae
9. Leaves rigid, (7-)10-15(-18) cm long, 1-1.4 mm wide; seed cones (3-)4-6(-7) cm
long ...................................................... 28. P teocote
7. Leaves in fascicles of (4-)5-6, rarely3 or 7-8; seed cones (4-)5-10(-12) cm long or much
larger.
10. Seed cones (4-)5-10(-14) cm long; seed scales with flat or slightly raised apophyses
(Fig. 6E-F).
i1. Leaves (6-)10-25(-30) cm long.

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68 Flora Neotropica

12. Seed cones ovoid to subglobose when closed, shorterthan wide when opened.
13. Cones falling with the proximalseed scales missing; leaves (8-)10-16 cm
long, 0.5-0.8 mm wide; leaf resin ducts internal ........................... 21. P praetermissa
13. Cones persistent,intact when falling; leaves (I -)14-25(-30) cm long; leaf
resin ducts septal ..............................................................20a. P oocarpa var. oocarpa
12. Seed cones ovoid-oblong to ovoid-attenuateor obliquely ovoid when closed.
longer than wide when opened.
14. Proximal seed scales not parting when the cone opens; leaf resin ducts
septal. ...................................................................................................... 23. P jaliscana
14. Proximalseed scales partingwhen the cone opens; leaf resin ducts medial.
15. Seed cones with ca. 150-200 scales; apophyses of seed scales more or
less flat, weakly keeled, light brown to purplishblack ................. 13. P hartfwegii
15. Seed cones with ca. 90-120 scales; apophyses of seed scales (slightly)
raised, prominentlykeeled, ochraceous to light (reddish-)brown..........
..................................................................................................... .. P duranger nsis
II. Leaves 20-35 cm long.
16. Leaves 0.6-1 (-1.1) mm wide, drooping to pendulous;seed scales thin, spread-
ing 90? or reflexed when the cone is open; leaf hypodermis without or with
few intrusionsinto the mesophyll................................................................18. P. taximinoi
16. Leaves 0.7-1.2 mm wide, spreadingor drooping; seed scales thick, spreading
<90? when the cone is open; leaf hypodermis with numerous intrusions into
the mesophyll, some connecting with the endodermis ........................... 17. P douglasiana
10. Seed cones (7-)8-35 cm long; at least several seed scales with prominently raised
apophyses (rarely all nearly flat) (Fig. 6G-H).
17. Fascicle sheaths 30-40 mm long, resinous: leaves 1.1-1.6 mm wide; outer walls of
leaf endodermalcells thin; seed cones 15-35 cm long .................................... 16. P devoaniana
17. Fascicle sheaths (15-)20-30(-35) mm long. usually not resinous; leaves 0.8-1.3
mm wide; outer walls of leaf endodermalcells thickened;seed cones 8-20 cm long.
18. Leaves with partly connate vascular bundles; seed cones usually obliquely
ovoid when closed............................................................................. 14. pseu ostrob s
18. Leaves with separate vascular bundles: seed cones usually ovoid-oblong to
ovoid-attenuatewhen closed .................................................................... 15. P montezt umae

Key to species in NE & E Mexico (IV)

(eastern Chihuahua,Coahuila, Nuevo Le6n, Tamaulipas,northernZacatecas, San Luis Potosi,
Guanajuato,Queretaro,Hidalgo, Tlaxcala, northernVeracruz,northernPuebla)
I. Leaf fascicle sheaths deciduous.
2. Seed cones with -50. very widely spreading,very moveable scales; cones not longer than wide.
3. Leaves in fascicles of 5 (very rarely4 or 6, count 15-20 fascicles); a low, spreadingshrub...44. P culininicola
3. Leaves in fascicles of 2-4(-5); a shrub or (small) tree.
4. Scales of fascicle sheaths recoiling strongly before falling; leaves in fascicles of (2-)3(-4), rarely
5, < mm wide .......................................................................................................................... P cem ro es
4. Scales of fascicle sheaths not recoiling; leaves in fascicles of 2(-3), 0 .......................................45. P remote
2. Seed cones with -50 scales, spreading -90?, rigid, or if flexible, spreading very little; cones longer
than wide.
5. Leaves in fascicles of 3, rarely4 (count 15-20 fascicles); seed cones ovoid-oblong or short cylindri-
cal, irregular,5-10 cm long; seed scales very moveable; seeds wingless when detached from the
scale.....................................................................................................................................................42. P pinceana
5. Leaves in fascicles of (2-)3-5(-6); seed cones ovoid or long cylindrical, mostly longer than 10 cm,
regular;seed scales rigid; seeds winged.
6. Leaves in fascicles of 5, rarely 6: seed cones cylindrical.
7. Seed wing shorterthan seed or vestigial (Fig. 6Mb); leaves 6-11 cm long, 0.8-1.2 mm wide.
8. Vegetativebuds resinous;fascicle sheaths up to ca. 15 mm long; seed cones 10-15 cm long.
...............................................................................................................................36. P fl exilis var. reflexa
8. Vegetative buds usually not resinous; fascicle sheaths 20-25 mm long; seed cones 12-30
cm long (or longer).........................................................................................................37. P s ro ifo is
7. Seed wings longer than seed (Fig. 6J); leaves (6-)8-15(-18) cm long, 0.6-1 mm wide. 34. P. !avacahuite
6. Leaves in fascicles of (2-)3-5(-6); seed cones ovoid.................................................................. . P leiophylla
I. Leaf fascicle sheaths persistent.
9. Bases of cataphylls (pulvini) not decurrent(Fig. IB); leaves in fascicles of 3, rarely 4, but
connate, appearingas one; seed cones on thick, long and curved, persistentpeduncles;seeds
wingless when detached from the scale ...............................................................41. P. nelsoii

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Systematic Treatment 69

9. Bases of cataphylls (pulvini) decurrent(Fig. IA); leaves in fascicles of (2-)3-5(-6), sepa-

rate; seed cones on (relatively) short peduncles;seeds winged.
10. Leaves (7-)9-15(-18) cm long; seed cones (semi-)serotinous;apophyses of seed scales
flat or only slightly raised (Fig. 6E).
II. Seed cones conspicuously pedunculate, in whorls of 1-3, (3-)4-6(-7) cm long;
leaves in fascicles of 3, but occasionally 2-5 (count 15-20 fascicles) ................ 28. P teocote
II. Seed cones appearingsessile, in whorls of 3-8, (6-)8-13(-15) cm long; leaves in
fascicles of 3..............................................................................................................32. P greggii
10. Leaves usually (except in high-altitudeP hartwegii) longer than 15 cm; seed cones
opening soon after maturity;apophyses of seed scales raised, or at least prominently
keeled (Fig. 6F-H).
12. Leaves very slender, drooping in 2 rows, 0.7-0.9(-1) mm wide, in fascicles of 3-
4(-5 )............................................................................................................................22. pa t l
12. Leaves >1 mm wide, if 0.8-1 mm, not drooping in 2 rows and predominantlyin
fascicles of 5.
13. Leaves in fascicles of (2-)3-4(-5) (count 15-20 fascicles).
14. Umbo of seed scales with a persistent spine or prickle; seed cones 8-15
cm long; leaves (18-)20-35 cm long ............................................... 12. P. engelannlii
14. Umbo of seed scales with a tiny, deciduous prickle; seed cones (4.5-)5-
10(-14) cm long; leaves 14-25 cm long....................... 1c. P arizonica var. storntilae
13. Leaves in fascicles of (3-)4-5(-6), predominantly5.
15. Leaves (6-)10-17(-22) cm long; seed scales of cones relatively thin.
slightly flexible ................................................................... 13. P h rtwegii
15. Leaves (15-)20-40(-45) cm long; seed scales of cones relatively thick,
mostly inflexible.
16. Fascicle sheaths 30-40 mm long, resinous; leaves 1.1-1.6 mm wide;
outer walls of leaf endodermalcells thin; seed cones 15-35 cm long..
........................................................................................................ 16.
16. P devonia
16. Fascicle sheaths (15-)20-30(-35) mm long, usually not resinous;
leaves 0.8-1.3 mm wide; outer walls of leaf endodermalcells thick-
ened; seed cones 7-20 cm long.
17. Leaves with connate vascular bundles; seed cones usually
obliquely ovoid when closed ......................................... 14. P pseudostrobhs
17. Leaves with mostly separatevascularbundles;seed cones usually
ovoid-oblong to ovoid-attenuatewhen closed..................... 15. P. montezuettle

Key to the species of central& southernMexico (V)

(eastern Michoacan,Mexico, Distrito Federal,Morelos, Hidalgo, Tlaxcala, Puebla, Guerrero,
Oaxaca, southernVeracruz,Chiapas)
1. Leaf fascicle sheaths deciduous.
2. Seed cones with -50, very widely spreading,very moveable scales; cones not longer than wide...........
................................................................................................................................................................43.P cembroides
2. Seed cones with 250, rigid scales, spreading-90?; cones longer than wide.
3. Leaves in fascicles of (4-)5(-6) (count 15-20 fascicles); seed cones ovoid, (4-)5-7(-8) cm long;
umbo on seed scales dorsal (Fig. 6Da, c) ...............................................................la. leiophll var. leiophl
3. Leaves in fascicles of 5, rarely 6; seed cones cylindrical, (6-)8-40(-50) cm long; umbo on seed
scales terminal(Fig. 6Ca).
4. Apophyses, at least those of proximalseed scales, recurvedor reflexed;seed cones usually 15-40
cm long. ..34. P avuacahuite
4. Apophyses of seed scales not reflexed; seed cones usually 8-16 cm long, rarely longer................
..................................................................................................................................38. P strobus var. chiapensis
1. Leaf fascicle sheaths persistent.
5. Seed cones asymmetrical,oblique at base or curved,4.5-35 cm long; seed scales opening >90?; apophy-
sis nearly flat to prominentlyraised (Fig. 6E-H).
6. Leaves relatively short, (6-)10-17(-22) cm long, spreading;apophyses of seed scales nearly flat or
slightly raised (Fig. 6E-F), light brown or purplishblack............................................................... 13. P hartwegii
6. Leaves very long, (15-)20-40(-45) cm, spreadingor drooping;apophyses of seed scales nearly flat
or prominentlyraised (Fig. 6E-H).
7. Cones 5-10(-12) cm long; falling intact (peduncle deciduous with the cone).
8. Seed scales of cones thin, flexible, in open cones usually stronglyrecurved;leaves very slender,
drooping,0.6-1(-1.1) mm wide .......................................................................................... 8. P ininoi

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70 Flora Neotropica

8. Seed scales of cones not flexible, not strongly recurved in open cones; leaves drooping or
spreading,0.7-1.2 mm wide.................................................................................. 17. P douglasiana
7. Cones (7-)10-35 cm long; leaving a few proximalscales on the branchwhen falling.
9. Fascicle sheaths 30-40 mm long, resinous;leaves 1.1-1.6 mm wide; outer walls of endodermal
cells thin; seed cones 15-35 cm long................................................................................... 16. P devoniana
9. Fascicle sheaths (18-)20-30(-35) mm long, usually not resinous; leaves 0.8-1.3 mm wide;
outer walls of endodermalcells thickened;seed cones 7-20 cm long.
10. Leaves with (partly) connate vascular bundles; seed cones usually obliquely ovoid when
closed ........................................................................................................................ 1. P. pseudostrobus
10. Leaves with usually separatevascular bundles;seed cones usually ovoid-oblong to ovoid-
attenuatewhen closed...................................................................................................15. P rnontezum ae
5. Seed cones symmetrical,ovoid or ovoid-oblong, sometimes slightly oblique, (4-)5-
10(-12) cm long; seed scales usually spreading<90?; apophysis flat or slightly raised
(Fig. 6E-F).
11. Cones broadlyovoid to subglobose when closed.
12. Cones semi-serotinous (only the distal seed scales parting), remaining on
the tree; leaves 0.8-1.6 mm wide; resin ducts in the leaves septal..........20. P oocarpa
12. Cones opening completely when mature,falling after 1-3 years; leaves 0.7-
1(-1.3) mm wide; resin ducts in the leaves medial............................24. P tecunumanii
[Link] ovoid to ovoid-attenuatewhen closed.
13. Leaves with (6-)9 or more lines of stomata on the abaxial face; leaf resin
ducts internalor internaland medial.
14. Cones semi-serotinous, persistent, leaving some basal scales on the
branchwhen falling; umbo of seed scales flat or depressed.............. 27. P pringlei
14. Cones opening at maturity,falling soon with peduncles attached;umbo
of seed scales pyramidal,prominentlyraised ................................... 26. P [Link]
13. Leaves with 3-7(-10) lines of stomataon the abaxial face; leaf resin ducts
only medial.
15. Leaves (7-)10-15(-18) cm long, 1-1.4 mm wide............................... 28. P teocote
15. Leaves (11-)15-25(-30) cm long, 0.7-0.9(-1) mm wide .....................22. P patula

Key to the species of Mesoamerica(VI)

(Chiapas, QuintanaRoo, Belize, Guatemala,El Salvador,Honduras,Nicaragua)
1. Leaf fascicle sheaths deciduous.
2. Apophyses, at least those of proximal seed scales, recurvedor reflexed; seed cones usually 15-40 cm
long........................................................................................................................................................ 34. P avacahuite
2. Apophyses of seed scales not reflexed; seed cones usually 8-16 cm long, rarely longer...........................
............................................................................................................................................38. P strobus var. chiapensis
I. Leaf fascicle sheaths persistent.
3. Long shoots usually multinodal;vegetative buds usually resinous; leaves in fascicles of 3 (rarely 2 or
4); leaf resin ducts internal ...................................................... c. P caribaea var. hondurensis
3. Long shoots uninodal;vegetative buds usually not resinous;leaves in fascicles of (3-)4-5, rarely6; leaf
resin ducts variously positioned.
4. Seed cones asymmetrical,oblique at base or curved, 4.5-35 cm long; seed scales opening to >90?;
apophysis nearly flat to prominentlyraised (Fig. 6E-H).
5. Leaves relatively short, (6-)10-17(-22) cm long, spreading;apophyses of seed scales nearly flat
or slightly raised (Fig. 6E-F), light brown or purplishblack .................................................... 13. P hartwegii
5. Leaves very long, (15-)20-40(-45) cm, spreadingor drooping; apophyses of seed scales nearly
flat or prominentlyraised (Fig. 6E-H).
6. Cones (4-)5-10(-12) cm long; falling intact (peduncle deciduous with the cone); seed scales
thin, flexible, in open cones usually strongly reflexed; leaves very slender,drooping, 0.6-1
(-1. ) mm wide ..................................................................................................................... 18. P maximinoi
6. Cones (7-)10-35 cm long; leaving a few proximal scales on the branch when falling; seed
scales thick, rigid, usually not recurvedor reflexed; leaves 0.8-1.6 mm wide.
7. Fascicle sheaths 30-40 mm long, resinous;leaves 1.1-1.6 mm wide; outer walls of endoder-
mal cells thin; seed cones 15-35 cm long ........................................................ P. 16. devoniana
7. Fascicle sheaths 20-30(-35) mm long, usually not resinous; leaves 0.8-1.3 mm wide; outer
walls of endodermalcells thickened;seed cones 8-20 cm long.
8. Leaves with (partly)connate vascularbundles;seed cones usually obliquely ovoid when
closed. .................................................................................................................... 14. P. pseudostrobus
8. Leaves with usually separatevascularbundles;seed cones usually ovoid-oblong to ovoid-
attenuatewhen closed................................................................................................ 15. P montezumae

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Systematic Treatment 71

4. Seed cones symmetrical, ovoid or ovoid-oblong, sometimes slightly oblique,

(3-)5-10(-12) cm long; seed scales usually spreading <90?; apophysis flat or
slightly raised (Fig. 6E-F).
9. Cones broadlyovoid to subglobose when closed.
10. Cones semi-serotinous (only the distal seed scales parting), remaining
on the tree; leaves 0.8-1.6 mm wide; leaf resin ducts septal .............20. P oocarpa
10. Cones opening completely when mature,falling after 1-3 years; leaves
0.7-1(-1.3) mm wide; leaf resin ducts medial..............................24. P tecunumanii

Key to the species of the Caribbean(VII)

(southernFlorida,Bahamas,Turks-CaicosIslands, Hispaniola,Cuba)
1. Leaves in fascicles of 2, rarely 3; resin ducts in the leaves septal (Fig. 2L); umbos on seed scales without
a prickle. ili.........................................................................................................................................................2.
P tropicalis
1. Leaves in fascicles of 2-4(-5); resin ducts in the leaves internal(Fig. 2H-I); umbos on seed scales with a
deciduous or persistentprickle.
2. Leaves (6-)10-15(-18) cm long; seed cones usually 4-7 cm long; seeds with adnate wing (Fig. 61). 7. P cuhensis
2. Leaves (11-)14-28(-30) cm long; seed cones 5-13 cm long; seeds with articulatewing (Fig. 6K-L).
3. Vegetativebuds not resinous; leaves in fascicles of 2-3 (count 15-20 fascicles) ............ 6. P elliottii var. densa(
3. Vegetativebuds usually resinous;leaves in fascicles of (2-)3-5.
4. Long shoots multinodal;fascicle sheath of newest leaves 15-20 mm long; leaves in fascicles of
(2-)3(-4 ), very rarely 5.................................................................................................................... c ri ea
4. Long shoots uninodal;fascicle sheaths of newest leaves (8-)10-15 mm long; leaves in fascicles
of 3-5 ........................................................................................................................................... occident lis

I. Pinus subgen. Pinus (pulvini). Cataphylls usually persistent, becoming

recurvedon shoots, falling with the leaf fascicles or
Pinus subgen. Pinus (spp. 1-33)
[Link] sheaths mostly persistent
Pinus I. Pinus MUnchhausen,Hausvater 5: 215. 1770; (deciduous in two
species). Leaves amphistomatic,
Pinus [group] Pinus Sweet, Hort. Brit., ed. 2, 475.
with serrulate margins; resin ducts variously posi-
1830, nom. nud.; Pinus Isubgen.] B. Pinus Endlicher,
Syn. Conif. 137. 1847; Pinus subgen. Pinus Parlatore tioned;
stele with outercell walls of endodermisthin
in Alph. de Candolle, Prodr. 16(2): 364, 378. 1868; or thickened,containingtwo vascularbundles,which
Pinus sect. Diploxylon Koehne, Deut. Dendrol. 30. are separateor (partly)connate. Pollenlcones (long)
1893; Pinus subgen. Pinaster Lemmon, Cal. State when mature,crowded near the proximal
Board Forest. Bienn. Rep. 2: 71, 85. 1888; Lemmon, cylindrical
[Link] Pacif. Slope, ed. 3, 24. 1895; Pinus end of new long shoots; microsporophyllsradially
div. CrassisquamaeMasters,J. Linn. Soc. London,Bot. striatedistally and with erose-denticulateuppermar-
35: 570. 1904; Pinus subgen. Diploxylon (Koehne) gin. Seed cones often persistent, symmetrical or
Rehderin Bailey, Cult. Evergr.311. 1923; Pinus subgen.
oblique. Seed scales numerous,with complex phyllo-
Eupitys (Spach) Rehder, Biblio. Cult. Trees 35. 1949; taxis;
Pinus subgen. Diplopinus Campo-Duplan,Trav. Lab. apophysis usually transverselykeeled; umbo
[Link] T. 2, sect. 1, 4, art. 1: 94. 1950, nom. dorsal, with or without a spine or prickle. Seeds
nud.; Pinus subgen. ScleropitysIshii, Kochi Univ. Nat. commonly with an articulate(rarely adnate) but ef-
Sci. Reports 2: 113, 125. 1952, sine diagn. lat.; Pinus fective wing which is largerthan the seed.
subgen. Eupinus Gaussen, Trav. Lab. Forest. Toulouse
T. 2, sect. 1, I, part 2: 94. 1960, sine descr. lat.
Lectotype species (vide Britton, N. Amer. Trees 5. 1. Pinus leiophylla Schiede ex Schlechtendal &
1908). Pinus sylvestris Linnaeus,Sp. PI. 2: 1000. 1753. Chamisso, Linnaea6: 354. 1831.
Pinus subgen. Pinea (Endlicher) Landry, Nat. Canad.
101(5): 774. 1974. Holotype species. Pinus pinea Lin- Tree, medium to tall, height to 15-30(-35) m,
naeus, Sp. PI. 1000. 1753. dbh to 20-85 cm. Trunkmonopodial, erect, terete,
Pinus subgen. SabinianaE. Murray,Kalmia 13: 18. 1983.
sometimes forked higher up, often clear of branches
Holotype species. Pinus sabiniana Douglas ex D. Don
in Lambert,Descr. Pinus, ed. 8, 2: p. s.n. inter 144 et for 3/5 of total height. Bark very thick, rough, scaly,
145, t. 80. 1832. divided into elongated, irregular,scaly plates and
deep, mainly longitudinalfissures,dark grey-brown,
Bark thin and scaly on branches and stems of on young trees and branchesscaly, flaking, reddish
young trees, soon variously furrowed or plated and brown to grey-brown. Branches long, slender,
scaly, exfoliating. Long shoots uninodal or spreadingor ascending,the lower ones curveddown,
multinodal, rough, with long decurrent leaf bases forming an open, rounded crown. Shoots more or

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72 Flora Neotropica

less scaly, with short, decurrent pulvini, initially dorsal, pyramidal, transverse-rhombic in outline, up
reddish brown or glaucous, soon grey-brown. Cata- to 5 mm wide, with a blunt prickle, darker than the
phylls small, ca. 6 mm long, lanceolate-subulate, apophysis. Seeds obliquely ovoid, slightly flattened,
soon recurved,light orange-brownwith hyaline-cili- small, 3-4(-5) mm long, dark grey-brown with black
ate margins,early deciduous. Vegetativebuds ovoid- spots. Seed wings articulate, effective, held to the
conical, obtuse, the terminalbuds 10-15 x 5-7 mm, seed by two claws, the membrane partly covering
the laterals smaller, not or slightly resinous, light the seed on one side, obliquely oval, 10-18 X 4-8
brown; the scales imbricate, subulate, with ciliate mm, yellowish brown, translucent, with a dark tinge.
margins and free apex. Fascicle sheaths 12-20 mm Cotyledons or seedlings not observed. Adventitious
long, with 5-6 initially imbricate, ligulate, ciliate- shoots sprout from the trunk or stump after fire or
margined, early deciduous scales, leaving the full- cutting and may develop into a small tree.
grown fascicles devoid of a sheath. Leaves in fasci- Distribution (Map 1). SW United States (SE Ari-
cles of (2-)3-5(-6), spreading in lax or rigid tufts zona, SW New Mexico); in Mexico along the Sierra
near the ends of branches, persisting 2-3 years, Madre Occidental, the "Eje Volcanico Transversal"
sometimes subpendulous,straight,variablein length and in the Sierra Madre del Sur as far SE as the
on the same branch,(4-)6-15(-17) cm X 0.5-1.3(- highlands of central Oaxaca.
1.5) mm, the margins (minutely) serrulate,acute or
acute-pungent, light green or glaucous-green. Sto-
Key to the varieties of Pinus leiophylla
mata on all faces of leaves, in (3-)4-8(-9) lines on
the convex abaxial face, in (2-)3-4 lines on each 1. Leaves in fascicles of (4-)5(-6), 0.5-0.9 mm wide,
with (3-)4-6(-7) lines of stomata on the abaxial
adaxial face, generally correlatedwith the width of face; numberof resin ducts (1-)2-3(-4)................
the leaf. Leaf anatomy: Cross section triangular, .............................................................
eio a. va .lla ph
with a convex abaxial side; hypodermis 1-2-layered, 1. Leaves in fascicles of (2-)3(-4, rarely 5), 0.9-
rarelythickerand intrudinginto the mesophyll;resin 1.3(-1.5) mm wide, with (4-)5-8(-9) lines of sto-
ducts (1-)2-6(-7), medial, occasionally 1-2 internal; mata on the abaxial face; number of resin ducts
(3-)4-6(-7) ................................... Ib. var. chiiuahuah na
stele terete or slightly oval in cross section; outer
cell walls of endodermisnot thickenedor thickened;
la. Pinus leiophylla Schiede ex Schlechtendal &
vascularbundles2, distinctbut connateor with partly
Chamisso var. leiophylla. Type. Mexico. Veracruz:
merged xylem strands. Pollen cones crowded near Cruz Blanca, 28 Nov 1828, Schiede 1109 (holotype,
the proximal end of a new shoot in a short, dense
HAL) Fig. 16A-F
cluster,subtendedby large, subulate,scariousbracts,
Pinus gracilis Roezl, Cat. [Link]. Mexic. 21. 1857;
cylindrical, 10-15(-20) x 4-6 mm, yellowish pink, Pinus comnonfortii Roezl var. gracilis (Roezl) Carriere,
maturingto light brown. Microsporophyllswith rela- Trait6G6n. Conif., ed. 2, 2: 565. 1867. Type. Mexico.
tively large, conspicuous pollen sacs. Seed cones "Habite le revers des cordilleres du c6t6 du Pacifique,
subterminal,solitary or in whorls of 2-5 on stout, a une altitude de 9500 a 10.000 pieds," 1857, Roezl
10-20 mm long, spreading or recurved peduncles, s.n. (lectotype, Fl, here designated).
Pinus comonlfortiiRoezl, Cat. Grain. Conif. Mexic. 22.
persistinga few years after seed [Link] 1857. Type. Mexico. Mexico: Huixquilucan, 1857,
cones broadly ovoid to subglobose, with pungent Roezl s.n. (lectotype, Fl, here designated).
spines, initially pink, later purplishgreen to purplish Pinus decandolleana Roezl, Cat. Grain. Conif. Mexic.
brown,maturingin threeseasons. Maturecones (nar- 22. 1857. Type. Mexico. Locality not stated, 1857,
Roezi s.n. (lectotype, P, here designated; isolectotype,
rowly) ovoid when closed, nearly symmetrical,
Fl).
(broadly) ovoid when opened, (4-)5-7(-8) X (3-) Pinus huisquilucaensisRoezl, Cat. Grain. Conif. Mexic.
4-5.5 cm when open. Seed scales ca. 50-70, parting 22. 1857. Type. Mexico. Mexico: Huixquilucan. 1857,
gradually,eventually spreadingwide, straightor re- Roezl s.n. (lectotype, FI, here designated).
curved nearthe base of the cone, oblong, with nearly Pinus monte-allegri Roezl, Cat. Grain. Conif. Mexic.
22. 1857; Pinus comonfortii Roezl var. monte-allegri
straightmargins,darkpurplishbrownturninggreyish (Roezl) Carriere, Trait6 Gen. Conif., ed. 2, 2: 566.
on the abaxial side, purplish brown with lighter, 1867. Type. Mexico. Michoacan: Near Xochiltepec at
greyish marks of seed wings adaxially. Apophysis Zitacuaro, 1857, Roezl s.n. (lectotype, Fl, here desig-
mostly uniformaroundthe cone, raised, transversely nated).
keeled, with the central section of the second sea- Pinus verrucosa Roezl, Cat. Grain. Conif. Mexic. 22.
1857; Pinus decandolleana Roezl var. verrucosa
son's growth distinctly marked as a narrow band
(Roezl) Carriere,Trait6G6n. Conif., ed. 2, 2: 570. 1867.
around the umbo, rhombic in outline, up to 15 mm Type. Mexico. Huixquilucan ("Huisquiluca"), 1857,
wide at mid-cone, dull brown to grey-brown. Umbo Roezl s.n. (lectotype, Fl, here designated).

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Systematic Treatment 73

Ma1Pu IepIavrleo/ila(ice)Plcol nvrcieahaa(rage)

A~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~'S

tth p

4 3 3
Map 1. 8.* is;
lelll (I -.. . . . I . I (a
Map

Pinus cedrus Roezl, Cat. Grain. Conif. Mexic. 23. 1857. reddishbrown, sometimes glaucous. Leaves in fasci-
Type. Mexico. Mexico: On a hillside nearthe road from cles of (4-)5(-6), 4 more often than 6, (6-)8-15
Mexico to Cuernavaca, 1857, Roezl s.n. (lectotype. P,
here designated: isolectotype, Fl).
(-17) cm X 0.5-0.9 mm. Stomata in (3-)4-6(-7)
Pinus dependens Roezl, Cat. Grain. Conif. Mexic. 23. lines on the convex abaxial face, in (2-)3-4 lines on
1857. Pinus decandolleana Roezl var. dependens each adaxial face. Resin ducts in the leaves (I-)2-
(Roezl) Carriere, Traite Gen. Conif., ed. 2, 2: 569. 3(-4), medial, occasionally I internal.
1867. Type. Mexico. Mexico: Near rd. from Mexico Distribution and ecology. Mexico: In NE Sonora,
to Cueravaca, 1857, Roezi s.n. (lectotype, P, here
W Chihuahua,Durango, Nayarit,Zacatecas, Jalisco,
designated;isolectotype, Fl).
Pinus lerdoi Roezl, Cat. Grain. Conif. Mexic. 23. 1857. Michoacan, Mexico D.F, Hidalgo, Morelos, Tlax-
Pinus comonfortii Roezl var. lerdoi (Roezl) Carriere. cala, Puebla, Veracruz,Guerreroand Oaxaca. In the
Traite Gin. Conif. ed. 2, 2: 567. 1867. Type. Mexico. north it is gradually replaced by var. chihuahuana,
Mexico: Sierra de Ajusco, 1857, Roezl s.n. (lectotype, which is presumablythe only taxon occurringin the
P, here designated; isolectotype, FI).
Pinus lumholtziiB. L. Robinson& [Link]- United States (see Perry, 1991). Pinus leiophylla var.
Ila Carvajal,Phytologia 59: 135. 1986. Type. Mexico. leiophylla is a widespreadconstituentof montaneto
Jalisco: San Martinde Hidalgo, Rancho El Cobre, 16 high-montane pine and pine-oak forests on deep,
Sep 1983, Carvajal 4031 (holotype, CREG). well-drainedsoils derived from various, but usually
Tree, usually tall, height to 20-30(-35) m, dbh to volcanic or metamorphic,rock. Its altitudinalrange
50-85 cm. Trunk monopodial, erect, straight. Shoots is (1500-) 19002900(-3300) m, graduallyincreasing

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74 Flora Neotropica

'i)1,

FIG. Pinus
16.A-F. var.'
leiophllaeiphylla 100EF,F
,Alman
(A-D, Padilla
Higman, &Stles [Link]
38). with

'
Ia xtwy^f^. \ B E F ,

young leaf fascicles. B. Shoot with leaf fascicles and pollen cones. C. Leaf fascicle. D. Cross-section of leaf. E. Ovulifer-
ous cones. F. Seeds. G-J. P leiophYllavar. chihuahuana(S. S. White3409). G. Branchwith leaf fascicles and ovuliferous
cone. H. Leaf fascicles with sheaths. I. Leaf fascicles. J. Cross-section of leaf. K. Seed scale (three views). (Magnifica-
tions: A-C,
tions: A-C. E---,
E-G, 1, K,
K, x 0.5; D,
XO.5; J, xX40;
D, J, 40; H,
H, x 2.)
x2.)

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Systematic Treatment 75

from north to south. It occurs most frequently with (FHO, M). TLAXCALA:
Conchatera,Aug 1901, Rose & Ha\
Quercus spp. and/or with Pinus patula, P pringlei,
5968 (A, K, NY,US). VERACRUZ:LasVigas,nearPerote,
20 May 1899,Pringle8182 (A, BM, E, F, GOET,JE, K.
P teocote, P lawsonii, P pseudostrobus, P. montezu-
M, MO, NY, P). UNKNOWN: "SanJose del Oro,"1839,
mae, P. douglasiana, P durangensis, and, at lower Ehrenbergs.n. (P).
altitudes, P oocarpa. In the NW of its range, P
arizonica, P engelmannii, and R leiophylla var. chi- lb. Pinus leiophylla Schiede ex Schlechtendal &
huahuana grow commonly with it. Locally, Juni- Chamisso var. chihuahuana (Engelmann) G. R.
perus spp. or Cupressus lusitanica are found with it. Shaw, [Pines Mexico] Publ. Arnold Arbor. 1: 14.
Annual precipitation varies greatly with locality and 1909. Fig. 16G-J
altitude, from a low of ca. 700 mm to 1950 mm. In
the north and at high altitudes, frost and snow are PinuschihuahuanaEngelmann in Wislizenus,
[Link]
N. Mexico 103. 1848; Pinus leiophyllaSchiedeex
common in winter. It is one of the few pines with
Schlechtendal
& [Link] (Engel-
a capacity to coppice. Phenology: Time of pollen mann)E. Murray, Kalmia12:23. [Link].
dispersal not recorded, it is probably variable; a few Chihuahua: Oct 1846, Wislizenus232
Cosiquiriachi,
herbarium specimens suggest March in the south of (holotype,MO 1817023,andcone MO 1635446).
the range. The ovuliferous cones take three seasons
Tree, medium size, height to 15-25 m, often not
to mature, which is exceptional in pines.
more than 10 m, dbh to 20-60 cm. Trunkmonopo-
dial, erect, often forked higher up. Shoots reddish
Representative specimens examined. MEXICO. CHI- brown, often glaucous. Leaves in fascicles of (2-)
HUAHUA:Madera,Rio Negro, 12 Jun 1958, Matuda32677
3(-4, rarely 5), (4-)6-12(-14) cm X 0.9-1.3(-1.5)
(FHO, MEXU); San Vicente, 30 Jun 1892, Hartman 538 mm. Stomata in (4-)5-8(-9) lines on the convex
(A, NY, UC, US); Arroyo Ancho, 7 Jun 1929, Mex(a
2607 (P); Mt. Mohinora, I Sep 1898, Nelson 4907 (F). abaxialface, in 3-4 lines on each adaxial face. Resin
DURANGO: 13 km N of Coyotes railway sta., 4 Nov 1970, ducts in the leaves (3-)4-6(-7), medial,occasionally
Breedlove 18726 (CAS, ME); El Salto, 30 Apr 1987, 1-2 internal.
Hughes 978 (FHO, MEXU); Lagunadel Progreso,46 km Distribution and ecology. United States: SE Ari-
N of Coyotes railway sta., 8 Aug 1955, [Link] 8380
(FHO, MICH);Quebradade San Juan,34 km N of Coyotes zona, SW New Mexico. Mexico. Along the Sierra
railway sta., 9 Aug 1955, Mavsilles 8435 (FHO, MICH). MadreOccidentalin NE Sonora,W Chihuahua,Du-
GUERRERO: 2 km E of Malinaltepec,2 Jun 1981, Almazdn rango, Nayarit,N Jalisco, and [Link] leio-
100 (FHO); along rd. from Chilpancingo to San Vicente,
phylla var. chihuahuanawas included with the spe-
25 Mar 1993, Higman. Padilla & Styles 60 (EAP, FHO,
cies in the distributionmap given by Critchfieldand
HEH, MO). JALISCO: Tapalpa,9 Oct 1892, Ellis 30 (MO);
10 km from CiudadG6zman, 21 Feb 1980, Stead & Styles Little (1966); it has been recognized as a distinct
528 (FHO, MO); 12 km from Ciudad Guzman, 16 Mar species by Perry (1991), who gives a generalized
1974, Styles 89 (FHO). Mexico: Desierto de Le6n, 2 Apr map roughly in accordance with our findings. Al-
1950, Matuda 19495 (FHO, MEXU, MO); Juchitepec, 15
thoughlargely sympatric,the altitudinalrangeof var.
Apr 1982, Lorea 1969 (FHO);Mt. Ixtaccihuatl,9 Jan 1894, chihuahuana is narrower- 500-2700(-2950) m-
Nelson & Goldmtans.n. (A, E, US); Sierra de Ajusco, 2
Nov 1895, Pringle 6180 (A, BM, E, F, GOET, JE, K, M. but this is in partdue to the fact that it does not occur
MO, NY, S). MICHOACAN: along Hwy. 37 to Uruapan,ca. on some of the high volcanos of central Mexico. In
5 km S of Carapan,6 May 1994, Farjon & Mejia 349 (E, the Sierra Madre Occidental its lower limit usually
FHO, HEH, K, MEXU, U); Cerro Patzcuaro,Nov 1890, is below the level of var. leiophylla. Here it may
Pringle 3377 (F, GH, MICH), Nov 1890, Pringle 4171 (F,
MICH, NY, UC, US), 11 Oct 1892, Pringle 4358 (BM, F, be found in semi-arid habitat with P. cembroides,
GH, GOET, JE, K, M, MO, P, S, UC, US); Tajimaroa Juniperus spp., and xerophytes like Opuntia and
("prope Tajimaroaprovinciae Michoacensis"), Dec 1829, [Link] up, its habitat is similar to
Schiede 1117 (HAL); San Felipe, near Zitacuaroalong rd. that described above. Its poor growth in some areas
to Angangueo, 8 Feb 1980, Stead & Styles 462 (FHO); 5
km N of Uruapan, 22 Feb 1974, Styles I (FHO, K).
is related to lower precipitation and rocky, poor,
OAXACA:Along rd. from Oaxaca to Tuxtepec (Hwy. 175), shallow soils, especially at lower and middle alti-
near San Juan Atepec, 23 Apr 1994, Farjon & Mejia 315 tudes. Phenology:The time of pollen dispersalis not
(E, FHO, HEH, K, MEXU, U); La Carbonera, 18 Mar recorded.
1993, Higman, Padilla & Styles 38 (EAP, FHO, HEH,
MO); National Park Benito Juarez,20 Mar 1993, Higman, Representative specimens examined. MEXICO. CHI-
Padilla & Styles 41 (EAP, FHO, HEH, MO); Miahuatlan, HUAHUA:Along rd. betweenCreeland La Bufa, 12 Sep
22 Mar 1993, Higman, Padilla & Styles 50 (EAP, FHO, 1957,Knobloch410 (MEXU);Cerrode la Ventana,16km
MO, ESNAC, HEH); 1841, Liebmann 14791 (MO, NY); NWof Minaca,10May1929,Mexia2514(CAS,E, MICH,
Llano Verde,E of Oaxaca, 4 Mar 1985, McCarter& Styles MO,NY, P); aboveArroyoAnchoon Mesade Gonzalez,
344 (FHO). PUEBLA: Paso Cortez, 17 Jul 1977, Mittak8507 8 Jun 1929,Mexia2620 (CAS, E, MICH,MO, NY. P):

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76 Flora Neotropica

YerbaBuenaaboveBatopilas,Nov 1885,Palner301 (GH); titious ("epicormic")shoots along the trunkof mature

SierraLa Silla (SierraMapula),5 Nov 1886,Pringle959 trees.
(BM,F, MEXU,NY);CascadaBasaseachic,11Nov 1989,
Carvajal (1986) described Pinus lumlholtziivar.
Spellenherget al. 10064(FHO,MEXU,NMC,ESAHE),
12 Nov 1989, Spellenberget al. 10081 (FHO, ID, MEXU, microphylla based on Carvajal 4031 (CREG!) but
NMC); Cascada Basaseachic, 12 Nov 1989, Spellenberget also referringto materialcollected tentatively as P
al. 10090 (FHO, NMC);ColoniaGarcia,1 Oct 1899, chihuahuana. His conclusion, that it resembles P
Townsend& Barber359 (BM, E. F, GH, K, MO,NY, P, lumholtziimore,
NW of El Salto, 12 Jul 1898, Nelson 4558 appearsto have been withdrawnin
UC). DURANGO:
1896, Palmer 83 Carvajal & McVaugh, 1992: 71, where its type is
(A, K, US); SantiagoPapasquiaro, Apr
(K, MO);nearDurango, Apr1896,Palmer774(K),Palher synonymizedwith P leiophylla [Link]. Pinus
775 (A, BM, F, K, MO,NY, US); Durango.2 Apr 1991, lumholtzii has sometimes short, non-pendulous
Stles, Favela et al. I (FHO). NAYARIT: SantaTeresa, 1897, leaves and can in those cases become confused with
Rose 2123B (A). SONORA:Region of the Rio de Bavispe,
P leiophylla var. chihuahuana(Martinez, 1948; see
LasTieritasdel Tamblor, 20 Aug 1940,Phillips659 (GH);
SierraEl Tigre, 18 Aug 1940, White3409 (FHO,GH, also the treatmentof P lumholtzii,this volume).
MICH);regionof the Rfode Bavispe,Ranchode la Nacha The numerous "species" described or named by
SE of El Tajo, 15 Aug 1941, White3942 (GH). ZACATECAS: Roezl, most of which were synonymizedby Parlatore
16 kmSE of Chalchihuite, 25 Jul 1982,Diggs& Nee 2997
(1868), have been typified for the first time from
(F. FHO, XAL); Sierrade los Morones,near Plateado. Roezl's materialin Fl and P. A
1897. Rose 2742A (GH). general comment on
Roezl and his pines is given under the treatmentof
Uses. Along with many other species of pine in Pinus montezumae.
the Sierra Madre Occidental, Pinus leiophylla has In the Botanical Appendix to Wislizenus (1848),
been heavily exploited for timber in the latter half Engelmann described three new species of pines
of this century. Because of its high resin content, it "entirely different from those found farther north,
does not provide high-qualitywood but at the same but perhaps identical with some species from the
time is a producer of good resin, for which it is Pacific coast." One of them is Pinuischihuahuana.
[Link] variety chihuahuana,an equally prolific No comparisonswere made with other pines already
producerof resin, is less exploited for timber. described from the Sierra Madre Occidental and
Under Coniferae, Schlechtendal and Chamisso occurringfarthersouth. In his revision of the genus,
(1831) describedtwo new species of Pinus, P patula Engelmann(1880a) classified P leiophylla in a dif-
and P leiophylla, based on Schiede (& Deppe) 1108 ferent subgroup from P chiuahuahuana, although he
and 1109 respectively, and both from (approxi- observed its deciduous fascicle sheaths. [Hartveg
mately) the same locality. Schiede (& Deppe) 1109 441 does indeed have persistent sheaths, but it be-
consists of a single sheet with two sterile branchlets longs to P teocote, not to P leiophylla, where Engel-
at HAL, which is the holotype of P leiophylla. mann (1880a) and Parlatore (1868) cited it]. Not
Deppe, mentioned as the second collector in the until Shaw (1904b, 1909) recognized the prolonged
protologue, does not appear on the labels. Another phenology as well as the sharedcharacterof adventi-
collection in HAL, Schiede 1117, bears a small cone, tious shoots and coppicing capacity were P. leiophy-
but in the protologue it is claimed no cones were lla and P chihuahuanacorrectly interpretedas con-
observed ("nobis deficiente"). Although meagre, specific.
Schiede 1109 is unambiguousand need not be sup- Engelmann(in Wislizenus, 1848) did not indicate
plemented with Schiede 1117 as an epitype; but as which of the collections of Wislizenus pertains to
another specimen annotated by Schlechtendal, the his new species, and neither does Parlatore(1868),
latterhelps to interprethis taxon more fully. Lambert who received some materialon loan from Engelmann
(1832, t. 21) had the species drawn for his third (see Carvajal& McVaugh, 1992: 69), as is annotated
edition from a specimen with a cone received from by Engelmannon the sheet of Wislizenus232 (MO
the collector(s). 1817023). This sheet and a cone (MO 1635446),
Shaw (1904a) was the first author to describe the returned from the loan to Parlatore,are to be consid-
prolonged growth of the ovuliferous cone, a feature ered the holotype of P chihuahuana.
shared by P pinea of the Mediterraneanbut unique Martinez (1948), who maintained both taxa at
in Latin American pines. It shares this feature with the rank of species, nevertheless admitted that P
its variety, P leiophylla [Link], chihuahuanain the southernpartof its range merges
the ability to coppice, shared by P rigida of the with P leiophvlla to such an extent that it becomes
eastern United States, is an uncommonphenomenon very difficult to distinguish them. We, too, have
presentin both varieties.A relatedfeatureare adven- found this to be the case: and since there seem to

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Systematic Treatment 77

be no other than quantitative foliar characters to lines on the convex abaxial face and in 6-8 lines on
differentiatethem, while they sharetwo very distinct the adaxial face (3-leaved specimens not observed).
and uniquequalitativefeatures,these combinedargu- Leaf anatomy: Cross section semi-circular (rarely
ments stronglysupporttreatmentat [Link] transverse-triangular); hypodermisuniform in thick-
the other hand, in certain areas in Durangothe two ness with 1-2 rows of cells but thicker in marginal
taxa retain their differences over large areas, where areas; resin ducts 6-10, very large, septal, or some-
typical P leiophylla with fascicles of five, slender, times a few external; stele semi-terete, large; cell
pendulous leaves grow in proximity to "pure" P walls of endodermisthin; vascular bundles 2, sepa-
leiophylla var. chihuahuana with fascicles of three, rateor the xylem strandsmore or less [Link]
thick, spreadingleaves (Styles, 1992). Furtherassess- cones crowdednearthe proximalend of a new shoot,
ment of the relationshipof these two pines and their ovoid-oblong to cylindrical, 20-30 x 5 mm, pink
putative relatives, including molecular genetics, is to yellowish, turning yellowish brown. Seed cones
desirable. subterminal,erect on short, thick peduncles, solitary,
in pairsor whorls up to 6, remainingerect or spread-
2. Pinus tropicalis P. M. A. Morelet, Rev. Hort. ing obliquely. Immaturecones (narrowly)ovoid, 10
C6te d'Or 1: 106. 1851. Type. Cuba. Isla de la x 5-7 mm, purplishred, maturingin two seasons.
Juventud[Isla de Pinos]: La Caniada,16 Feb 1916, Mature cones narrowly ovoid to ovoid-attenuate
Brittonet al. 14416 (neotype, NY, here designated). when closed, ovoid with a flattened base when
Fig. 17 opened, then 5-8 x 4-5.5 cm, persistent for some
Pinus cubensisGrisebachvar. terthrocarpa Grisebach, years but falling with the peduncle attached. Seed
[Link].217. 1866;Pinulsterthrocurpa (Grisebach) scales ca. 100-120, parting to release the seeds
G. R. Shawin Sargent,Trees& Shrubs1: 149,t. 75. except those at the base, oblong, straightor strongly
[Link],1860,Wright recurved, dark brown, with light brown marks of
3189(lectotype,[Link] marked 3189a,here seed
[Link], K, S).
isolectotypes. wings on the adaxial side. Apophysis flat or
designated:
slightly raised, transverselykeeled, rhombicto pen-
Tree, medium to large, height to 30 m, dbh to tagonal in outline, up to 12 mm wide, radially stri-
100? cm. Trunk monopodial, erect, terete, usually ated, light brown or reddish brown. Umbo dorsal,
slender, in forest stands with clear bole for /3-3/4 of flat or slightly raised, rhombic, ca. 3 mm wide,
height. Barkthick, rough, scaly, breakinginto irregu- without a prickle, greyish brown to grey. Seeds
lar, polygon plates, divided by deep longitudinal obliquely ovoid, slightly flattened,5 x 4 mm, light
fissureson the lower partof the trunk,reddishbrown, [Link] wings articulatebut effective, held
weatheringgrey, on young trees and branchesscaly, to the seed by two claws, obliquely obovate-oblong,
flaking, reddish brown. Bratches of first and higher 12-15 x 5-6 mm, yellowish with a grey or black
orderssparse, thick, spreadingto ascending, forming tinge. Seedling: Numberof cotyledons not observed.
an irregular,open crown. Shoots uninodal, thick, Suppressedterminalgrowth and a thick radicle pro-
very rough with large, prominent and persistent, duce a "grassstage"as an adaptationto groundfires.
decurrentpulvini, lustrous orange-brownin Ist and Leaves are normalbut less straight.
2nd year, then grey, scaly with flat bases of pulvini. Distribution and ecology (Map 2). W Cuba: in
Cataphylls subulate, recurved, scarious, with erose- Pinar del Rio and on the Isla de la Juventud(often
ciliate margins and caudate apex, brown, caducous. still referred to as Isla de Pinos (Isle of Pines) in
Vegetativebuds ovoid-oblong, acute, the terminal botanical literature).Pinus tropicalis is a lowland
bud 15-25 mm long (on vigorous leading shoots of pine, occurringon the coastal plains and low foothills
young trees larger),the lateralssmaller,not resinous; at 1-150(-300) m altitude,on nutrient-poorsandy or
the scales more or less spreading, reddish brown gravelly alluvial soils which are dry due to rapid
with ciliate, hyaline margins. Fascicle sheaths of [Link] climate is tropical,with annualprecipi-
juvenile leaves ca. 20 mm long, papery, yellowish tation of ca. 1200 mm and a prolongeddry season.
brown with ciliate, whitish margins, persistent but It is in partsympatricwith P caribaea [Link],
reduced to ca. 10 mm on mature leaf fascicles, which has a greateraltitudinalrange. Pine savannas
weatheringgrey. Leaves in fascicles of 2 (rarely 3), are open, grass-dominatedlowland areas which burn
spreadingobliquely,persisting2 years, very uniform, frequently; P tropicalis has an advantage over P
straightand rigid, (15-)20-30 cm X 1.5 mm, with caribaea in that it has a "grassstage" which enables
serrulate margins, acute, light or yellowish green. the seedling to survive successive fires. Thus it be-
Stomataon all faces of leaves, in 6-8 inconspicuous comes frequently the only pine in this vegetation

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78 Flora Neotropica

FIG. 17. Pinus iropicalis [a, C, D, Little 14048; B, E-H, Lambs.n. (div. coll.)]. A. Branch with leaf fascicles and two
immatureovuliferous cones. B. Seedling in "grass stage." C. Leaf fascicle. D. Cross section of leaf. E-F. Ovuliferous
cones. G. Seed scale (three views). H. Seeds. (Magnifications:A-C, E-H, X0.5; D, X25.)

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Systematic Treatment 79

Map 2. Pinus tropicalis (circles); P. cubensis (triangles);P. occidentalis (squares).

type. Phenology: Time of pollen dispersal not re- used in plantationforestry,mainly in Cuba, but also
corded; presumably later than P caribaea. to a limited extent elsewhere.
Morelet (1851) recognized from his field observa-
Representative specimens examined. CUBA. ISLADE
LAJUVENTUD [ISLADE PINOS]:Loma Vista, near Nueva
tions that there are two distinct species of pines in
Gerona, 26 Nov 1920, Ekman 12385 (K, S); 25 km WSW W Cuba, both on the "Isla de Pinos" and on the
of Nueva Gerona, 22 Jan 1951, Little 14048, 14052 (US); mainland. They were not to be equated with P
locality not indicated, 22 Mar 1905, Shaw s.n. (M). PINAR occidentalis Swartz, the only pine previously de-
DELRfo: San Vicente, 12 Sep 1910, Britton et al. 7319
scribedfrom the CaribbeanIslands,of which Morelet
(NY); Herradura,12 Feb 1906, Hermann 2653 (NY, US);
Vinales, between Pinar del Rio and San Vicente, 13 Jul said that it occurs on Hispaniola("Saint-Domingue")
1968, Lamb s.n. (FHO); Galalon, 15 Jul 1968, Lamb s.n. and E Cuba (Sierra de Cobre).
(FHO); Sandino, 30 Jun 1990, Maas et al. 7750 (U); near Grisebach(1866) describeda (tentative)varietyof
city of Pinar del Rio, 19 May 1951, Rutten-Pekelharing P cubensis as var. terthrocarpa,a name coined by
431 (U); Herradura,30 Apr 1903, Shafer 425 (K, NY);
C. Wrightat species rank on the label of his collec-
Laguna el Jovero, 12 Dec 1911, Shafer 10905 (NY, U,
el
US); Laguna Bufeo, 16 Dec 1911, Shafer 11017 (NY, tion no. 3189. Wrightonly mentioned "Cuba"as its
U, US); Herradura,9 Feb 1903, Shaw s.n. (K, MO, P, US); origin, but apparentlyit is from easternCubabecause
21 km N of Pinar del Rio, 18 Jun 1955, Schultes et al. 4 P cubensis is restricted to E Cuba. Shaw (1903),
(NY). WITHOUTLOCALITY:1856, Christy s.n. (K); 12 the GH duplicate of Wright3189 and other
May 1911, Clinton-Baker s.n. (K). citing
collections, took up the specific epithetfromWright's
Uses. Pinus tropicalis is an important regional label and published it as a new species. He main-
source of timber mainly used by local sawmills. Its tained this at first (Shaw, 1904a), but later (Shaw,
wood is dense and durable but resinous. Despite 1914) synonymized it underP. tropicalis. Engelmann
extensive exploitation, it is still common. Regenera- (1880a), who, like others, seems not to have known
tion seems to be good in many areas, especially of Morelet's repeated publication (Morelet, 1851,
where it is protected from grazing pressures. It is 1855) of two new Cuban pine species, mentioned

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80 Flora Neotropica

Pinus murrca'anaBalfour in Murray,Bot. Exped. Oregon

8: 2. 1853; Pinus contorta D. Douglas ex J. C. Loudon
subsp. murravana (Balfour) W. B. Critchfield, Publ.
Maria Moors Cabot Found. Bot. Res. 3: 106. 1957.
Type. United States. California: Siskiyou Mts., s.d.
[1852]. J. Jeffrey 740 (holotype, E; isotypes, Fl, K).

Tree, in Mexico of medium size, occasionally tall,

height to 25-30(-33) m, dbh to 100-150(-200) cm.
Trunkmonopodial, erect, massive, some trees may
be forked higher up, scrubby on summit ridges,
slender in dense stands. Barkthin, scaly, with numer-
ous small, flaking plates, colour variable from light
orange- or pink-brownto greyish brown, on young
trees thin, papery. Branches of first order relatively
short, spreading to ascending; branches of higher
orders slender and flexible, spreading or ascending,
forming a narrowly conical or pyramidal,dense or
more open crown. Shoots multinodal (especially
s~.. \..
leading shoots), lateral shoots on older trees mostly
uninodal, rough, ridged between bases of leaf fasci-
cles, more or less glaucous; pulvini [Link]-
phylls small, ligulate, curved, with erose margins,
early deciduous. Vegetative buds ovoid-conical, the
terminal buds ca. 15 x 6 mm, the laterals smaller,
resinous; the scales imbricate, appressed, acute, or-
ange-brown. Fascicle sheaths short, 5-8 mm long,
reduced to ca. 5 mm in maturefascicles, persistent,
light reddish brown with hyaline-whitish margins,
weathering grey. Leaves in fascicles of 2, in dense
tufts spreading obliquely from the shoot, persisting
Map 3. Pinus contorta var. murravana. 5-8 years, rigid, usually curved and contorted, 4-7
cm long (variable on a single branch, decreasing
Wright's name as published by Grisebach and, appar- with altitude), 1.2-2 mm wide, entire or sparsely
ently thinking that its "grass stage" was the reason for serrulate,acute-acuminate,pungent,light (yellowish)
its distinction, which he considered taxonomically green. Stomata on all faces of leaves, in 10-14
insignificant, dismissed it. grooved lines on both the convex abaxial and the
Previous classifications (e.g., Little & Critchfield, slightly concave adaxial face, conspicuous. Leaf
1969; Farjon, 1984) group P tropicalis (and P res- anatomy:Cross section semi-circularto semi-elliptic;
inosa of the eastern United States and Canada) with cuticula and epidermis thick; hypodermis uniform,
the predominantly Eurasian subsect. Sylvestres on 2-layered; resin ducts (1-)2(-?), medial toward the
grounds of morphological features of the seed cones leaf margins;stele elliptic in cross section, with outer
and leaf numbers. A phylogenetic approach using all cell walls of endodermisthickened;vascularbundles
available information, preferably including molecu- 2, distinctly [Link] cones crowded near the
lar data, should be used to determine if P. tropicalis proximal end of a new shoot, subtended by 5-8
indeed belongs in this rather homogeneous subsec- subulate bracts, ovoid-oblong to cylindrical, 8-14
tion. Unfortunately, Wang and Szmidt (1993) did mm long, yellowish to yellowish orange. Microspor-
not incorporate this species in their cpDNA-based ophylls ca. I mm diam. with a radiallystriateabaxial
phylogeny assessing many of the purportedly related surface; pollen slightly largerthan in other varieties.
Asian species. Seed cones subterminal,usually in pairs or whorls
of 3-5, on short, recurvedpeduncles, persisting only
3. Pinus contorta D. Douglas ex J. C. Loudon var. 1-3 years after release of seeds. Immature cones
murrayana (Balfour) Engelmann in Watson, Bot. ovoid, with prominent,erect spines, ca. 10 mm long,
California 2: 126. 1880. Fig. 18 reddish purple, maturing in two seasons. Mature

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Systematic Treatment 81

FIG. 18. Pinus contorta var. inurravana (A-C, E-G, Hughes & Styles 174; D, Jeffrey 740). A. Branch with leaf
fascicles and ovuliferous cone. B. Leaf fascicle. C. Cross section of leaf. D. Ovuliferous cones, closed. E. Ovuliferous
cones, opened, with lateralview of seed scale. F. Seed scale (two views). G. Seeds, with articulatewings. (Magnifications:
A, B, D-G, x 0.5; C, x20.)

cones (narrowly) ovoid when closed, symmetrical, light brown marksof seed wings on the adaxial side.
broadlyovoid with a more or less oblique base when Apophysismore or less symmetricalaroundthe cone,
opened,(3-)4-5.5 x 3-4 cm when open. Seed scales slightly raised, transversely keeled, somewhat gib-
ca. 90-1 10, partinggraduallyafter cone maturation, bous on the basal scales, irregularlypentagonal in
oblong, straight or recurved, thin woody, dark pur- outline, ochraceous,[Link], rhombic-
plish brown on the abaxial side, dark brown with pyramidal, 2-3 X 3 mm, with a prominent. thin,

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82 Flora Neotropica

curved and persistentprickle, greyish brown. Seeds yet recognized as a distinct species, it was later
obliquely ovoid, 4-5 mm long, greyish brown. Seed described by Loudon (1838) from a collection made
wings articulate but effective, held to the seed by by David Douglas in 1825 near the mouth of the
two oblique, thin claws, obliquely ovate, 8-12 X 4- Columbia River in Washington, U.S.A. (Mirov,
5 mm, yellowish brown, [Link]- 1954). The specimen at K! (D. Douglas s.n., ex herb.
dons 4-5 (number on average higher by I than in Bentham)-annotated by Douglas as "P contorta
other varieties). Sabine. ... In swampy ground near the sea coast.
Distribution and ecology (Map 3). United States: Abundantly near Cape Disappointment and Cape
Cascade Range of SW Washingtonand W Oregon; Lookout. 1825.6.7."-is here designatedas the lecto-
southward in the Sierra Nevada of California and type of Pinus contorta Douglas ex Loudon.
extreme W Nevada, disjunct in high mountainsof S Based on collections sent by J. Jeffrey under
[Link]: in Baja CaliforniaNorte, Sierra no. 740 to Edinburgh, Balfour (in Murray, 1853)
San Pedro [Link] southernCaliforniaand Mexico describedP murrayanafrom the Siskiyou Mountains
it occurs in the upper zone of the Mixed Conifer in southernOregon or northernCalifornia,accompa-
Forest, with Abies concolor and Pinusjeffreyi as the nied by a figure done by Greville (who is often
most common conifers, and also Pinus lambertiana erroneouslycited as one of the authors;in fact, the
and Populus tremuloides on deeper soil and more new species are ascribed by Murrayto the "Oregon
mesic sites, [Link] SierraSan PedroMartir Committee"of which Balfour was chair).
is a graniticbatholithwith a steep easternescarpment Engelmann(1880b) reducedthis species to a vari-
and extensive western [Link] altitudinalrange ety of P contorta (in which he included P contorta
of Pinus contorta var. murravana is here 2300- var. latifolia Engelm. from the Rocky Mountains).
3000+ m; there is no climatic tree line, but due to Apart from stressing the taller habit (especially in
steepness of the highest summit, Picacho del Diablo the Sierra Nevada, where most conifers are indeed
(3095), only a few erraticsaplings of Abies concolor taller), he describedthe longer and wider leaves and
occur there (Minnich, 1987). Annual precipitationis the less oblique, often opening and deciduous cones
a moderate500 mm, of which 50% may be snowfall. as the differentiatingcharactersagainst P contorta
Frosts are common during the winter months. Phe- [Link]., which he confined to the Pacific coast. Critch-
nology: Pollen is dispersed during May-June. field (1957) changed the rank of this and other
infraspecifictaxa to subspecies, based on allopatric
[Link]
Representative distributionand growth performancerelatedto geog-
CALIFORNIA NORTE: Sierra San Pedro Martir, at
"Vallecitos," 29 Nov 1987, Franklin & Chandler 5673 raphyand climate ("provenance").Many of his argu-
(NY), 26 Jan 1972,HallerJr. 3427 (UC), 25 May 1987, ments (Critchfield,1957, 1963, 1980) are of ecologi-
Hughes & Styles 174 (FHO), Sep 1961, Lindsay s.n. (SD), cal consequence, rather than taxonomic, and the
Sep 1940, M. Martinez 3464 (INIF. MEXU), 2 Jun 1968, morphologicaldifferencesappearto be clinal in areas
Moran15076(MEXU,SD);SierraSanPedroMartir,near
3 May [Link]. & RomeroT 13296 where these taxa meet. Some of them, such as non-
Observatory,
(MEXU). serotiny of cones ascribed to var. murrayvaa, also
occur in certain individualsof populationsbelonging
Uses. Pinus contorta is one of the most important to other infraspecifictaxa. Distinctionat varietalrank
timber trees of North America (Lotan & Critchfield, seems the most appropriatetaxonomic approach in
1990), but since it is in Mexico restrictedto the high such cases.
plateaus and ridges of the Sierra San Pedro Martir,
it is of local importancethere. Much of this disjunct
4. Pinus herrerae Martfnez,Anales Inst. Biol. Univ.
population, including the best-developed stands Nac. Mexico 11: 76. 1940.
aroundhigh-montanemeadows such as "La Encan- Fig. 19
tada,"is now in the SierraSan Pedro Martir National
PitnusteocoteSchiedeex Schlechtendal & Chamissovar.
Park and enjoys protection at least from lumber herrerae(Martinez)Silba, Phytologia68: 63. 1990.
harvesting. [Link]:SierraLalo,Feb 1940,[Link]-
Pinus contorta is highly variable genetically and nez 3427 (lectotype, MEXU; isolectotypes, F, US).
morphologically throughout its vast range, and is
only second in geographic extent to P sylvestris Tree, usually tall, height to 30-35 m, dbh to 75-
(Critchfield,1980). As a consequence, several forms 100 cm. Trunkmonopodial, straight or sometimes
of different regions have been described as species. tortuous,terete, often with a branch-freebole 2/3 or
Although first collected in Alaska (at Sitka), but not /4 its length. Bark thick, rough, with scaly plates

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Systematic Treatment 83

F -
B3B
E A

FIG. 19. Pinus herrerae (A-C, Stles 69; D-G, Hughes & St'les 137). A. Branchwith leaf fascicles and pollen cones.
B. Leaf fascicle. C. Cross section of leaf, with enlarged endodermal cells showing thick outer walls. D. Immature
ovuliferous cones. E. Matureovuliferous cones. F. Seed scale (two views). G. Seeds, with articulatewings. (Magnifica-
tions: A, B, D-G, x0.5; C, x60.)

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84 Flora Neotropica

and shallow, longitudinalfissures, reddish brown to 5-8 x 3-5 mm, translucent, yellowish with a dark
grey-brown, in young trees and on branches thin, tinge. Number of cotyledons not observed.
exfoliating in thin flakes, reddish brown. Branches Distribution and ecology (Map 4). Mainly in
long, slender, arching or spreadinghorizontally,the disjunct populations in the west and south of Mexico,
higher-orderbranches flexible, slightly pendulous. along the Sierra Madre Occidental and more abun-
Shoots smooth, with well-developed, long decurrent dantly along the Sierra Madre del Sur; in SW Chihua-
pulvini, [Link],scarious, hua, Sinaloa, Durango, W and S Jalisco, Michoacan,
recurving,brown, weakly ciliate-hyaline at margins, and Guerrero. Its altitudinal range is (1100-) 1500-
early deciduous. Vegetative buds ovoid-acute to 2600 m; its lowest altitude apparently is reached in
ovoid-oblong or cylindrical, the terminalbud 10-15 the Sierra de Cuale in W Jalisco. It occurs in the
mm long, the lateralssmaller,not resinous;the scales mesic forest belt, with annual precipitation of ca.
subulate,reflexed, weakly ciliate-hyalineat margins. 900-1600 mm but with a dry season lasting from
Fascicle sheaths of flushing leaves up to 20 mm November to May. From N to S annual precipitation
long, torn by elongating and spreading leaves; the increases. Mixed pine and pine-oak forest is its usual
scales chartaceous,yellowish white; sheaths persis- habitat, in which it is associated with many other
tent but reduced to 8-15 mm in mature fascicles, pines according to their ranges, further with Pseu-
light brown to [Link] in fascicles of 3, dotsuga locally. Other broad-leaved trees-e.g., Ar-
in open or dense tufts persisting 3 years, slender, butus, Alnus, Clethra, Juglans, Persea, Clusia, and
lax, drooping or spreading,(10-)15-20 cm X 0.7- Tilia-are sometimes common. In many areas the
0.9 mm, serrulate at margins, acute, light green to forests are greatly influenced by man-made changes
yellowish green. Stomata on all faces of the leaves, in their composition, e.g., by selective felling of
in 6-8 lines on the abaxial face and ca. 3 lines larger trees of Pinus or by recurrent fires and grazing.
on each adaxial face. Leaf anatomy: Cross section Phenology: Pollen dispersal has been observed in
transverse-triangular,with a convex abaxial side; March in Jalisco and Michoacan.
hypodermis2-3-layered, or thickerin marginalareas;
resin ducts 1-3(-4), internal,rarelyseptal; stele oval Representative specimens examined. MEXICO. CHI-
HUAHUA: Cascada Basaseachic National Park, I km S of
in cross section; cells of endodermiswith thickened the falls, 12 Nov 1989, Spellenberg et al. 10086 (FHO,
outer walls; vascular bundles 2, (widely) separated. MEXU, NMC). DURANGO: ChavarriaNuevo, 3 Apr 1991,
Pollen cones crowded along the proximal end of a Styles, Favela et al. 6 (FHO, MEXU). GUERRERO: 4 km
new shoot, subtended by short bracts, spreading, SW of Puerto El Jiguero, 25 Oct 1987, Aquino 153 (FHO,
MEXU); 3 km NE of Cruz de Ocote, 23 Mar 1985, Lorea
ovoid-oblong to cylindrical, 15-18 x 5 mm, yellow- 3483 (FHO); Sierra Madre del Sur, Las Lumbreras,3 Jan
ish green, tinged red, turninglight [Link] cones 1938. Mexial9088 (B, K, MO, NY, S, UC); Omiltemi, Las
subterminal,solitary or opposite, rarely in whorls of Trinceras,2 km N of Pablado, 6 Apr 1985, Millan Espin
3, on 10-15 mm long peduncles, at first erect, soon 582 (FHO, MEXU); Carrizalde Bravo, 8 Feb 1981, Stead,
on recurved peduncles which fall with the cones. Styles et al. 731 (FHO). JALISCO: Loc. not stated, det. by
M. Martfnezas "P herreraisp. nov.," 1938-1939, collector
Immaturecones (broadly) ovoid, 8-13 mm long, not specified, (MEXU No. 2526); SierraManantlin, 16 Apr
purplish red, maturing in two years. Mature cones 1987, Hughes & Styles 137 (FHO); Pihuamo, Mar 1939,
narrowlyovoid when closed, nearly symmetricalor M. Martinez3599 (NY); Sierra de Cuale, along rd. to the
mines E of El Tuito, 16-19 Feb 1975, McVaugh 26346
slightly curved, (broadly) ovoid when opened, (2-)
(MICH);Tecatitlan,64 km from Ciudad Guzmain,13 Mar
3-3.5(-4) x 2-3.5 cm when open. Seed scales ca. 1974, Styles 67 (FHO, K), 13 Mar 1974, Styles 68, 69
50-80, partingexcept those at the base of the cone, (FHO). MICHOACAN: Dos Aguas, 24 Sep 1960, Courbas-
thick woody, oblong, straight or recurved, on the son & Mahieux s.n. (P); ca. 3.5 km S of Paracho de
abaxial side purplish, greyish at base, with light Verduzco,at km 35 along Hwy. 37, 6 May 1994, Farjon &
brown marksof seed wings and a darkerbrowndistal Mej'a 351 (E, FHO, HEH, K, MEXU, U); Dos Aguas,
along rd. to Coalcoman, 11 Apr 1985, Soto N. et al. 8144
part adaxially. Apophysis slightly raised, mostly so (EAP, NY); Dos Aguas, forest station W of Aguililla, 2
on a few proximal scales on one side of the cone, Mar 1974, Styles 41 (FHO). SINALOA: Mun. Rosario, ca. 6
transverselykeeled, the apical marginentire or cre- km W of El Palmitoalong Hwy. 40, 31 Jan 1962. Breedlove
nate, radiallystriate,light brown. Unbo dorsal,pyra- 1704 (CAS); Mun. Badiraguato,5 km N of Los Ornos
along rd. to Ocurahui, 2 Oct 1970, Breedlove & Thorne
midal, irregularlyrhombicin outline, 3-4 mm wide, 18361 (CAS); Los Loberos,along main rd. between Mazat-
with a minute,deciduous prickle,greyish. Seeds (ob- lin and Durango,30 Apr 1987, Hughes 970 (FHO, MEXU).
)ovoid, slightly flattened, 2.5-4 X 2-3 mm, dark
[Link] wings articulate,effective, held to Uses. Along with other tall growing pines, it is
the seed by two small, thin claws, obliquely ovate, (heavily) exploited for timber throughout almost all

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Systematic Treatment 85

;'~~

'' 1.. ~ ~ ~ ~ ~ .8 S..,. ""

-?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
*
?
..,... ....
iiii i ii i

Map
[Link]~~~~~~;rrre(ice)[Link].

a; . ?" Z' .

p4 . ....

of its range. It is considered to be of good wood species. Perry(1991) noted virtuallythe same differ-
[Link] has been startedand is ences between the two taxa. Molecular evidence
recommended by foresters (Dvorak & Donahue, (cpDNA) may providenew, non-morphologicalchar-
1992) to be conducted on a largerscale. acters to more firmly establish (or refute) species
This species was, according to Martinez (1940), delimitations, but, apart from P patula (Perez de la
first collected in Jalisco in 1931 and again in 1938- Rosa et al., 1995), no molecular data are as yet
1939 as P. teocote and [Link], [Link]- available for these species.
field and Little (1966) did not recognize it and
mappedit with P. teocote. Pinus herrerae is in many 5. Pinus caribaea Morelet, Rev. Hort. C6te d'Or 1:
characterssimilar to P teocote, yet distinct in a few 107. 1851.
constant characters, most notably the very slender,
lax, and (slightly) longer leaves and the smaller Tree, medium to tall, height to 20-35(-45) m, dbh
ovuliferouscones which are distinctly narrowerwhen to 50-100 cm. Trunk monopodial, erect, slender,
still closed. Other differentiatingcharactersare the terete. Bark rough, scaly, on lower part of trunk
resinducts in the leaves, which are fewer and internal breaking into irregularly square plates divided by
in P. herrerae and medial in P teocote, and the less shallow or deep fissures (in part influenced by fire
ciliate cataphylls (margins hyaline) of the former damage, see Hunt, 1962), grey-brown, inner bark

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86 Flora Neotropica

reddish brown. Branches of first order slender, in outline, transverselykeeled, to 15 mm wide, och-
spreadingor ascending;of second and higher orders raceous, light or dark brown, lustrous. Umnbodorsal,
similar, or drooping. Crown broad conical, open or flat or slightly raised or pyramidaland curved up-
irregular; usually a clear bole for 2/3 or -/4 of tree ward, 3-4 mm wide, up to 3 mm high, with a minute,
height. Shoots multinodal,very rough, resinous;with persistentprickle,dull brown,weatheringgrey. Seeds
large, short decurrentand persistent pulvini. Cata- obliquely (narrowly)obovoid, slightly flattened,ob-
phylls large, 15-20 mm long, subulate, strongly re- tuse or angular, 5-7 x 2.5-3.5 mm, light grey-
curved, scarious, with hyaline-ciliate margins, dark brown with dark spots, or dark brown to blackish.
brown, apex caudate, persisting some time after the Seed wings articulateor adnate,effective, held to the
leaf fascicles have fallen. Vegetative buds ovoid- seed by two claws or more commonly enveloping
oblong to cylindrical, 20-25 mm long, the laterals the seed on one side (adnate), obliquely ovate or
smaller, ovoid-acute, (slightly) resinous; scales oblong, 10-20 X 5-8 mm, thin, semi-transparent,
spreadingor recurved,scarious, with ciliate margins, yellowish grey or light brown with dark stripes.
brown. Fascicle sheaths 15-20 mm long, retaining Cotyledons(4-)5-8(-9), (12-)20-35 mm long. Seed-
their length or reducedto ca. 10 mm at maturity;the lings with an elongated stem, primaryleaves green
scales papery, with erose-ciliate margins, yellowish or glaucous, more or less ascending or spreading,
brown,turningdarkred-brownor [Link] soon replacedby secondaryleaves or these delayed.
in fascicles of (2-)3(-4, very rarely 5) in dense, Distribution (Map 5). Caribbean: Bahamas,
irregularlyspaced tufts, persisting3 years, spreading, Turks-CaicosIslands, W Cuba [including Isla de la
straight, (slightly) twisted, rigid, (12-)15-26(-28) Juventud(Isla de Pinos)]. Mesoamerica:Mexico (S
cm X (1.2-)1.4-1.8 mm, margins serrulate, apex QuintanaRoo), N Guatemala,Belize, Honduras(incl.
acute-pungent,light or darkgreen (occasionallyglau- Islas de la Bahia), Nicaragua.
cous-green), moreor less [Link]-
ous on all faces of leaves, in 8-11(-14) lines on the
Key to the varieties of Pinus caribaea
convex abaxial face and 4-6 lines on each adaxial
face. Leaf anatomy:Cross section transverse-triangu- i. Leaves in fascicles of 2-3, mostly 3, fascicle
sheathsreducedin lengthat maturityfrom15-20
lar; hypodermis multi-layered (3-5 rows of cells); mm to 10 mm ................................. 5b. var. hlhamensis
resin ducts 2-5(-8), internal,rarely 1-2 medial;stele I. Leaves in fascicles of 3-4, mostly 3, very rarely2
elliptic in cross section; outer cell walls of endo- or 5, fascicle sheaths persistently long, ca. 15-20
dermis not thickened; vascular bundles 2, distinctly mm.
clustered near 2. Seedlings with green, ascendingprimaryleaves,
separate. Pollen cones the proximal in a few years replaced by secondary leaves....
end of a new shoot, sometimes more spreading by ..........................................................5a. var. caribaea
multinodal habit, subtended by imbricate, scarious, 2. Seedlings with glaucous, spreading primary
reddishbrown bracts,cylindrical,often curved when leaves, development of secondary leaves de-
mature, 20-30 x 5-6 mm, pink or yellow, turning layed ...........................................5c. var. hondurensis
yellowish or [Link] cones subterminal,
often in several whorls (nodes) in one season, mostly 5a. Pinus caribaea Morelet var. caribaea. Type.
in pairs or whorls of 3-5(-8), on 2-2.5 cm long, Cuba. Isla de la Juventud [Isla de Pinos]: Santa
curved, deciduous Barbara,Jun 1957, Luckhoff11608 (neotype, PRF
peduncles (sometimes breaking
off at the base); immaturecones ovoid to narrowly designated by Luckhoff, 1964). Fig. 20A-F
ovoid, 15-20 x 10-15 mm with pyramidal or Pinus recurvataRowlee, Bull. Torrey Bot. Club 30: 107.
1903. Type. Cuba. Isla de la Juventud [Isla de Pinos]:
curved, acuminateumbos terminatingin an erect or
Santa Rosalie, Camp Columbia. 29 Dec 1901, Rowlee
incurvatespine, ochraceous, lustrous, maturingtime 232 (holotype, CU).
two seasons. Maturecones pedunculate,ovoid-coni- Pinus cubensis Grisebach var. anolmala Rowlee, Bull.
cal or ovoid when opened, more or less symmetrical TorreyBot. Club 30: 108. 1903. Type. Cuba. Isla de la
but the base obliquely flattened, (4-)5-12(-13) x Juventud[Isla de Pinos]:JucaroLanding,Dec 1901,
Rowlee 233 (holotype, CU).
(3-)4-6(-7) cm when open. Seed scales ca. 120-
200, partingto release the seeds except at the base, Tree,medium to tall, height to 20-35(-45) m, dbh
thin or thick, oblong, straightor recurved,dark pur- to 60-100 cm. Fascicle sheaths 15-20 mm long,
plish or blackish brown on the abaxial side, dark retainingtheir length at maturity;the scales papery,
brown with faintly visible marks of seed wings on with erose-ciliate margins,yellowish brown, turning
the adaxialside. Apophysisslightly or stronglyraised dark red-brownor [Link] in fascicles of
or nearly flat on basal scales, rhombicto pentagonal 3(-4), very rarely 2 or 5, spreading, straight,

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Systematic Treatment 87

i I
|| e . ,

?X A

...........
^...I..............
...... .... .. . . . . . . . . . .

Map 5. Pinus carib(ea var. caribaea (squares); P. caribaea var. bahatwensis (circles): P caribaea var. hotndurensis
(triangles);P. elliottii var. clensa (stars).

(slightly) twisted, rigid, (13-)15-26 cm X (1.2-) below 400 m. The growing is season continuous in
1.4-1.8 mm, light or dark green, more or less lus- a warm tropical climate with long dry spells. Annual
trous. Stomata on all faces of leaves, in 8-11 lines precipitation varies mainly with altitude, ca. 1000-
on the convex abaxial face and 4-6 lines on each 1800 mm, with a winter dry season. Frost does not
adaxial face. Seed cones (4-)5-10(-12) x (3-)4- occur. Phenology: Pollen dispersal is in January and
6(-7) cm when open. Seed scales ca. 120-170, more February.
or less thin woody. Apophysis slightly raised or
nearly flat on basal scales, rhombicto pentagonalin Representative specimens examined. CUBA. ISLADE
outline, ochraceous or light brown, lustrous. Umbo LA JUVENTUD[ISLA DE PINOS]: Lanier, 20 Nov 1920,
dorsal, flat or slightly raised. Cotyledons (4-)6-7 Ekman12279 (S): Nueva Gerona,22 Jan 1951. Little 14047
(-9), 12-25 mm long. Seedlings with an elongated (US). PINARDELRio: San Diego de los Bahos. 31 Aug
1910, Brittonet al. 6766 (NY); Bahia Honda,9 Mar 1920.
stem, primaryleaves green, more or less ascending,
Ekiman10440 (NY. S); Cortez. 27 Mar 1921, Ekman12802
soon replacedby secondary leaves. (S); Topes de Collantes, Las Villas, 12 Jun 1968, Lambs.n.
Distribution and ecology. W Cuba: Pinardel Rio, (FHO); San Diego de los Bafios, at El Luiar, 14 Apr 1904,
Isla de la Juventud (Isla de Pinos). Forming pure, Le6n 4250 (NY); Guane, 28 Dec 1911. Shrfer 11209 (NY,
U, US); Los Palacios. 10 Jan 1912, Shafer 11697 (B, NY.
open, dry fire-climax forest or open woodland with
US); Pinal de la Catalina, 18 Jan 1912, Shafer 11867 (NY,
undergrowthof grasses or scatteredshrubson sandy U); San Marcos, 27 Jan 1912, Shafer 11968 (K. NY. US).
or gravelly, well-drained, acidic soils. Altitudinal WESTERNCUBA:Loc. unknown, 1860, Wright 1462 (S);
range from 1-700 m, with most extensive stands loc. unknown, 1863, Wright1462B (GOET, K).

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88 Flora Neotropica

I I :i@i J

,I I,

t
tzyI

FIG. 20. A-F. Piols carihaea var. carilbea (A-C. Lmnhb 34; D-F, Lal)mbs..). G-I. P. caribhte var. bahal ensis (G-
H. Brlis 15). J-L. P caribhea var. hiondurensis(Farjon & D. Meji4 356). A. Branch with leaf fascicles and immature
cones. B, G. Leaf fascicles. C, H, J. Cross sections of leaves. D, K. Ovuliferous cones. E. Seed scale (three views). F.
Seeds with adnate wings. I, L. Seeds with articulateand adnate wings. (Magnifications:A, B, D. F, G. 1, K. L. XO.5; C.
H. J. x30: E. x I.)

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SystematicTreatment 89

Remarks on the species. Morelet (1851) de- wings more frequentlyarticulate,also [Link]-
scribed this species as one of two new pines from dons (4-)6-7(-9), 12-25 mm long. Seedlings with
the Isla de la Juventud, Cuba, with rather small an elongated stem, primary leaves green, more or
(ca. 6 cm) cones but otherwise giving an adequate less ascending, soon replacedby secondary leaves.
descriptionof its [Link] Morelet'sCu- Distribution and ecology. Bahamas: on Grand
ban collections were deposited at P, no original Bahama, Great Abaco, New Providence, and North
materialof P. caribaea could be found there. Luck- and South Andros Islands in the north; on Great
hoff (1964) designateda neotype from SantaBarbara InaguaIslandin the [Link] on the Turks-Caicos
on Isla de la Juventud. Islands. On flat, eroded limestone rock with pockets
Probablydue to the obscurityof Morelet'spublica- of sandy soil (pH >7), on sandy spits and old
tion in regional horticultural(1851) and naturalhis- beaches, forming pure, open fire-climaxstands, usu-
tory (1855) periodicals,the name Pittuscaribaea has ally with scatteredor dense undergrowthof shrubs
been erroneouslytaken up for the slash pine of the (often Sabal palnetto); also invadingopen scrubland
SE United States, P. elliottii Engelm. (e.g., de Vail, forming secondaryforest. Altitudinalrange 1-10 m.
1941), a closely related, but nevertheless distinct Annual precipitationdecreases from ca. 1500 mm
species. Subsequently,Loock (1950) compared the on GrandBahamato ca. 1000 mm on Andros in the
Mesoamerican(Belize) populations of P. caribaea northernBahamas, mean annual temperatureis ca.
with "P. caribaea" from the United States, not from 25?C and frost does not occur. Hurricanes are a
Cuba or Isla de la [Link] concluded that the recurringdestructive factor. Phenology: Pollen dis-
pines in Belize "show . . . some distinct botanical persal occurs from Januaryto March.
differences, which are considered importantenough Representative [Link]-
to warrantthe separationof the Belize plants from DROS:San Andros,25 Aug 1988, Baylis 20 (FHO); Duncan
P. caribaea, Morelet, under the name, P. hondur- Point, 30 Aug 1988, Baylis 21 (FHO); Conch Sound. 22
ensis." One can hold the view, as Shaw (1914) did, May 1890, Northrop& Northrop84A (NY); Lisbon Creek,
16 Jan 1910, Small & Carter 8497 (K, NY, US); Nicholl's
that there is just one species incorporatingpopula- Town, 4 Feb 1910, Small & Carter 8973 (K. NY, US).
tions from SE United States to Mesoamerica, but GRAND BAHAMA: McLean's Town, 16 Aug 1988. Baylis
such a broadconcept is not now generally accepted. 10 (FHO); Little HarbourKey, 20 Aug 1988. Baylis 14
The pines from Isla de la Juventud, for lack of a (FHO); Freeport, 19 Aug 1988, Baylis 15 (FHO): Garden
Cay, Neat End, 16 Apr 1905. Brace 3670 (NY). GREAT
holotype specimen, are as near as we can get to a ABACO:Sandy Point, 1 Aug 1988, Baylis 6 (FHO);Cedar
referencepoint for Morelet's Pinus caribaea. Harbour,12 Aug 1988, Baylis 7 (FHO);MarshHarbour,11
Dec 1904, Brace 1634 (NY). NEWPROVIDENCE:Nassau.6
5b. Pinus caribaea Morelet var. bahamensis Sep 1988, Baylis 27 (FHO); Blue Hills, 7 Apr 1904. N. L.
(Grisebach) W. H. Barrett & Golfari, Caribbean Britton25 (NY); loc. unknown, 12 Mar 1907. E. G. Britton
6589 (NY, US); loc. unknown.22 Feb 1888, Eggers 4172
Forest. 23: 69. 1962. Fig. 20G-I (E, M, US).
Grisebach,Fl. Brit.W.I.:503. 1864.
Pinusbahtimensis TURKS-CAICOS ISLANDS. Pine Cay. 25 Aug 1975,
NewProvidence,
[Link]. 20 Nov 1960,Golfari Correll43153 (NY); GrandCaicos,Banberra. Feb 1978,
by Barrett& Golfari,1962, Correll 49463 (NY); Conch Bar. E of Conch Bar airstrip,
77571(neotype,designated
BAB). 14 Jun 1974, Gillis & Proctor 12301 (FHO): Pine Cay. 17
Dec 1907. Wilson7694 (K, NY).
Tree, medium size, height to 20 m, dbh to 50 cm.
Fascicle sheaths 15-20 mm long, reduced to ca. 10 5c. Pinus caribaea Morelet var. hondurensis (Sen-
cm at maturity; the scales imbricate, with erose-
eclauze) W. H. Barrett& Golfari,CaribbeanForest.
ciliate margins, dark red-brownturning grey-black. 23: 65. 1962. Fig. 20J-L
Leaves in fascicles of 3(-2), on GrandBahama2-3,
Pinushondurensis S6enclauze, Conif. 126. [Link].
spreading,straight,(slightly) twisted, rigid, (13-)15- [Link] Cayo,5 Nov 1960,Barrett77582(neotype.
26 cm X (1.2-)1.4-1.8 mm, light or dark green,
designatedby Barrett& Golfari,1962,BAB).
more or less [Link] all faces of leaves, PinushonthrensisLoock,[[Link]]
in 8-11 lines on the convex abaxial face and 4-6 S. Afr. Dept. [Link]. No. 35: 210. 1950, non
lines on each adaxial face. Seed conles (4-)5-10 [Link]:StannCreek,Loock10280
(holotype,PRF).
(-12) x (3-)4-6(-7) cm when open. Seed scales ca.
120-170, moreor less thin woody. Apophvsisslightly Tree,mediumto tall, height to 20-35(-40) m, dbh
raised or nearly flat on basal scales, rhombic to to 60-100 cm. Fascicle sheaths 15-20 mm long,
pentagonal in outline, ochraceous or light brown, retainingtheir length at maturity;the scales papery,
lustrous. Umbo dorsal, flat or slightly raised. Seed with erose-ciliate margins,yellowish brown, turning

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90 Flora Neotropica

dark red-brownor [Link] in fascicles of along rd. to El Volcan, 14 Mar 1972, Styles & Kemp 104
3 (rarely 2, 4, very rarely 5), spreading, straight, (FHO). ZACAPA: Just E of El Lobo, 8 Mar 1978, Stead &
Styles 234 (EAP, FHO, K, M), Stead & Styles 235 (FHO).
(slightly) twisted, rigid, (12-)16-28 cm x (1.2-) BELIZE. BELIZE:Stoney Camp, II Aug 1929, Lundell
1.4-1.8 mm, light green, occasionally glaucous- 677 (F, MO, NY, S, US), Lundell678 (K, NY, US). CAYO:
green, more or less [Link] on all faces of Mountain Pine Ridge, Augustine, 18 Dec 1959, Hunt 62
leaves, in 8-14 lines on the convex abaxial face and (BM, P); MountainPine Ridge, Cooma Cairn, 19 Jul 1966,
Lamb 32 (FHO), 25 Jul 1966, Lamb 34 (FHO); Mountain
4-6 lines on each adaxial face. Seed cones (4-)5-
Pine Ridge, ThousandFoot Falls, 5 Nov 1960, Little 17985
12(-13) X 3.5-7 cm when open. Seed scales ca. (US); Augustine, 31 Nov 1960, Little 17986 (F, NY, US).
120-170, moreor less thin [Link] ORANGEWALK:4 km from San Antonio, 4 Mar 1977,
or strongly raised, but less so on basal scales, rhom- Stead & Styles 116 (FHO, K). STANNCREEK:All Pines,
bic to pentagonalin outline with uppermarginirregu- 29 Jan 1930, Schipp 709 (BM, F, K, MICH, MO, NY, S);
Cabbage Haul Creek, 14 Jan 1988, Stevens 1128 (NY).
larly undulate,chestnut-brown,lustrous. Umbo dor- TOLEDO:Las Lomitas, 18 Jul 1974, Chaplin c60, c61
sal, slightly raised or curved upward,up to 3 X 3- (FHO).
4 mm. Cotyledons 5-8, 20-35 mm long. Seedlings HONDURAS. COMAYAGUA: N of Siguatepeque, 11 Feb
with an elongated stem, primary leaves glaucous, 1977, Stead & Styles 78 (FHO). COPAN:Santa Rosa, 6
Mar 1970, Kemp 38 (FHO). CORTES:Lago de Yojoa,
more or less spreading, development of secondary
Canaveral,on hillside near NE shore of lake, 14 May 1994,
leaves delayed.
Farjon & Mejia 356 (E, FHO, HEH, K, MEXU, U); Lake
Distribution and ecology. Mexico (S Quintana Yojoa, on shore of lake, 26 Nov 1983, Herndndez &
Roo), Belize, N Guatemala,Honduras(includingthe Cornejo s.n. (HEH). EL PARAISO: Yuscaran,Danli-Yusc-
Islas de la Bahia), [Link] distributedon aran jct., 3 Jan 1951, Kemp 57 (FHO); Ojo de Agua, 22
the lowlandcoastal plains withinthe Atlanticclimatic May 1957, Molina R. 8506 (US); Los Limones, 17 Feb
1972, Styles & Kemp 8 (FHO); Yuscaran, 17 Feb 1972,
influence, from the edges of mangrove swamps to Styles & Kemp 14 (FHO, K). FRANCISCO MORAZAN: El
lower upland "bunch-grass/pinesavannas."In Nica- Zamorano, 15 Feb 1972, Styles & Kemp 42 (FHO, K).
ragua it is the southernmostpine in the Americas. It INTIBUCA: 19 Feb 1981, Stead & Styles 725 (FHO); Otoro,
19 Feb 1981, Stead & Styles 726 (FHO). ISLASDE LA
occurs on well-drained, sandy or gravelly, acidic
BAHIA:Good Winter,"IslandW of Guanaja,"26 Jul 1988,
soils, forming pure (or sometimes mixed with P. Cornelius 36 (HEH); Roatan I., Port Royal, 1 Sep 1988,
oocarpa and/or P tecunumanii) fire-climax forest, Cornelius38, 42 (HEH);Guanaja1., 30 May 1982, Hughes
or pine-oak forest, with undergrowthof grasses, 165 (FHO); Guanaja I., n.d., Molina R. 20786 (EAP, F,
Pteridium aquilinum, or Sabal palmetto nearer the US). OLANCHO: Agalta National Park, Catacamas, 1954,
Johannessen470 (IFGP);Aguacate,7 Feb 1965, Johannes-
coast, possibly replacing broad-leaved rain forest sen 969 (UC). SANTABARBARA: Pinalejo, near Quimistan,
underhumaninfluencein many [Link] altitudinal 16 Mar 1977, Stead 22 (FHO).
range is 1-700(-1000?) m. Annual precipitationvar- NICARAGUA. NUEVASEGOVIA: Santa Clara, 3 Mar
ies greatly, from the highest measurement,ca. 4000 1972. Styles & Kemp 78 (EAP, FHO). ZELAYA: Near Tala
Has and Puente Mango (over Rio Kisalaya), 18 Apr 1978,
mm, at Lagunadel Pinarin Nicaraguato the lowest,
Stevens 7638 (ENCB, FHO, MO. NY); between El Em-
ca. 660 mm, at Los Limones in the interior of
palme and Limbaika,24 Feb 1979, Stevens 12787a (FHO.
Honduras; it generally increases from the interior MO, NY). UNKNOWN: "Froma sawmill."no location given,
valleys to the coastal plain and from north to south 10 Oct 1934, Prettymans.n. (NCU).
along the coast. The dry winter season decreases in
durationin a similar [Link] annual tempera- Uses. Pinus cubensis, and especially the Mesoam-
ture is 21-27?C and no frosts occur. Phenology: erican variety hondurensis, is an important timber
Pollen is shed in the early part of the dry [Link] forming easily accessible, extensive stands in
Styles et al. (1982) cited reportsof pollen dispersal
level to moderately hilly terrain. As such it is heavily
in November and Januaryin different areas. exploited, and although not in danger of extinction
in any of its major areas, many local and especially
Representative specimens examined. MEXICO.
outlying populations that are likely to be genetically
QUINTANA ROO:Ejido Caobas, 7 Feb 1981, Avila s.n.
(ENCB, MEXU). distinct are now severely degraded (Dvorak & Do-
GUATEMALA. ALTAVERAPAZ:Secanquim, 3 Jan nahue, 1992). One of these is Ejido Caobas, Quintana
1905, Maxon & Hay 3141 (GH, NY, US). BAJAVERAPAZ: Roo, Mexico, which is the northernmost occurrence
Sierra de las Minas, Mittak 8283 (FHO). CHIQUIMULA: of P caribaea var. hondurensis.
Loc. unknown, Bernoulli & Cario 1090 (B, GOET). EL
Three varieties of Pinus caribaea are recognized,
PETEN:Poptun, 21 Jun 1971, Harmon & Fuentes 5848
(NY); 1.5 km NW of Poptun, Lickhoff 11940 (PRF). as is customary also in forestry literature and prac-
IZABAL: Montana del Mico, Stevermark38548 (F, US); tice. We have not found sufficient evidence (i.e.,

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Systematic Treatment 91

discontinuous,fixed characters)to separatevar. hon- variancewith the presentconcept of the species and
durensis from P caribaea as a distinct species, de- cannot be confused with any other taxon in Belize
spite its greatervariability. or coastal [Link] is no reason, therefore,
Rowlee (1903) was apparentlynot awareof More- to reject the name in favour of Loock's later hom-
let's species, which had been publishedin a compara- onym. No original materialappearsto be extant (the
tively unknownperiodicaland was not mentionedin description may have been based on live material
later nineteenth-centurygeneral treatmentsof coni- with no specimens preserved) and a neotype has
fers. His P recurvatais a taxonomic synonym of P been designatedby BarrettandGolfari(1962). Loock
caribaea var. caribaea, not of P tropicalis as it is (1950), like other writers before him (e.g., de Vail,
sometimes cited. Rowlee also described P cubensis 1941) comparedP. caribaea auct. from Mesoamerica
var. anomala from near "Jucaro Landing" on the with P elliottii (then still commonly known, espe-
eastern side of Isla de la Juventud. Pinus cubensis cially among foresters, as P caribaea Morelet) and
does not occur on this island, and this variety is concluded that he had a case for treating them as
also synonymous with P caribaea var. caribaea. different species. Unaware of both Morelet (1851,
Its "anomalous"leaf-like cataphylls described from 1855) and Seneclauze (1868), he proposedthe obvi-
young shoots appearto be not different from those ous epithet [Link] is no reason to doubt
in the species generally. Liickhoff (1964) considers that Loock referredto the same taxon as Seneclauze
the adnateseedwing of P caribaea from Cuba(Pinar had in mind. Morphologicalcharactersare especially
del Rio), "as opposed to the clearly articulateseed variable in Mesoamerican Pinus caribaea, which
wings from all other areas," the most outstanding has the greatest geographical range and ecological
differencebetween seeds of the variousprovenances. amplitude of the species as a whole (Styles &
His observationsare based on examinationof large Hughes, 1983). These authors,as well as Barrettand
samples and methodsof "seed handlingor cleaning." Golfari (1962) and Luckhoff (1964) report differ-
Adnate seeds occur in the Bahamas as well as in ences in colour and growth pattern of seedlings
Honduras,as has been pointed out by Loock (1950) between P caribaea var. hondurenlsisand the two
for Hondurasand observed in this study. This casts Caribbean varieties. We have used these, besides
serious doubts on the "infallible means of identi- mature foliage characters,to key out the varieties.
fying" P caribaea var. caribaea by this character Additionally,the more stronglydeveloped apophyses
state, as Lickhoff (1964) suggested. of the cone scales in P caribaea var. hondurensis
Grisebach (1864) described Pinus bahaimensisas are usually a reliable characterstate.
a new species from the Bahamas,based on Grisebach Hybridizationbetween P caribaea var. hondur-
s.n. (holotype, K, lost). Grisebach, too, was appar- ensis and P oocarpa has often been inferred from
ently not aware of Morelet's descriptionof P cari- intermediatemorphology(see Styles et al., 1982, for
baea. Barrettand Golfari (1962) designated a neo- review) and is possible in areas where both taxa are
type. They mention several differential character sympatricand their phenologies overlap in time, as
states from P caribaea and conclude that the pines is the case in NW [Link] hybrids with
of the Bahamas should be classified as a variety of P. tecunumanii(as P oocarpa [Link],vide
it. Apartfrom lower leaf numbers,they find a greater Perry, 1991) as well as with P oocarpa are reported
size range in the cones and predominantlyarticulate from MountainPine Ridge in Belize and other areas.
seed wings. From our observations, it appears that
such differences are present,but less pronouncedas
6. Pinus elliottii Engelmann var. densa Little &
suggested by Barrett and Golfari (1962) and on K. W. Dorman,J. Forest. (Washington)50: 921, f.
the whole with considerably overlapping character
states. Both the lower average numberof leaves per 1, 2. 1952. Fig. 21
fascicle and the more often articulateseed wings are
Pinus elliottii Engelmannsubsp. densal(Little & K. W.
charactersthat P caribaea var. bahamensis shares Dorman)E. Murray,Kalmia 12: 23. 1982: Pinus densa
with P elliottii (var. densa) of the SE United States. (Little & K. W. Dorman) Silba, Phytologia Mem. 7:
Seneclauze (1868) describeda new species of pine 50. 1984; Type. United States. Florida:HendryCo.. SE
based on material he had received in 1854 for his of La Belle, 19 Jan 1951, Little & Dorman 14033
(holotype, US; isotypes, F, NY).
nurseryat Bourg-Argental(Dept. Loire, France)with Pinus densa (Little & K. W. Dorman)Silba var. austro-
the name Pinus [Link] description,though kevsensis Silba, Phytologia 68: 49. 1990. Type. United
very concise and based only on foliage, is not at States. Florida:Little Pine Key, I Mar 1911, J. K. Small

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92 Flora Neotropica

2ii

itA A

FIG. 21. Pinus elliotii var. lenst (A, C, D, Hoffinan296; B, Killip 43267; E, G, H, J. K. Small & Nash s.n.; FJ. K.
Small et al. 3635). A. Branch with leaf fascicles and immaturecones. B. "Grass stage" seedling. C. Leaf fascicle. D.
Cross-section of leaf. E-F. Ovuliferous cones. G. Seed scale (two views). H. Seed with wing. (Magnifications:A-C, E-
H. xO.5; D, x30.)

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Systematic Treatment 93

et al. 3635 (holotype, NY; erroneously cited in the Seed cones subterminal,commonly in whorls of 2-
protologueas 3655). 5 or more, sometimes 2(-3) whorls on multinodal
Pinuscaribaeaauct.,nonMorelet(1851),proparte,e.g.,
shoots, on 10-20 mm long, stout, scaly peduncles.
Small,Floridatrees2. 1913;Shaw,[genusPinus]Publ.
ArnoldArbor.5: 70. 1914;De Vail,[Link]. erect at pollination but soon spreadingor recurved,
Sci. 5: 121-132. 1941; Loock, [Pines Mexico Brit. deciduous. Immaturecones ovoid to ellipsoidal, II-
Honduras] S. [Link]. 35: 210-211. 13 x 6 mm, pinkish to pale purple, with greenish
1950.
Pinusheterophylla (S. Elliott)Sudworth,non K. Koch apophyses,maturingtime two seasons. Maturecones
Bull.
(1849),nec Presl(1849),proparte,e.g. Sudworth, ovoid-attenuateto ovoid-conical when closed, ovoid
TorreyBot. Club 20(1): 45. 1893;Sargent,Silva N. to broad cylindrical when opened, symmetrical,
Amer.11: 157-159,t. 591-592. 1897. (5-)7-12(-15) X (4-)6-8.5 cm when open. Seed
scales ca. 120-150, readily parting except near the
Tree,small to mediumor occasionally tall, rapidly base, thin woody, flat, oblong, straight or slightly
growing at early age (ca. 20 years), height to 15- recurved, nearly symmetrical, dark brown on the
25(-30) m, dbh to 50-(X) cm. Trunkmonopodial, abaxial side, with lighter coloured marks of seed
erect, straight or somewhat tortuous. Bark thick, wings on the adaxial side. Apophysis transversely
scaly, breaking into large, irregularplates, divided rhombicand keeled, moreor less flattenedor slightly
by deep fissures,reddishbrown,weatheringgrey; on raised, ochraceous to (reddish) brown, lustrous.
young trees rough, scaly, reddish brown. Branches Umbo dorsal, raised, with a persistent,stout, curved
of first order few, long, spreading or ascending, 1-2 mm long prickle, darker than the apophysis.
tortuousin older trees; of higher orders spreadingor Seeds obliquely ovoid, acute at proximal apex, or
ascending, the ultimatebranchesassurging,forming angular,6-7 X 4 mm, mottledgrey to nearlyblack.
an open, roundedor flat-toppedcrown. Shoots often Seed wings articulatebut effective, held to the seed
multinodal,on lateral shoots uninodal, thick, rigid, by two claws, oblong or obliquely obovate. size
orange-brownto brown, with small, decurrent,cadu- variable with cone(scales): 15-30-(35) x up to 10
cous pulvini. Cataphylls small, subulate, scarious, mm, light brown usually with darkertinge. Seedlings
recurved, brown, with hyaline-erose margins, early with a very thick hypocotyl, cotyledons 5-8; devel-
deciduous. Vegetativebuds ovoid-conical, the termi- oping into a "grassstage"with delayed shoot elonga-
nal buds 15-25 mm long, the laterals smaller, not tion, a short stem near the ground, a thick taproot
resinous;the scales imbricate,linear-acuminate,red- and maturefoliage.
dish brown with white, ciliate margins. Fascicle Distribution and ecology (Map 5). United States:
sheaths initially 10-15 mm long, persistent but In S Florida and north along the coasts to central
slightly reduced to ca. 8 mm in mature foliage; Florida; also on eight of the Lower Florida Keys.
scales reddish brown with white, ciliate margins, Formingpure fire-climaxforest or scatteredin grass-
weathering grey. Leaves in fascicles of 2(-3), in land on sandy lowland plains or on calcareous rock
dense tufts near the ends of branches, spreading SW of Miami and in the FloridaKeys. There is often
obliquely,rigid, straightor slightly twisted, (16-)18- a dense undergrowthof the palm Sabal palmettoand
25(-30) cm x ca. 1.5 mm, margins serrulate,apex it can grow mixed with taller palms locally. Its
acute-acuminate,pungent, dark green, slightly lus- altitudinalrangeis 1-20 m. The climate is subtropical
trous. Stomataon all faces of leaves, in 10-12 lines to tropicalon the Lower FloridaKeys: annualprecip-
on the convex abaxial face and ca. 10 lines on the itation is 100-150 mm, with a six-month dry winter
adaxial face. Leaf anatomy: Cross section semi- period. It is especially well-adapted to frequent
circular, with three leaves in a fascicle transverse- ground fires due to its "grass stage" strategy,which
triangular;hypodermis with (2-)3-4(-5) layers of is found in the seedlings up to sapling age (Little &
cells; resin ducts 3-9(-11), internal, sometimes 1- Dorman, 1954). Phenology: Pollen dispersal takes
2(-4) medial, variablein diam.; stele elliptic in cross place from late FebruarythroughMarch,depending
section, outercell walls of endodermisnot thickened; on weatherconditions (Little & Dorman, 1954).
vascular bundles 2, distinctly separate, sometimes
connate. Pollen cones crowded near the proximal Representative specimens examined. UNITED
end of a new shoot in short clusters, subtendedby STATES. FLORIDA: Near Miami, 21 Mar 1904. Britten
101 (NY); Florida Keys, Big Pine Key. 4 Jul 1956. Briz-
triangular,reddish brown bracts, ovoid-oblong, then
ickv& Stern378 (NY);Lee County,SanibelIs.. E partof
cylindrical,30-50 x 5-8 mm, at first purplish,then island, 29 Dec 1953. Cooley 2485 (NY); Dade County,
reddish brown; microsporophyllspeltate, the distal "edge of Everglades." 30 Dec 1940. Gifford et al. s.n.
part radially striate, with erose-denticulatemargins. (NCU); Florida Keys. Big Pine Key. near Blue Hole at

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94 Flora Neotropica

intersection of Key Deer Blvd. & Miami Ave., 16 Mar as he considered the characteristicsof the seedling
1982, Hill 10974 (NY); MyakkaRiver State Park,Myakka.
28 Mar 1987, Hoffman296 (NY); Florida Keys, Big Pine
"evolutionarilyimportant,"but by failing to cite the
Key, along newly cut rd. direct from Inn to Palm Willow,
basionym he enabled Silba (1984) to publish the
25 Apr 1953, Killip43267 (NY); EvergladesNationalPark, combination validly under the Code. A specimen
between Coconut Grove and Cutler, Oct 1903, Small & with small cones and relatively long (ca. 30 cm)
Carter 1249 (NY); near Miami, Oct 1901, Small & Nash leaves, preservedin NY (for the cone, see Fig. 21F),
s.. (NY);FloridaKeys,Big PineKey,27 Feb 1911,Smaill was described
et al. 3575 (NY). by Silba (1990) as a distinct variety
occurring in the Florida Keys. There is no such
Uses. Pinus elliottii var. densa, the South Florida distinction occurringas fixed charactersin the Flor-
slash pine, is a regionally important timber tree ida Keys, as other collections from there amply
prized for its dense wood. Substantial areas are testify.
logged regularly,but in most cases regenerationis
rapidand successful. Substantialvirgin stands (natu- 7. Pinus cubensis Grisebach, [PI. Wright.] Mem.
rally subject to cyclic fires) are protected in Ever- [Link]. Arts, ser. 2, 8: 530. 1862. Fig. 22
glades National Park;on the other handsuburbanisa-
tion around Miami has been a factor in its decline Pints occidentalis [Link] (Grisebach)Silba,
there. PhytologiaMem. 7: 55. 1984. Type. Cuba. Eastern
Cuba:Localityunknown,1856,Wright598 (lectotype,
For a considerable time botanists and foresters GOET,heredesignated; isolectotypes,GH, K, NY).
have applied the binomial Pinus caribaea to the PinuswrightiiEngelmann, [Link]. [Link]. 4:
slash pine (P elliottii) of the SE United States, and 185. 1880. Type. Cuba. EasternCuba:Localityun-
to the in southern as known.1860,Wright3190 (lectotype,[Link]-
particularly species Florida,
nated;isolectotypes, GH,GOET,K, S).
well as to pines growing on the islands in the Carib- PinusmaestrensisBisse, Ciencias(Havana),ser. 10, 2:
bean and in Mesoamerica. However, Engelmann 2. 1975; Pinus occidentalis Sw. [Link](Bisse)
(1880a) already had renamed, fully described, and Silba, Phytologia 68: 57. 1990. Type. Cuba. Eastern
Cuba:SierraMaestra,nearLa Francia,28 Mar 1969,
(beautifully) illustratedP taeda L. var. heteroplhlla Olivia& Herrera13904(holotype,HAJB-n.v.;isotype,
Elliott as P elliottii, realizing that he could not use
JE).
the epithet heterophylla at species rank. The lecto-
type of P elliottii: United States, South Carolina,20 Tree, medium to large, height to 25-30 m, dbh to
Mar 1873, Mellichamp s.n. (MO no. 3941460) was ca. 1() cm. Trunk monopodial,erect, usually slender,
designatedby Farjon(1993). Sudworth's(1893) rais- in forest stands with clear boles for 2/3or /4 of total
ing of Elliott's varietal epithet to species rank was height. Bark thick, rough, scaly, on the lower partof
both superfluouswhen publishedand illegitimate,as the trunk breaking into irregular, squarish plates
it created a later homonym. divided by deep fissures, or forming vertical ridges
Little and Dorman (1952) restrictedthe concept between fissures, grey-brown, weathering grey, on
of P caribaea to Caribbeanand Mesoamericanpines, young trees and branchesrough and scaly. Branches
recognizing a single variabletaxon rangingfrom the of first orderslender,spreadingirregularly;branches
Bahamas to Honduras, and further recognized P of higher orders similar, forming a small, irregular,
elliotrii with two varieties in the United States. Con- broadlydomed and open crown. Shoots more or less
sequently, they named the southern Florida taxon multinodal,rough with small, short-decurrent,semi-
Pionuselliottii var. densa. The botanical differences persistent pulvini, glaucous to pruinose in the first
with P elliottii var. elliottii include a prolonged, and second years. Cataphylls small, subulate, scari-
well-defined "grass stage" of the seedling which ous, early recurving, with ciliate margins, brown,
results from apical growth inhibition of the stem. deciduous shortly after the leaves or falling simulta-
This is apparentlyan inherited trait (Little & Dor- neously. Vegetativebuds ovoid-oblong, the terminal
man, 1954) as it is retainedin seedling experiments buds 10-15 mm long, the lateralssmaller, acute, not
outside naturalconditions. Initial growth is invested resinous or sometimes slightly resinous; the scales
in a thick hypocotyl; subsequentlya large taprootis imbricate,more or less appressed, scarious, narrow,
formed and the hypocotyl grows to a short "stem" with ciliate margins, brown. Leaffascicles initially
up to 3 cm thick and 4 cm long (Fig. 21B). Other 9-12 mm long, on mature leaves reduced to 7-
differences appear to be more clinal (Price, 1989), 9 mm, persistent, with imbricate, silvery-brown to
such as the fewer leaves per fascicle and the thicker greyish brown scales. Leaves in fascicles of 2(-3),
hypodermisin the leaves of var. densa. in dense, spreadingtufts, persisting 2-3 years, more
Gaussen (1960) raised the variety to specific rank, or less rigid, or flexible, straight or slightly curved,

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Systematic Treatment 95

1.1

w...
~~~~~~~~~~~~"?i ~'
ZE;~~~~"

FIG. 22. Pinus cubensis (A-C. Marie-Victori & Cleiment 21428; D, E. Lamib .sn. (Moa Mine); F-H. Shafer 3216).
A. Branch with leaf fascicles and immatureovuliferous cone. B. Leaf fascicles. C. Cross section of leaf. D. Immature
cones on recurved peduncles. E. Growing cones. F. Mature cone. [Link] scale (three views). H. Seed (two views).
(Magnifications:A, B, D-F, O.5;C, x40; G. H. x 1.)

(6-)10-15(-18) cm X 0.8-1.3 mm, margins serru- the convex abaxial face and 4-5(-7) on the adaxial
late, acute, light green, occasionally glaucous-green. face(s), weakly correlatedwith width of leaves. Leaf
Stomata on all faces of leaves, with 5-10 lines on anatoiny: Cross section semi-circularor transverse-

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96 Flora Neotropica

triangular(3-leaved fascicles), with convex abaxial s.. (FHO, NY); SierraMaestra,N of Pico Bayamesa,May
side; hypodermis multi-layered with 2-3 rows of 1968, Bisse & Duek 9118 (JE); Santiago de Cuba, E of
Gran Piedra, 26 Apr 1969, Bisse & Lippold 15040 (JE);
cells; resin ducts 3-6, internal; stele more or less Santiago de Piedra, Gran Piedra,Jun 1967, Bisse & Rojas
elliptic in cross section, outer cell walls of endo- 3762 (JE); Sierra Maestra,28 Jun 1914, Ekman 1533 (K,
dermis not thickened; vascular bundles 2, distinctly S); Sierra Maestra. near Reuni6n, 30 Mar 1915. Ekman
7026 (NY, S); Sierra de Nipe, 6 Dec 1919, Ekman 10137
separate. Pollen cones clustered near the proximal
end of a new shoot, cylindrical,often recurvedwhen (K, NY, S): Sierra Maestra, along trail to Pico Turquino,
11 Jul 1985, Gentry & Lavin 50962 (MO); Monte Cristi, 5
mature, 15-20 X 5 mm. Microsporophyllsdistally Aug 1968. Limb s.n. (FHO); Moa Mine, 6 Aug 1968,
peltate, radially striate on the abaxial side, with Lambs.,. (FHO); Braza de Cauto, Jul 1927, Le6n & Roca
erose-denticulate margins. Seed cones subterminal, 10326 (P); Rio Yao. Mar 1943, Marie-Victori, 600/0 (US);
vic. of Moa. 27 May 1943, Marie-Victorin & Clement
solitary or in pairs, less commonly in whorls of 3- 21428 (NY, US); MonteVerdenearYateras.29/30 Apr
4, on 1-2 cm long, bracteate, recurved or straight 1907, Matxon 4341 (MO); Sierrade Nipe, Apr 1966, Sanmek
peduncles, deciduous. Immature cones (narrowly) s.n. (FHO);Sierrade Nipe, 19 Dec 1909, Shaifr 3216
ovoid or oval, 10-15 x 6-9 mm, ochraceous,often (US);nearmouthof Rio Yamaniquey, I Mar1910,Shaftr
4254 (NY); CampLa Gloria,S of SierraMoa, 24 Dec
pruinose;umbos rhombic, with a minute spine. Ma-
ture cones (narrowly)ovoid to ovoid-attenuatewhen 1940, Shlafrr8128 (K, NY, P, US); along rd. from Arriba
to Bermejal,31 Jan 1911,Shafer8453 (K, NY,U. US);La
closed, (broadly) ovoid when opened, (3.5-)4-7 x Perla.19Feb1911,Shafer8842(K,NY);localityunknown,
(2-)3.5-5 cm when open. Seed scales ca. 90-110, 1860-1864,Wright1462B(MO);localityunknown,1863,
partingreadilyexcept the basal ones, oblong, straight Wright1462C(GOET);nearMonteVerde,1859, Wright
or slightly recurved,thin woody, darkpurplishbrown 1462D(GOET,K, MO),Wright1462(MO).
to nearly black on the abaxial side, dark brown with Uses. Pints cubensis is the only pine occurringin
faintly distinct marks of seed wings on the adaxial the eastern partof Cuba (see below); no pines occur
side. Apophysismore or less symmetricalaroundthe naturallyin Cuba between Eastern Cuba and Pinar
cone, slightly raised, transverselykeeled, transverse- del Rio in the extreme west of the island, and as
rhombic to pentagonal in outline, with curved or such the pines are of importanceeconomically as a
irregularupper margin, up to 11 mm wide, radially timber source to this partof Cuba.
striate,darkochraceous to dull brown. Umibodorsal, Grisebach (1862), in his enumeration of plants
depressed or slightly raised, transverse-rhombic,3- collected by CharlesWrightin Cuba,describedPinus
4 mm wide, brown, weathering grey; the minute cubensis as a new species, based on Wright 598
prickle early deciduous. Seeds obliquely ovoid, flat- and 1462. Number 1462 being an ambiguous
tened, 5-6 x 3-3.5 mm, mottled [Link] collection in part mixed with P caribaea (GOET,
wings usually adnate (with membranecovering the seen by B. T. Styles), no. 598 (GOET) is here
seed on one side), or articulate (wing held to the designatedas the lectotype; it has been annotatedby
seed by two claws), obliquely ovate-oblong, 12-16 Grisebach.
x 5-6 mm, grey-brown, with a graphite-liketinge. Engelmann (1880a) described as a new species
Numberof cotyledons not observed. No. 45 Pinls wrightii, citing Wright 1462B and
Distribution and ecology (Map 2). EasternCuba: Wright 3190. The specimen subsequently marked
From the Sierra Maestra and the Sierra de Nipe E 1462C (GOET) belongs here; it is also a syntype
into the highlands terminatingthe eastern partof the of P cubensis. The lectotype of P wrightii is desig-
island. It occurs in foothills and highlandsas well as natedhere to be no. 3190 (MO). Engelmannappears
in "pinebarrens"along the coast. Its altitudinalrange to have confused his species no. 44 P cubensis
is 100-900(-1200) m. Pinus chbensis forms mostly with P caribaea Morelet ("Isle of Pines, Bahama
pure but open stands or is invasive in disturbedsites Islands"); and his P wrightii, which describes P
on serpentine or serpentine-derived,often ferrugi- cubensis from "Mountains in Eastern Cuba" is a
nous soils ("Nipe latosol" or "Nipe clay") or on superfluousname.
alluvial sediment near the coast. In the highlands Sudworth (1893), erroneously equating P taeda
there is abundantrainfall (1800 mm or more annu- var. heterophvlla Elliott, described from Georgia,
ally), but a dry winter season is a typical aspect of United States, with P cubensis, proposedon nomen-
the tropical to subtropicalclimate. Phenology:Time claturalgrounds (American rules) the "older"bino-
of pollen dispersal not recorded. mial P heterophylla(Elliott) Sudworthfor the "Cu-
Representative specimens examined. CUBA. EASTERN ban pine." Under the present Code, no name has
CUBA:Sierra de Nipe, Mayari. 13 Apr 1969, Betancol,rt priorityoutside its own rank. Shaw (1914) synony-

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Systematic Treatment97

mized P cubensis and P wrightiiwith P occidentalis turning grey-brown to grey. Branches of first and
Sw. by applicationof a much wider species concept higher orders slender, spreading or ascending, in
than one would accept today. the lower crown curved and drooping, forming an
Bisse (1975) provideda key for the pines of Cuba irregular,open crown. Shoots uninodal,glaucous to
in which the first couplet divides them into 2- and pruinosein the firstyear, later very rough with large,
3-leaved pines. Fromthe SierraMaestrahe described short-decurrent, persistentpulvini,[Link]
a new species: P maestrensis,with leaves in fascicles slender, subulate to caudate, straight or recurved,
of 3. The leaves are -I1 mm wide, glaucous-green, scarious,with erose margins,brown,persistingsome
and very flexible. The reportedthin leaves (0.5-0.7 time after the leaf fascicles have fallen. Vegetative
mm), predominantlyin fascicles of 3, are the only buds ovoid to ovoid-oblong, acute, the terminalbud
different feature of this taxon, which was said to 10-15 mm long, the lateralsshorter,usually slightly
occur in various places between 300-1700 m to- resinous; the scales imbricate, more or less ap-
gether with P cubensis. The measurementsof Bisse pressed, scarious, with ciliate margins, lustrous
(1975) have not been observed on the isotype (JE), brown. Fascicle sheaths persistent, (8-)10-15 mm
which has leaves 0.8-0.9 mm wide, nor on any other long, scales lustroussilvery-brownwith erose-ciliate
materialat JE determinedby Bisse as P mnaestreesis margins,[Link] in fascicles
(with up to 1.1 mm wide leaves). One of his speci- of 3-5, in dense, spreading tufts at the ends of
mens (Bisse & Duek 9118) has all leaves in fascicles branches, persisting to the third year, straight or
of 2, and another(Bisse & Rojas 3762) primarilyso. slightly curved, more or less rigid, (I -)14-18(-20)
Silba (1984) reducedP cubensis to a varietyof P cm x 1.2-1.4 mm, with serrulate margins, acute.
occidentalis, and P maestrensislikewise to a variety pungent,light green. Stomata on all faces of leaves,
of P occidentalis, repeating the original diagnosis in 6-8 lines on the convex abaxial face and ca. 4
but without having seen any of the original material lines on each adaxial face. Leaf anCatomyv:Cross
or giving an explanation. As Darrow and Zanoni section triangularor transverse-triangular, with con-
(1991) have discussed, the pines of the SierraMaes- vex abaxial side; hypodermis multi-layered (3-4
tra in E Cuba are not P occidentalis, for which they rows of cells); resin ducts (2?-)3-5, internal;stele
have long been mistaken (e.g., map 46 in Critch- elliptic to transverse-obovatein cross section; outer
field & Little, 1966; Mirov, 1967)-that species is cell walls of endodermis not thickened; vascular
native to [Link] do they belong with P bundles 2, distinctly [Link] cones clustered
caribaea. Their placement with P cubensis is in near the proximal end of a new shoot, cylindrical.
agreement with Florin's determination of Ekmian often recurvedwhen mature,15-25 X 5 mm, pinkish
1533, with morphologicalcharacterstates found in yellow, turningyellowish brown. Seed cones subter-
that species east of the Sierra Maestra, and finally minal, solitaryor in pairson 10-20 mm long, straight
with overall geographic patternsof Caribbeanpine or recurved, bracteatepeduncles, persisting several
taxa. years after seed dispersal, falling with the peduncle
attached. Immaturecones ovoid to ovoid-attenuate
8. Pinus occidentalis Swartz, Prodr. 103. 1788. when growing, 10-15 x 8-10 mm, with inflexed
Type. Haiti. "Hispaniola: aux Pins, quartier des spines, ochraceous,maturingin two seasons. Mature
Nippes, du c6ot septentrional,"1785, Swartz s.n. cones ovoid to ovoid-conical, straight or curved.
(lectotype, BM, designatedby Farjon, 1993). nearlysymmetrical,largecones with a flattenedbase,
Fig. 23 (4-)5-9(-11) x 3.5-6.5 cm when open. Seed scales
Pinus occidentalis Swartz [Link] Silba, Phyto- ca. 110-150, partingto release the seeds except those
logia58: 368. [Link] [Link]- at the base, oblong, straightor recurved,thin woody,
nales:Sierrade Baoruco,nearAceitillar,14 May1976. dark purplishbrown abaxially,dark brown with in-
Judd 1487 (holotype. A). distinct lighter brown marks of seed wings on the
Tree, medium to tall, height to 30-40 m, dbh to adaxial side. Apoplhsis slightly raised, transversely
100-120 cm. Trunkmonopodial,erect, terete,usually keeled, rhombic to pentagonal in outline, up to 12
slender, in forest stands with clear bole for 2/3--4 of mm wide, on the proximal scales more or less gib-
height. Barkthick, on lower trunk30-40 mm, rough, bous, (lustrous)darkbrown,radiallystriate,weather-
scaly, breakinginto irregularlysquare plates divided ing dull grey. Umbo dorsal, raised and often curved,
by deep fissures, longitudinally fissured toward the usually armedwith an inflexed, 2-3 mm long spine,
base, on young trees and branches light brown, [Link] obliquely obovoid, flattened,5-6

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98 Flora Neotropica

! ~E a

1, a a

FIG. 23. Pinuusoccidentalis (A-Ba, Hughes 481; Bb, C, Ea, Ga, Zanoni et al. 40761; D, Eb, Gb, Zanoni et al. 41212).
A. Branch with leaf fascicles. B. Leaf fascicles. C. Cross-section of leaf. D. Growing immatureovuliferous cone. E.
Ovuliferous cones. F. Seed scales. G. Seeds. (Magnifications:A, B, D-G, x0.5; C, x 30.)

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Systematic Treatment 99

X 3-4 mm, light, mottled grey-brown. Seed wings 32443 (NY). LA VEGA:Constanza, 23 Apr 1976, Juddl
articulate but effective, held to the seed by two 1166 (NY); Constanza, I Jul 1964, Lamb 10 (FHO); Rio
Jimenoa, 7 km from Jarabacoa,22 May 1986, Zanoni et
claws, obliquely ovate or oblong, 12-18 X 4-6 al. 36453 (NY): 2.5 km SE of Aguacate. 26 May 1988,
mm, ochraceous with black or grey tinge or stripes. Zanom i et cl. 41212 (FHO). PEDERNALES:Sierrade Baor-
Numberof cotyledons not observed. uco, 26 Jul 1981, Fisher-Meerow687 (NY); island W of
Distribution and ecology (Map 2). Hispaniola: Hoyo de Pelempito,NE of Cabo Rojo, I I Feb 1981, Zanoni
Both in Haiti and the Dominican Republic, where et al. 10889 (F, NY): Sierrade Baoruco,33 km N of Cabo
Rojo, 10 Apr 1988, Zatnoti et al. 40760, 40761 (FHO).
the largerpart of its range is; reportsof the species PERAVIA: 42 km NW of San Jose de Ocoa, 30 May 1984,
from EasternCuba (Oriente)are probablyall errone- Mejia et al. 608 (NY), 30 May 1984. Mejia et al. 612
ous and likely pertain to P cubensis. It occurs in (MO, NY, US). SANCRISTOBAL: Los Cacaos, 3 May 1984,
diverse habitatsfrom the lowlands, at about 200 m, Zcaoni et al. 29734 (FHO). UNKNOWN: Rio Bao, Dec
to the highest mountainridges (Pico Duarteand Pico 1952,Humtbert27625(P,S); Rio Bao,locationpresumably
fartherupstreamthan Humbert27625 (P), Dec 1952.
La Pelona) on the island, at almost 3200 m. The Humbert27699 (P); loc. not recorded, 1853. Schomlburgk
more extensive and pure stands occur at 900-2700 43 (P).
m (Mirov, 1967; Farjon, 1984), but in more accessi-
ble areas these are much depleted. Soils are either Uses. Being the only species of Pinis on Hispan-
derived from limestone at lower altitudes, or more iola and formerlyabundantover much of the island,
acid, clay-like, and shallow in the CordilleraCentral. it has been heavily exploited for [Link]
Pinus occidentalis consequentlyis found in a variety to Darrow and Zanoni (1991) it has been depleted
of vegetation types, mostly occupying the shallow, from an estimated3 million ha of primeval,more or
nutrient-poorsoils and rock outcrops, where it may less pure pine forests to perhapsless than 5% of that
occur in open or dense, pure stands or mixed with area, but accurate estimates of even the present
various broad-leavedtrees and shrubs. In disturbed forestextent are lacking. Protectionin the Dominican
(grazed)areasPteridiumaquilinumcan dominatethe Republic is inadequatebut existent, contraryto the
groundcover; in frequentlyburntareas,grasses (e.g., virtually uncontrolledsituation in Haiti (Darrow &
Danthoniadomingensis,Andropogonspp.) and Pteri- Zanoni, 1991).
dium replace shrubs and small trees. Thick bark on This species has been describedby Swartz (1788)
the lower trunks of pines are a protection against from "Hispaniola,"where he explored the western
quickly passing fires, and regenerationcan be rapid part of the island (now Haiti) from December 1784
from seed. Annual precipitationvaries greatly with throughJune 1785 (Steam, 1980). He gives a concise
exposition, but is in the range of 1200-1600 mm description of a 5-leaved pine and a reference to
where most pine forests occur and exceeds 2300 mm "Plum.t. 161,"which is plate 161 in Burman(1758).
in the N and E of the [Link] is a Darrowand Zanoni (1991) refer to a collection in S,
3-5-month dry season during winter, which may but this specimen,althoughfrom Swartz'sherbarium,
bring frost, but rarely snow, at the higher altitudes is a 4-leaved branchletof unknownorigin. A speci-
above 1600-1800 m. Yet there are also pines in men in BM, which is a 5-leaved branchletin accor-
tropical and subtropical regions at low altitudes dance with the protologue, bears on the reverse
where no frostoccurs. A detailed accountof distribu- of the sheet the inscription (in L'Heritier'shand):
tion and ecology is given by Darrow and Zanoni "Hispaniola:aux Pins, quartierde Nippes, au c6ot
(1991) from which much of the above information [Link] Swartz."This specimen has been
is [Link]:Time of pollen dispersalis in designatedas the lectotype (Farjon, 1993).
early spring,probablyvaryinggreatlywith altitude;a Shaw (1914) considered the pines of E Cuba
second period has been reported in October (see conspecific with those indigenousto Hispaniolaand
Darrow& Zanoni, 1991). thereforecited P cubensis Griseb. in synonymy with
P. occidentalis Swartz. His observationof an "erect
[Link]- conelet and a reflexedcone"
ite: Marmalade,1903,Nash825 (NY). NORD:Briquis,26 may have been accurate
Nov 1985, Zanoni & Mejia 35801 (NY). OUEST:Furcy,21 in the specimens he studied,but conelets on recurved
Aug 1917, Cook, Scofield & Doyle 25 (FHO, US). PORT- peduncles occur as well, and this "character"is
AU-PRINCE:Morne la Visite, 35 km S of Port-au-Prince,15 thereforeof little value [Link] collec-
Jan 1985, Hughes 480, 481 (FHO). UNKNOWN: "Sourcail," tions from E Cuba had been determined
20 May 1828, Jaeger 155 (S, US). by B. T.
DOMINICAN REPUBLIC. AZUA:Sabana de Miguel Styles as P. occidentalis because of Shaw's treatment,
Martin,3 May 1988, Zanoni et al. 40832, 40833 (FHO). which seems to have been based on a mixture of
DAJABON:Sierra Chacuey, 27 Nov 1984, Zanoni et al. taxa (Darrow & Zanoni, 1991). No authentic P

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100 Flora Neotropica

occidlentalis collected in Cuba from the wild has cous. Cataphylls 10-15 mm long, subulate, scarious,
come to the attentionof the first author(A. Farjon), recurved,with hyaline-erose to ciliate margins, red-
who agrees to restrict P occidentallisto Hispaniola dish brown, weathering blackish, mostly abscising
in the absence of positive evidence to the contrary. with the leaf fascicles. Vegetativebuds ovoid-acute
Silba (1985b) described as new to science P occ-i- to broad-ovoid, the terminal bud 15-25 x 10-15
deitalis var. baorucoensis, with leaves in fascicles mm, the laterals smaller, not resinous; the scales
of 5 (rarely4), and with "swollen, sometimes protu- spreadingor recurvedat base of bud, subulate-cau-
berant apophyses with a fine, upcurved, prominent date, lustrousred-brownwith darkerapex and white
spine" from the Sierrade Baoruco in the Dominican or brown ciliate margins. Fascicle sheaths initially
Republic and other localities both in the Dominican ca. 20 mm long, consisting of ca. 10 imbricate,
Republic and Haiti. Similar cone characters have appressed,orange- to red-brownscales with hyaline
been observed in specimens with 3-leaved fascicles margins, in mature fascicles persistent but reduced
(e.g., Zanoni et al. 40760, 40761 from the Sierrade to 10-15 mm, with a thickened base, weathering
Baoruco, FHO), whereas 5-leaved specimens may blackish grey. Leaves in fascicles of 2-3 (in some
have cones with more-flattenedapophyses and less trees predominantly2 or 3), in dense, spreadingtufts
prominentspines. The holotype specimen (A), deter- toward the ends of upturnedbranches,persisting 2-
mined by Zanoni Dec 1981 as P occidentalis, has 2.5 years, straightor curved, rigid, (10-)15-25(-27)
3-, 4-, and 5-leaved fascicles. As with virtually all cm X 1.3-1.6 mm, with serrulate margins, acute-
of Silba's "new"taxa, this one seems to be based on pungent to acuminate, light or dark green. Stomata
odd observationsof only a few specimens (a scan of on all faces of the leaves, in 8-10 lines on the convex
the relevant materialin the herbariawhere the types abaxial face, in 3-4 lines on each adaxial face.
of his taxa are deposited usually reveals this), and it Leaf anatomy:Cross section transverse-triangular (3-
was proposed without consultation of the botanist leaved fascicles), or semi-circular (2-leaved fasci-
(Zanoni, in this case) who knew intimatelyboth the cles), with a convex abaxial side; hypodermis uni-
area and its pines. form, with 2-3 layers of cells; resin ducts 2-6,
medial;stele oval in cross section; outercell walls of
9. Pinus ponderosa D. Douglas ex C. Lawson var. endodermisthickened;vascularbundles 2, distinctly
scopulorum Engelmannin S. Watson,Bot. Califor- separated. Pollen cones densely clustered near the
nia 2: 126. 1880. Fig. 24 proximalend of a new shoot, subtendedby subulate
Piins ponderosa D. Douglas ex C. Lawson subsp. scopu- bracts with white ciliate margins, ovoid-oblong to
E. Murray,Kalmia12: 23. 1982;
loruni(Engelmann) cylindrical, 15-30 X 5-7 mm, yellowish, turning
Pinus scopultorum (Engelmann) J. G. Lemmon, Gard. yellowish [Link] cones subterminal,solitary or
Forest10: 183. [Link]: in whorls of 2-3 on short, stout, recurvedpeduncles,
"ColoradoTerritory,
lat. 39-41?. Alpineand Subal-
pine,"Jun 1862,Parrys.n. (lectotype,MO,heredesig- persistent some time after seed dispersal, leaving a
nated). few basal scales on the pedunclewhen falling. Imma-
PilnusbrachYptera
Engelmann in Wislizenus,[Link] ture cones broadly ovoid to subglobose, with sharp,
N. Mexico89. [Link] Mexico: spreading or curved spines, (15-)20-25 x 12-18
"RockCreekto SantaFe,"5 Jul 1846,WislizeYnus
534 mm, purplish brown, glaucous, maturing in two
(holotype,MO).
years. Mature cones ovoid to subglobose when
Tree, usually tall, height (Mexico) to ca. 40 m, opened, symmetrical or slightly oblique toward the
dbh to 80-120 cm. Trunkmonopodial,straight,erect, base, 5-10 X 4.5-7 cm when open. Seed scales ca.
usually [Link] thick, roughand scaly, dividing 90-120, partingwhen mature,thin, flexible, oblong,
into large, slightly elongate plates separated by with thin margins,straightor recurved,darkpurplish
broad, shallow fissures, clear orange-pinkishor red- brown with a light brown edge below the apophysis
dish brown with dark fissures on the lower part of abaxially,light brownwith faint marksof seed wings
the bole, on young trees and branches thin, scaly, adaxially. Apophysis slightly to prominentlyraised,
brown or darkreddishbrown. Branchesof firstorder transverselykeeled, rhombic or irregularlypentago-
relatively short,spreadingor curveddown, numerous nal in outline, ochraceous to reddish brown. Unbo
and persistent;branches of higher orders ascending dorsal, flat to prominently raised and transversely
or spreading, the lower branches subpendulous,as- keeled, 5-7 mm wide with a persistent, curved, to
surgent at tips. Shoots stout, rough, with persistent, 3 mm long spine. Seeds obliquely ovoid, slightly
decurrentpulvini which leave prominentmarksafter flattened,5-7 x 4-5 mm, light grey-brown, some-
the fascicles have fallen, reddish brown, often glau- times with dark spots. Seed wings articulate,effec-

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Systematic Treatment 101

. I
III-o ~

FIG. 24. Pinus ponderosa var. scopulorun (A, Ba, C, Wynd& Mueller 650; Bb, D, E, Parry s.n. (lectotype); F-H,
MacKelveY194, FHO, from Arizona, not cited). A. Shoot with leaf fascicles. B. Leaf fascicles. C. Cross section of leaf.
D, E. Shoots with leaf fascicles and growing ovuliferous cones. F. Matureovuliferous cone. G. Seed scale (three views).
H. Seeds, with articulatewings. (Magnifications:A, B, D-H, X0.5; C, x30.)

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102 Flora Neotropica

"-r~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~i....
...

A --- I I
i~.._~l,,,,; ',,. : / ,,, u ,,-~ : ~ _. ,, , ....???...
-':
I,

i~ . I~
~~.---?--I ..

Map 6. Pinus [Link] var. scopulorumn(squares); P. arizonica var. cooperi (triangles); P arizonica var. .torniae
(circles).

tive, held to the seed by two narrow,flimsy claws, 1500 m) to receive a minimum of ca. 200 mm
obliquely oblong, 15-20 x 6-10 mm, translucent, precipitationduringthe growing season.
light yellowish brown. Cotyledons6-9(-1 I). The vegetation on the two mountains in Mexico
Distribution and ecology (Map 6). The range of where P. ponderosa has been found is similar to that
Pinus ponderosa extends from SW Canada to the N of the border,with open, park-likestands of Pinus
Mexican border. Commonly, the species is recog- interspersedwith Quercus, Arbutus, shrubs such as
nized to occur in two wide-ranging varieties, var. Arctostaphylos, and grassland. Phenology: Pollen
ponderosa and var. scopulorum(with P. arizonica as dispersaloccurs in May-June,a dry period.
a third variety in the United States, which is here
recognizedas a species). Pinus [Link]- [Link]-
lorumis the interioror "Rocky Mountain"form, and huahua:W sideof theSanLuisRange,in the firstcanyon
S of DiabloCanyon,3 Sep 1952, Tucker2596 (A, UC).
it is this taxon that is found in two separatelocalities COAHUILA: Sierradel Carmen,highpeakadjoiningCanon
just S of the Mexican border, in Chihuahua and Sentenela,10 Jul 1936, Wynd& Mueller650 (MO, NY,
Coahuila. In the region of the SW United States P).
adjacentto the Mexican border,P. ponderosa occurs
in the "yellow pine belt" above a zone characterised Uses. Ponderosapine is one of the most important
by Pinus edulis, Juniperus monosperma,and Quer- timbertrees of the United States. Due to the remote-
cus, on mostly isolated mountainsseparatedby broad ness of the areas where it is found in Mexico, no
intermontanevalleys of sufficient elevation (above commercial exploitation is likely to occur; its use

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SystematicTreatment 103

would otherwise be comparablewith that of P ari- Pinus ponderosa [Link]. is recognized here as oc-
zonica. curringin two wide-rangingvarieties:[Link]
Pinus ponderosa var. scopulorum was distin- and var. scopulorum (Kral, 1993). Trees with mor-
guished from P. ponderosa of Oregon and California phologicalcharacterstates as given in the description
by Engelmann (1880b), who also referred to it as above occur in the San Luis Range in extreme NE
"Pinus ponderosa of the Rocky Mountain flora's." Chihuahua,close to the United States-Mexico bor-
The materialupon which Engelmannbased this vari- der, and in the Sierra del Carmen, just south of
ety included a collection by C. C. Parry,made in the Rio Grande in Coahuila. They belong to var.
June 1862 and bearing a label imprinted "Rocky scopulorum. Bailey and Hawksworth(1983) antici-
MountainFlora,"which is in Engelmann'sherbarium pated such occurrencesnear the border.
(MO), with two small brancheswith leaves up to 10 Martfnez(1948), when referringto P. ponderosa
cm long in fascicles of 2-3 and immatureto half- in Baja California, confused it with P jeffreyi, a
grown seed cones. Since Engelmanndescribed also species he also recognized as distinct. The southern
maturecones and seeds, he must have had additional California"race"of P ponderosa (Conkle & Critch-
material,but it has not been found in MO. The Parry field, 1988) has its southernmostextension in the
sheet is neverthelessdesignated as the lectotype be- Laguna Mountainsin San Diego County (Griffin &
cause Engelmann used to send out other bits to Critchfield,1972;A. Farjon,pers. obs., 1992), where
Parlatore and other botanists, which may yet be it is sympatricwith P jeffrevi.
found. Engelmann (1880a) distinguished P pon-
derosa [Link] both [Link] (= P 10. Pinus arizonica Engelmann,in Rothrock,Rep.
jeffreyi) and [Link] by its smaller cones and U.S. Geogr. Surv., Wheeler,6, Bot.: 260. 1879.
shorterleaves, "often in pairs."This lattercondition
is, according to Conkle and Critchfield(1988), not Tree, usually tall, height to 30-35 m, dbh to 100-
found in the "Southwesternregion,"i.e., Arizonaand 120 cm. Trunkmonopodial,erect, straight,tereteand
New Mexico, where only 5-15% of leaf fascicles slender, often clear of branches 2/- to 3/5 its length.
are foundto be 2-leaved and where 3-leaved fascicles Bark thick, rough, scaly, breaking into large plates
[Link] extension of usage of the epithet divided by broad, shallow or deep fissures, reddish
scopulorumto the Ponderosapines of this region fits brown to dark brown, outer bark grey; on young
the clinal patternof this trait from north to south in trees and branches thin, rough and scaly, reddish
the interiorpopulations(Conkle & Critchfield,1988: [Link] firstorderheavy, long, spreading
fig. I) and is accepted here. horizontallyor curved down; of second and higher
Shaw (1909, 1914) used a broad circumscription orders ascending or spreading, the lower branches
of Pinus ponderosa, which included P arizonica, P subpendulous,[Link] young trees pyrami-
engelmannii, and P jeffreyi as varieties in Mexico dal, open, in old trees roundedor flat-topped,open
(1909), and later also P scopulorum (1914). In the or dense. Shoots at first orange brown or glaucous,
United States, P arizonica is still widely considered rough with persistent,decurrentpulvini. Cataphylls
a variety of P ponderosa (Little, 1979; Burns & up to 15 X 3-4 mm, subulate, scarious, soon re-
Honkala, 1990; Kral, 1993), but that view seems flexed, with erose-ciliate marginsand caudate apex,
to be based primarily on the populations in the dark brown. Vegetativebuds ovoid to ovoid-acute,
southwesternUnited States and not on an evaluation the terminalbuds 15-30 X 10-15 mm, the laterals
of the complex throughout northern Mexico. The smaller,not resinous, brown;the scales spreadingor
only authorof these who has a broad knowledge of recurvedat apex, with long-ciliate, white or brown
Mexican pines is Little, but he favoureda conserva- [Link] sheaths initially long, 15-30 mm,
tive approachto species recognitionin his checklists, with ca. 10 imbricate, orange-brownto red-brown
which is of U.S. trees only (Little, 1953, 1979). scales with ciliate margins,persistentbut reducedto
There may be naturalhybridizationin Arizona and ca. 10 mm, with a thickenedbase, weatheringgrey-
New Mexico where the two taxa occur together black. Leaves in fascicles of 3-5, spreadingat a wide
(Conkle & Critchfield, 1988), complicating recogni- angle from the shoot, persisting 2-3 years, rigid to
tion of these taxa there. Monoterpenecomponents slightly lax, straightor slightly curved and twisted,
seem to be as variable between P arizonica and P (8-)10-20(-25) cm X 0.9-1.8 mm, margins serru-
ponderosa as within the latter species (Conkle & late, apex acute to pungent,light yellowish green or
Critchfield,1988) and provide inconclusive evidence glaucous-green. Stomata on all faces of leaves, in
for species delimitation. (3-)4-8(-12) lines on the convex abaxial face and

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104 Flora Neotropica

'
~~~~~~~~~~~'
'
~" i' .,- I.. '
?~~~~9
'.~i~ .
.~ ~?~~ . .z
.
???."~'-N~
" ?

i ~.
'-??

7. Ma Pin arznc . va.

arizonica?-? (
gr c(Iranges
rle)
P~..?U
e 1 ggii

in (3-)4(-5) lines on each adaxial face (the abaxial slightly curved, (4.5-)5-10(-14) X 3.5-6(-8) cm
numberof lines correlatedwith width of leaf). Leaf when open. Seed scales ca. 90-140, most numerous
anatomy: Cross section (transverse-)triangular;hy- at the base of the cone, partingto release the seeds
podermis multi-layered, with intrusions into the except at the base, thick, rigid, broadlyoblong, nearly
mesophyll; resin ducts (3-)5-10, medial, usually 3 straight or recurved especially near the base, dark
larger(external) ducts present;stele elliptic in cross purplishto blackish brown on the abaxial side, light
section; outer cell walls of endodermis thickened; brown with faint marksof seed wings on the adaxial
vascularbundles 2, more or less connateor distinctly side. Apophysisraised in the central part, rhombicto
separate. Pollen cones densely clustered near the pentagonal in outline, mostly symmetrical around
proximalend of a new shoot, subtendedby subulate the cone, transversely keeled, ochraceous to light
bractswith ciliate margins,ovoid-oblong to cylindri- reddish brown. Umbo dorsal, raised and slightly
cal, 15-20 X 5 mm, yellowish to yellowish brown. recurvedor nearly flat, rhombic,transverselykeeled,
Seed cones subterminal,solitary, in pairs or whorls 4-8 mm wide, grey, with a minute,deciduous prickle.
of 3-5, on short, curved, persistent peduncles re- Seeds obliquely ovoid, slightly flattened,4-7 x 3-
taininga few scales on the [Link] 5 mm, light brown with black spots. Seed wings
ovoid-globose, ca. 15 X 10-12 mm, with sharp, articulate but effective, held to the seed by two
curvedor spreadingspines, purpleto purplishbrown, flimsy claws, obliquely ovate, 12-20 x 4-8 mm,
maturingtime two seasons. Maturecones seemingly semi-transparent,greyish brown with black tinge.
sessile, ovoid or broadly ovoid with parted scales, Numberof cotyledons 10-12.
asymmetrical at obliquely flattened base, often Distribution (Maps 6, 7). SW United States: Ari-

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Systematic Treatment 105

zona, New Mexico. Mexico: Mainly in the Sierra Arbor.1: 24. 1909;PinusponderosaD. Douglasex P.
Madre Occidental, south to S Durango, scatteredin [Link](Engelmann) E. Murray,Kal-
mia 12:23. [Link]:Santa
Coahuila, NE Zacatecas, and Nuevo Le6n. RitaMts., 1874,J. T Rothrock
652 ([Link]).
Remarks on the species. Pinus arizonica was
based on a collection made by J. T. Rothrockwhile Tree, usually tall, height to 30-35 m. Leaves in
on one of the U.S. Army Survey expeditions in the fascicles of 3-5 (predominantly3-4), rigid to slightly
Santa Rita Mountains of southern Arizona. Engel- lax, straightor slightly curved and twisted, (8-)10-
mann (1879) stressed the difference in leaf number 20(-23) cm x 0.9-1.4(-1.6) mm. Stomata on all
with P ponderosa (5 vs. 3), but found it otherwise faces of leaves, in (3-)4-8 lines on the convex
very similar to certain forms of that species. Shaw abaxial face and in (3-)4(-5) lines on each adaxial
(1909, 1914) reduced it to a variety of P. ponderosa face (the abaxial number of lines correlated with
when it became known that the "leaf-fascicles (are) width of leaf). Seed cones ovoid to broadly ovoid,
heteromerous,with the largernumberin the southern often slightly curved, (4.5-)5-7 x 3.5-6 cm when
partsof its range"(referringto P ponderosa s.l.). As open. Seeds obliquely ovoid, slightly flattened,4-6
discussed under P ponderosa var. scopulorum, P. X 3-3.5 mm. Seed wings obliquely ovate, 12-15 x
arizonica has often been treated under that species 4-6 mm.
as a variety,but more recent treatmentsrecognize it Remarks. There seems to be clinal variation in
as a distinct species, with possible introgressionin the numberof leaves per fascicle, with trees having
Arizona and New Mexico. Table IV compares and predominantly3(-4) leaves more frequentin Sonora,
contrasts the morphological characters of the two Chihuahua,and Coahuila;those with predominantly
taxa. 5 leaves are more frequent in Durango. Exceptions
The cones, otherwise very similar, are generally do occur: 3-leaved in Sinaloa: H. S. Gentry 7242
somewhat smaller in P arizonica, but here again (MICH);(4-)5-leaved in Sonora:J. T. Marshall 134
considerable overlapping is found and with no ob- (UC), S. S. White3360 (MICH).
served clinal trend. Distribution and ecology. SW United States:Ari-
zona, New Mexico. Mexico: Mainly in the Sierra
Madre Occidental south to S Durango, scatteredin
Key to the varieties of Pinus arizonica
Coahuila, NE Zacatecas, and Nuevo Le6n. Forming
1. Leaves long (14-25 cm), thick (1.4-1.8 mm), with
pure stands or more commonly mixed with Quercus
8-12 lines of stomataon the abaxial side, 3-4(-5)
per fascicle......................................... [Link]. stormiae
spp., other pines (e.g., Pinus engelmannii,P strobi-
1. Leaves short or variable,<1.4(-1.6) mm thick. formis), and occasionallyJuniperusflaccidaat lower,
2. Leaves variable in length (8-)10-20(-23) cm, or J. deppeana at higher, altitudes. On various sub-
in fascicles of 3-5, predominantly3-4............ strates, but the best stands in valleys and on mesas
.......................................... 10a. var. arizonica with
2. Leaves short, (5-)6-10(-12) cm, in fascicles of deep soil, in moderately dry to mesic forest
(3-)4-5, predominantly 5................ lOb. var. with light winter frost occurring, at (1300-)2000-
cooperi
2700(-3000) m. Annual precipitationis low to mod-
10a. Pinus arizonica Engelmannvar. arizonica erate, 700-900 mm, mostly falling duringthe winter
months. Phenology: Pollen is dispersed in spring-
Fig. 25
time, dependenton altitude and latitude;exact time
Pitnus ponderosa D. Douglas ex P. Lawson var. arizonica
not recorded.
(Engelmann)G. R. Shaw, [Pines Mexico] Publ. Arnold
Representative specimens examined. MEXICO. CHI-
HUAHUA: SW of San Juanito,10 Aug 1954,
Mojarachic.
Table IV Knobloch 1164 (MICH); El Vergel, 5 Oct 1960, Little
17899 (US); Barrancade Cobre, 7 Oct 1960, Little 17914
Characterstates in the genus Pinus
(US); MesaHuracan,8 Oct 1960,Little17936(US); ca.
65 km SW of CasasGrandes,2 Jul 1952. Marshall54
Character P arizonica P ponderosa (UC);Madera,nearthe Rio Colorado,7 Jul 1958,Matuda
var. scopulorum 32667 (MEXU);vic. of Madera,May-Jun1908,Palmer
Leaves per fascicle 3-5 2-3 449 (MO);Ocampo,CascadaBasaseachicNationalPark,
Widthof leaves 0.9-1.8 mm 1.3-1.6 mm 12 Nov 1989, Spellenberget al. 10083 (FHO, NMC);Cerro
Stomatallines on abaxial Mohinora,16 km S of Guadalupe
y Calvo,11 Aug 1960,
side of leaf 8-10 Straw & Forman 1968 (MICH);Cerro Mohinora, 16 km S
(3-)4-8(-12)
Resin ducts in leaf (3-)5-10 2-6 of Guadalupey Calvo, 12 Aug 1960,Straw& Forman
Seed scales at maturity Thick, rigid Thin, flexible 2003 (MICH);63 km W of CasasGrandes,29 Aug 1952,
Tucker2582 (UC). COAHUILA:
Cuatro Ci6negas, Canon

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106 Flora Neotropica

x
Z
';

, ' '

i rai', ~'"'
~:(::::'''" "';''~";"';'":;~"as~

Os 9~~~~~~~~~~~';M,'':'i

Cb C

FIG. 25. Pinus arizonica var. arizonica (A-C, Straw & Forman1968; D, Pringle 10139). A. Branchwith leaf fascicles.
B. Leaf fascicles. C. Cross sections of leaves in (a) fascicle of 5 and (b) fascicle of 3. D. Ovuliferouscones: (a) immature
cone, (b) mature cone, (c) seed scale (three views), (d) remnants of cone bases on branch. E. Seed with wing (two
views). (Magnifications:A, B, D, E, X0.5; C, x30.)

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Systematic Treatment 107

del Agua, 8 Sep 1939, Mueller 3208 (MO, UC, US); 1977, Bye & Weber8298 (E, MEXU, NCU). COAHUILA:
"mountainsnear General Cepeda," 7 Oct 1905, Pringle Serraniasdel Burro, 7 Jun 1983, Marshall 83-28 (FHO).
10139 (E, GOET, MEXU, MO, NY, UC). DURANGO: 50 DURANGO: La Ciudad, 30 Sep 1960, Courbasson & Ma-
km WSW of El Salto at trail S to "Mexiquillo,"31 Aug hieux s.n. (P); El Salto, 23 Nov 1961, G6mez P. 612
1951, Mavsilles 7580 (MICH);Cerro Huehueto,S of Hua- (MEXU); El Salto, 18 Sep 1961, G. Guzmdns.n. (MICH.
chicheles, 23 Jul 1955, Maysilles 7981 (MICH); El Salto, SD); 5 km E of El Salto, 8 Jul 1958, Hendricks513 (MO);
4 Aug 1955, Mavsilles 8311 (MICH);Lagunadel Progreso, 58 km W of Durangoalong Hwy. 40, 7 Mar 1966, Hess &
58 km N of railroadat Coyotes, 10 Aug 1955, Maysilles Hall 554 (MICH); 10 km E of La Ciudad, 30 Apr 1987,
8337 (MICH);ChavarriaNuevo, 3 Apr 1991, Styles, Favela Hughes 977 (FHO, K, MEXU); Cerro Huehueto, 2 Jul
et al. 9 (FHO, MEXU). NUEVOLEON: Unknown loc.. 1950, Mavsilles 7262 (MICH);Cerro Prieto, ca. 35 km W
1900, Nelson 4502 (A). SINALOA:Sierra Surotato, Los of Otinapa, 10 Jul 1950, Mavsilles 7375 (MICH);Coyotes,
Pucheros,canyon below Los Pefiascos, 17 Mar 1945, Gen- 10 Aug 1952, Mavsilles 7697 (MEXU, MICH); Cerro
try 7242 (MICH, NY); "SantaLucia," Sierra MadreOcci- Chupadero,W of Pueblo Nuevo, 28 Jul 1955, Mavsilles
dental, 16 Nov 1926, Mexia 496 (MO). SONORA:Sierra 8079 (MEXU, MICH); Lagunadel Progreso, 55 km N of
Huachinera,nearsummiton E slope, 18 Jun 1953, Marshall railroadat Coyotes, 10 Aug 1955, Maysilles 8152 (MICH);
134 (UC); NW of Aribabi, Puerto de Hu6pari,in curve of WSW of El Salto, between Las Adjuntas and Rucias, 4
Rio de Bavispe, 7 Sep 1939, White2748 (MICH); Puerto Aug 1955, Maysilles 8312 (MEXU, MICH); El Salto, 24
de los Asseraderos,4 Aug 1940, White 3176 (MICH); El Aug 1957, Solbrig & Ornduff4637(NY); ChavarriaNuevo,
Temblor,in the Sierra El Tigre, 18 Aug 1940, White3360 5 Apr1991,Styles,Favelaet al. II (FHO);Coyotes.5 Apr
(MICH). 1991, Styles, Favela et al. 17 (FHO); El Madrofo, 5 Apr
1991, Styles, Favela et al. 18 (FHO, MEXU).
0Ob. Pinus arizonica Engelmann var. cooperi
(C. E. Blanco) Farjon, comb. et stat. nov. lOc. Pinus arizonica [Link] Mar-
Fig. 26A-D tinez, Anal. Inst. Biol. Univ. Nac. Mex. 16: 284.
Pious cooperi C. E. Blanco, Anales Inst. Biol. Univ. Nac. 1945. Fig. 26E-I
Mexico 20: 185. 1950; Pinus lutea C. E. Blanco ex
Martinez,Anales Inst. Biol. Univ. Nac. Mexico 16: 233. PinusponderosaDouglas ex [Link](Mart-
1945, non Walter,1788, nec G. Gordon& Glendinning, inez) Silba, Phytologia68: 59. [Link]. Mexico.
1858. Type. Mexico. Durango:El Salto, Mar 1940 by Coahuila:Sierrala Corcordia de los Patos")S
("Sierra
C. E. Blanco, M. Martinez 3442 (no original material of GeneralCepeda[locationis not in accordance with
found in MEXU; lectotype, A, here designated). thatgivenin Martinez(1948)forthisspecimen: Coahu-
Pinus lutea C. E. Blanco ex [Link] Marti- ila, [Link],Las Margaritas;probablyM's num-
nez, Anales Inst. Biol. Univ. Nac. Mexico 16: 237. bershavebeenmixedup],Jul 1941,M. Martinez3455
1945; Pinus cooperi C. E. Blanco [Link] (Marti- (lectotype,MEXU,heredesignated).
nez) C. E. Blanco, Anales Inst. Biol. Univ. Nac. Mexico
20: 185. 1950. Type. Mexico. Durango: El Salto, M.
Tree of medium size, height to 15-20 m. Leaves
Martinez3441 (holotype, MEXU).
in fascicles of 3-4(-5), thick, coarse, rigid, usually
Tree, usually tall, height to 30-35 m. Leaves in curved and twisted, 14-25 cm X 1.4-1.8 mm. Sto-
fascicles of (3-)4-5, predominantly 5, rigid to mata on all faces of leaves, in 8-12 lines on the
slightly lax, usually curved, (5-)6-10(-12) cm X 1- convex abaxial face and in (3-)4 lines on each
1.3 mm. Stomata on all faces of leaves, in 4-7 lines adaxial face (the abaxial numberof lines correlated
on the convex abaxial face and in (3-)4 lines on with width of leaf). Resin ducts in the leaf 3-5(-8).
each adaxial face (the abaxial number of lines corre- Seed cones ovoid to ovoid-oblong, often curved,
lated with width of leaf). Seed cones ovoid to broadly (4.5-)5-10(-14) x 3.5-6(-8) cm when open. Seeds
ovoid, often slightly curved, 5-10(-12) X 4-6 cm obliquely ovoid, slightly flattened,5-6 x 3.5-4 mm.
when open. Seeds obliquely ovoid, slightly flattened, Seed wings obliquely ovate, 12-16 x 6-8 mm.
5-7 x 4-5 mm. Seed wings obliquely ovate, 15-20 Distribution and ecology. Mexico: Mainly in S
X 6-8 mm. Nuevo Le6n, a few localities in S Coahuila, Zaca-
Distribution and ecology. Mexico: Mainly in Du- tecas, and possibly San Luis Potosi (dot map proba-
rango, but scattered northward in the Sierra Madre bly incomplete, but materialfrom several localities
Occidental into Chihuahua. At least in part sympatric attributedto [Link] here been determined
with var. arizonica. Similar habitat and altitudinal as var. arizonica). Habitat similar to var. arizonica,
range (mainly 2000-3000 m); associated pines are, but often recorded from more arid sites; forming
e.g., P. durangensis, P. leiophylla, P. strobiformis, P. pure stands or mixed with Quercus,Juniperus,and/
teocote. or other pines, e.g., P cembroides (especially dry
Representative specimens examined. MEXICO. CHI. sites) and P pseudostrobus. Altitudinal range ca.
HUAHUA:Bocoyna, SW of Creel near San Ignacio, 17 Oct 1500-3000 m.

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108 Flora Neotropica

,,{ A

j,,

A-D. Pinus arizonica var. cooper (A-C, Masilles 8152; & Favela 11). E-. arizonca var.
stormiae (E-G, Zobel 18; H, 1, Zobel 20). A. Branch with leaf fascicles. B, F. Leaf fascicles. C, G. Cross sections of
leaves. D. Ovuliferous cone. E. Branch with bud, immaturecones and leaf fascicles. H. Ovuliferous cone: (a) mature
cone, (b) seed scale (two views). 1. Seeds with wings. (Magnifications:A, B, D-F, H, 1, x 0.5; C, G, x 30.)
FIG. 26. D. Styles P

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Systematic Treatment 109

Representative specimens examined. MEXICO. is therefore more in accordance with the evidence
NUEVO LEON:3 km S of Pablillo,20 Jul 1958,Correll& presentlyavailable.
Johnston19914(NY);Canondel Aserradero, Miquihuana, Martinez (1948) recognized Pinus arizonica var.
Nov 1939,M. Martinez3455 (numberprobablyincorrect,
see above under type) (NY); Galeana, 20 km E of San stormiae, which he described as different from P
Roberto,4 Nov 1983, McCarter& Hughes105 (FHO); arizonica in its larger (longer and wider) leaves,
Puertode Reyna,14Jul1977,Mittak8492(FHO);Galeana, fewer resin ducts in the leaves, and largercones. The
14 Jul 1977, Mittak 8493 (FHO); I km from Iturbide, 12 other characterstates mentioned seem to be closer
Mar 1980, Stead & Styles 614 (FHO, MEXU); 30 km from
to P ponderosa var. scopuloruinthan to P arizonica
[Link] Iturbide,13 Mar1980,Stead& Styles
628 (FHO);Ascensi6n,Dec 1955,Zobel18 (UC);Galeana, var. arizonica, but Martinez did not consider P
Dec 1955, Zobel 20 (UC). ponderosa as occurringin Mexico. Unlike var. coo-
peri, var. stormiae has a more disjunct distribution
Uses. Pinus arizonica is an importantconstituent from the mainly western range of the species.
of the pine and pine-oak forests of northernMexico,
especially in the Sierra Madre Occidental. As such 11. Pinus jeffreyi J. H. Balfour in A. Murray,Bot.
it is heavily exploited as a timbertree in much of its Exped. Oregon 8: 2. 1853. Fig. 27
range, especially in the more accessible regions, and PinusponderosaD. Douglasex P. [Link]
stands with large trees in these localities are now (J. H. Balfour)Vasey,[Link].S. 31. 1876
rare (Perry, 1991). [& U.S. [Link]. 1875:179. 18761;Pinus
Three varieties of Pinus arizonica have been rec- ponderosaD. Douglasex P. [Link](J.
H. Balfour)E. Murray,Kalmia12: 23. 1982. Type.
ognized, based mainly on a combination of leaf [Link]: ShastaValley,("Chastey Val-
characterswhich are quantitativeand mostly continu- ley,Lat.41?30'N"),24 Oct 1852,Jeffrey731(holotype,
ous. We have not found any distinct charactersto E; a poorspecimenwith4 leaf fascicles,a budanda
separate P. cooperi C. E. Blanco from P arizonica brokenseed anda few seed wings).[Epitype:United
[Link]: TrinityCo., ScottMt.,2 Sep 1958,
Engelm. to consider at the rankof species. Alava et al. 2439A (E), here designated].
Martinez(1945) gave a Latindescriptionof a new [Link]. Torreyin Emory,U.S. [Link].
species of pine from Durango (El Salto). It was Surv. Rpt. 2(1): 209, t. 56. 1859; Pinus jeffreyi J. H.
based on a specimen, accompaniedby a description, [Link](J. Torrey)J. G. Lemmon,Calif.
sent to him by C. E. Blanco under the suggested [Link].2: 74, 100. [Link].
[Link]: "Mountains E of SanDiego,"
name Pinus lutea, presumablynamed for its yellow
1850,Parrys.n. (lectotype,NY,heredesignated).
wood. This name is a later homonym and for this Pinus jeffreyi J. H. Balfour var. baja-californica Silba.
reason the new species was renamed P. cooperi by Phytologia 68:52. [Link]
Blanco (1950). A search by A. Farjon for the type Norte:BetweenOjosNegrosandNeji Rancho,16 Sep
1929, 1. L Wiggins & Gillespie 4123 (holotype, NY;
specimen in MEXU has been [Link]
P isotypes, F. MEXU. MICH, SD).
(1945, 1948) compared this species with rudis (=
P hartwegii) and P. montezumnae, from which it is Tree, medium to large, height to 20-30 m, dbh to
more distinct morphologically (especially in cone 100 cm. Trunk monopodial, straight, terete. Bark
characters)than from P arizonica. Martinez(1945) thick, rough and scaly, divided by deep, longitudinal
also proposedthe variety P lutea [Link], being and anastomosing fissures into thick, elongated
a 5-leaved form with dark green, longer leaves and plates, light buff to light brown, the fissures dark
largercones. We found these charactersto be incon- grey-brown,on young trees and branchesrough and
sistent (e.g., Styles, Favela et al. 17 (FHO) with scaly, [Link] of first order long,
leaves 6-8 cm and a maturecone 8 cm long; Styles, slender, spreading horizontally or assurgent at the
Favela et al. 18 (FHO) with leaves 11-13 cm and a ends, persistent; branches of higher orders sparse,
cone 5 cm long). This taxon was later recombined curved or assurgent,forming an open pyramidalor
by Blanco (1950). Unlike Perry(1991: 109, t. 3.10), flat-topped crown. Shoots stout, assurgent, very
we do not think that the bark between these taxa rough and scaly with persistent, decurrentpulvini,
differs consistently in the manner he described, light orange-brown,often glaucous. Cataphyllslarge,
which follows from comparison of his published ca. 20 mm long, triangular-subulate, reflexed, scari-
photographswith those given by Martinez (1948: ous, with erose-ciliatemargins,light brown,weather-
fig. 195) alone. Other differences, such as those ing greyish black. Vegetativebuds ovoid-acute or
mentionedfor several cone characters,have not been subglobose, the terminal buds 15-20 mm long, the
confirmedin the collections we have [Link]- laterals smaller, not resinous; the scales imbricate,
ment of P cooperi as a variety of P arizonica subulate,with free apices and ciliate, white margins,

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110 Flora Neotropica

I
/ I

\,s/ t? aC 2D

FIG. 27. Pinus jeffreyi (A, B, Hughes & Styles 170: C-E, Hughes & Styles 181). A. Branch with leaf fascicles and
cataphylls. B. Leaf fascicle and cross section of leaf. C. Ovuliferouscone. D. Seed scale (three views). E. Seed. (Magni-
fications:A, C-E, x 0.5; B. x25.)

light [Link] sheaths of young leaves in older fascicles, weatheringgrey-brownto blackish.

20-25 mm long, with 8-10 pale orange-brown,im- Leaves in fascicles of 3, sometimes a few fascicles
bricate scales, persistent but reduced to 10-15 mm of 2, in dense, spreading tufts on thick branchlets,

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Systematic Treatment 111

persisting(3-)4-5(-6) years, thick, rigid, straightor

slightly curved, sometimes twisted, (12-)15-22(-25)
cm x 1.5-1.9(-2) mm, with serrulate margins,
acute-pungentto acuminate,light yellowish green to
greyish green. Stomata on all faces of leaves, in 7-
11 grooved lines on the convex abaxial face, in 3-5
conspicuous lines on each adaxial face. Leaf anat-
omy: Cross section transverse-triangular, with a con-
vex abaxial side; epidermis thick; hypodermismulti-
layered, slightly intrudinginto the mesophyll; resin
ducts (2-)3-5, medial, the three marginalducts larger
than any subsidiaryducts; stele oval in cross section;
the outer walls of endodermalcells thickened;vascu-
lar bundles 2, distinctly separated. Pollen cones
densely clustered near the proximal end of a new
shoot, subtendedby subulatebracts,ovoid-oblong to
cylindrical, 20-35 x 6-7 mm, yellow or purplish
yellow, maturinglight brown. Seed cones subtermi- Q
I
nal, solitary or in pairs on short, stout, persistent
peduncles, spreadingand seemingly sessile at matu-
rity, leaving a few basal scales on the branch when ^V^ \ o
falling. Immaturecones broadly ovoid, ca. 30 X 20
mm, with spreading spines, purplish, soon light
brown, maturing in two seasons (16-18 months).
Mature cones broadly ovoid to subglobose, with a
slightly oblique, flattenedbase, (10-)12-17 X 9-14
cm when open. Seed scales ca. 150-175, in a low
spiral, parting soon and wide, thin woody, straight
or slightly curved, up to 20 mm wide, light or dark
brown abaxially, with distinct, lighter marksof seed
wings on the adaxial side. Apophysis slightly raised, Map 8. Piniis jeffreyi.
transverselykeeled, broadly rhombic in outline, on
the proximal scales more gibbous, often resinous, northernregions, the persistence of leaf fascicles is
ochraceousto light brown. Umbo dorsal, moderately also reported(Bradshaw, 1941) to be longer, up to 8
raised, transverselykeeled, terminatingin a distinct, years. The almost brick-redto red-brownbark seen
curved, 3-5 mm long spine. Seeds obliquely ovoid, by A. Farjon in parts of the Sierra Nevada has not
slightly flattened, 9-12 mm long, light yellowish been observed in the Mexican populations.
brown with faintly darker spots. Seed wings articu- Distribution and ecology (Map 8). United States:
late, effective, held to the seed by two thin claws, Primarily in California, extending into SW Oregon
oblique, with a straightand a curved side, 20-25 X and extreme W Nevada; its main distributionis in
10 mm, light yellowish brown to light brown. Num- the Sierra Nevada; there are disjunct populationson
ber of cotyledons 7-13, usually 10. mountainsin S [Link]: In Baja Califor-
Remarks on morphology. The above description nia Norte, scattered in the Sierra Juarez, but more
pertainsto Pinus jeffreyi in Mexico only. Generally, common in the northernpartof the SierraSan Pedro
both tree size and sizes of leaves and cones can be Martir,where it reaches its southernmostlimit at ca.
larger in the United States, especially in the Sierra 30?40'N. Its altitudinalrange in the Sierra Juarez is
Nevada of California. There trees up to 50 m or (1 100-)1400-1800 m; in the SierraSan Pedro Martir
more occur, to which figures given by Martinez it occurs considerablyhigher, 1800-2500(-2700) m.
(1948) and Perry (1991) must be referring,although Here it forms an open, mixed coniferous forest with
theirbooks deal with Mexican pines. Similarly,cones other conifers. Common are also Abies concolor,
in the Sierra Nevada can be up to 25-30 cm long, Pinus lambertiana, and P contorta var. murravana
but we have not seen such large cones on our visits in the upper zone. In the Sierra Juarez it is often
to the populationsof this tree in Mexico. From more associated with Quercus spp. Both mountainranges

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112 Flora Neotropica

are graniticand provide acid soils. Annual precipita- view (e.g., Kral, 1993) that the two taxa are suffi-
tion is moderate, up to 500 mm, of which about ciently distinct to be recognized as species. Interme-
half comes as winter snow at the higher altitudes. diate forms have been recognized as naturalhybrids
Phenology: Pollen is dispersed in June-July, de- (Haller, 1962), but these are [Link] con-
pending on altitude and related climatic circum- trolled conditions, crossability is relatively low
stances. (Burns & Honkala, 1990). As it has been established
that P. ponderosa is absent from Baja California
[Link] (Duffield & Cumming, 1949), no such hybrids are
CALIFORNIA NORTE:Sierra Juarez, San Faustino, 15 km
S of RanchoEl Compadre,off Tecate-Rumorosa rd., 26 to be found there.
May 1987,Hughes & Styles 181 (FHO); Mun. Ensenada. The more or less disjunct populations in Baja
La Cienega, Jan 1942, M. Martinez 2148 (NY); Sierra California have been described by Silba (1990), on
Juarez,80 km S of Tecate,Sep 1939,M. Martinez3451 narrow evidence, as P.
very var.
jeffreyi baja-califor-
(MEXU); 1.5 km N of San Faustino, Moran 16605 (SD);
nica. His observation that the leaves are "mostly
SierraSan PedroMartir,20 May 1893, Brandegees.n.
(UC); 2 km N of Vallecitos,1 Feb 1971, Echenique46 two per fascicle" is contradicted by the holotype
(MEXU);SierraSanPedroMartir, [Link] observatory, specimen (/. L. Wiggins & Gillespie 4123, NY) and
1790m, 25 Apr 1987,Hughes& Styles170 (FHO),2320 by all other materialwe have seen in and from Baja
m, 25 Apr 1987,Hughes& Styles175 (FHO,K); Sierra California, in which
SanPedroMartir,1889,Townsend s.n.(US);LaEncantada, only an occasional fascicle has
18 Sep 1930, Wiggins & Demaree 4920 (F, NY, US). 2 leaves. Also are the leaves not wider than in
specimens from [Link] size appearsto be
Uses. Although not an importanttimber tree in smaller on average, but this character is of little
Mexico, its geographical isolation from other Mexi- taxonomicvalue (Duffield & Cumming, 1949). Vida-
can pine forests makes it of local or regional impor- kovic (1991) mentions a variety"peninsularis"from
tance. Especially in the Sierra Juarez, exploitation Baja California with "darkergrey and deeply fur-
may pose a threat to the limited and more scattered rowed bark"but does not cite a valid publicationof
populations. Many of the stands in the Sierra San this name.
Pedro Martirare within a national park and reason-
ably well-protected. 12. Pinus engelmannii Carriere,Rev. Hort., ser. 4,
Pinus jeffrevi was found by the Scottish plant 3: 227. 1854. Fig. 28
collector John Jeffrey in northern California, in
Shasta Valley, on 24 October 1852. He had been Pinusmacrophylla Engelmann,non Lindley(1839), in
Wislizenus, Mem. Tour N. Mexico 103. 1848; Pinus
sent out by the Committee of the Oregon Botanical
ponderosa D. Douglas ex P. Lawson var. macrophylla
Expedition, on behalf of its subscribers,to collect (Engelmann)G. R. Shaw, [Pines Mexicol Publ. Arnold
seeds of exotic plants. When the first batch of mate- Arbor. 1: 24. 1909. Type. Mexico. Chihuahua:Cosiquir-
rial finally arrived in Edinburghin the summer of iachi, Oct 1846, Wislizenus 233 (lectotype, MO
3377671, here designated; isolectotypes, MO 3377672,
1853, a list of its contents, including new species MO 3377673, MO 3534934).
with descriptions,was publishedas a pamphlet(No. Pinus latifolia C. S. Sargent, Gard. Forest 2: 496, fig.
8) "by order of the committee," by its secretary, 135. 1889; Pinus lmavrianaSudworth, U.S.D.A. Div.
Andrew [Link] Murray(1853) ascribes Forest. Bull. 14: 21. 1897; Pinus ponderosa D. Douglas
ex P. Lawson var. mavriana (Sudworth)C. S. Sargent,
the new conifer taxa to Balfour (who was the chair;
Silva N. Amer. II: 81. 1897. Type. United States.
Greville, often cited as co-author, did the illustra- Arizona: S slopes of Santa Rita Mts., 1887, H. Mavr
tions), the authoritywith the new species is cited as s.n. (holotype, A).
"Oreg. Corn.," which has led to the interpretation Pinus apacheca J. G. Lemmon. Erythea 2: 103, t. 3.
that Murray,not Balfour, was the validating author 1894. Type. United States. Arizona: Chiricahua Mts.,
Jun 1892, Lemmons.n. (lectotype, A, here designated).
(Farjon, 1993). This view is here revised in accor- Pinus [Link] Martinez,An-
dance with Art. 46.2 of the Tokyo Code (Greuteret ales Inst. Biol. Univ. Nac. Mexico 15: 345. 1944,
al.. 1994). The holotype, Jeffrey 731 (E), is such a "'macrofila";Pinus engelnannii Carriere var. hlancoi
poor specimen that use has been made of the new (Martinez) Martinez, Pinos Mexic., ed. 2, 288. 1948.
in the Code to an Type. Mexico. Durango:(Near) El Salto, Aug 1940, M.
provision (Art. 9.7) designate Marlinez 3450 (lectotype, MEXU 2509, designated by
epitype to accompany it. Carvajal& McVaugh, 1992: 59).
Formerly classified as a variety under P pon-
derosa, more detailed research(Bradshaw,1941; Jo- Tree, medium to large, height to 20-25(-27) m,
hansen, 1953) has led to the now generally accepted dbh to 70-90 cm. Trunk monopodial, straight, erect,

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Systematic Treatment 113

\\\\ \\

FIG. 28. Pinus engelmrannii(A-C, Mueller 3436; D, Pringle 13614; E, F, Mexia 2515; G, Styles, Far'elaet al. 3). A.
Branchwith leaf fascicles. B. Leaf fascicles. C. Cross section of leaf. D. Leaf fascicles and immatureovuliferous cone.
E. Ovuliferous cone. F. Seed scale (three views). G. Seed (two views). (Magnifications:A, B, E-G. x0.5; C, x 20; D.
x 1.)

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114 Flora Neotropica

usually free of branchesonly in the lowest part('/4- to 20 mm wide, dark purplishor blackish brown on
'I/) or to '/2 of height. Bark thick, rough, scaly, either side, with light brown marks of seed wings
divided into long, irregularplates by wide, shallow, on the adaxial side. Apophysis prominently raised,
dark fissures, reddish brown on branchesand young transverselykeeled, often recurvedand slightly larger
trees, then dark brown, weathering grey. Branches on one side of the cone, rhombic or pentagonal in
of first order thick, long, spreadingor assurging;of outline, ochraceous or light brown, often with dark
second and higherorderssparse,stout, spreadingand radial lines or fissures. Umbo dorsal, large, rhombic
assurging, curved, forming a broad, rounded, open to transverse-rhombicin outline, with a persistent,
crown. Shoots stout, very rough, with large, decur- curved spine up to 3 mm long, slightly darkerbrown
rent, persistentpulvini. Cataphyllslarge, 15-20 mm than apophysis, weathering grey-brown. Seeds
long, subulate, recurved, with scarious lamina and obliquely ovoid, slightly flattened,keeled, 5-8 X 4-
erose-ciliate margins, dark brown to blackish, often 5.5 mm, light grey-brown, often with dark spots.
caducous. Vegetativebuds large, the terminal buds Seed wings articulate,effective, held to the seed with
20-30 x 15-20 mm, ovoid-conical, the laterals two claws or (partly)covering the seed on one side,
smaller, not resinous; the scales spreading or re- ovate-oblong to obliquely ovate, 18-25 x 7-10
flexed, subulate-caudate,with long-ciliate margins, mm, translucentlight-ochraceous,often with a darker
brown. Fascicle sheaths persistent and remaining tinge nearthe apex. Seedlings: Numberof cotyledons
long, (15-)25-35(-40) mm, consisting of 10-12 im- 8-10; a conspicuous "grass stage" is reported for
bricate, distinct bracts, reddish brown with greyish this species (Bailey & Hawksworth, 1983).
ciliate margins, weathering brown to blackish. Distribution and ecology (Map 9). United States:
Leaves in fascicles of (2-)3(-4), rarely 5, in dense, SE Arizona and extreme SW New Mexico. Mexico:
spreading tufts toward the ends of branches, per- Extending south from the populations in the SW
sisting 2-3 years, thick, rigid, straight or slightly United States through the Sierra Madre Occidental
drooping, (18-)20-35 cm x 1.5-2 mm, margins in Sonora, Chihuahua, NE Sinaloa, Durango, and
serrulate, apex acute-pungent, light (yellowish- more scattered in Zacatecas. It has been cited from
)green. Stomata on all faces of leaves, in (6-)8-15 the SierraFriabetween ZacatecasandAguascalientes
lines on the convex abaxial face and 4-8 lines on (see Critchfield & Little, 1966: map 49), but no
each adaxial face. Leaf anatomy:Cross section trian- collections from there have come to our attention;
gular or transverse-triangular, with a convex abaxial Perry (1991) could not find it there. It also appears
side; hypodermis multi-layered, slightly intruding to occur in Nuevo Le6n, in the mountains near
into the mesophyll; resin ducts (4-)5-8(-14), medial, Galeana, from where two collections could be veri-
the 3 marginalducts usually largerand always pres- fied. Other reportsof disjunct populationsappeared
ent; stele oval in cross section; outer walls of endo- to be based on specimens of Pinus durangensis or
dermalcells not or slightly thickened;vascularbun- could not be verified and have been omitted. Pinus
dles 2, separate or the xylem strands more or less engelmannii occurs on moderately dry, summer-
connate, always distinct. Pollenlcones clustered near warm open mountainslopes or plateaus at (1200-)
the proximal end of a new shoot, subtended by 1500-2700(-3000) m, most abundantly at 2000-
subulate bracts, ovoid-oblong to cylindrical, up to 2500 m. It occurs on poor rocky (volcanic) soils as
40 mm long and 12 mm wide, yellowish pink, turning well as on alluvial coarse sand/gravelor loamy sand.
yellowish brown. Microsporophyllswith a 1.5 mm The climate is temperate,with annual rainfall 400-
wide peltate, more or less cordate distal part, which 700 mm, [Link] 2000 m, frost
appearsradially [Link] cones subterminal,in and snow are common in winter. It is a constituent
pairs or in whorls of 3-5, on thick, short, curved of open pine and pine-oak woodland, sometimes of
peduncles,persistingsome time, leaving a few scales mixed pine forest, with, e.g., P leiophylla, P lumhol-
on the branch when falling. Immaturecones ovoid, tzii, and P pseudostrobus, on drier sites with P
with prominent, spreading or slightly recurved cembroidesand Juniperussp., and usually with vari-
spines, light brown, often glaucous, maturingin two ous species of Quercus present. Phenology: Pollen
seasons. Maturecones seemingly sessile, ovoid-ob- dispersal is reportedto occur in May (Arizona); the
long, curved, with an oblique base when opened, 8- time is likely to be dependenton altitudeand can be
15 x 6-10 cm when open, often resinous. Seed some weeks later at the highest elevations.
scales ca. 100-140, parting to release the seeds
except at the base, oblong, straightor recurved,thick Representative specimens examined. MEXICO. CHI-
woody, with irregularlyundulating,thin margins,up HUAHUA:Guasarachic, 26 Sep 1898, Goldman 163 (A,

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SystematicTreatment 115

I I
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~?,

t~~~~~~~~~~~~~?13 1~~~~~~~~~~~~~~~~~~~~~~~~~~~'
?
15~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
?

:..' .:
'i~?
IVr

i 'l

'
t ,i .,,,,.UIL L ? U '' ? , I
...I..
? ?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ ..
r I: I F1~ "~ rl??L!???1-?T-? ,. ,......,,..,.....
r ?...... N .......
' ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~
~~~~~~~~~~~~~~~~~~~~~~~~~~.'
'~ ~~~~~~~~~~~~~~~.......~
"?~~~~~~~~~~~~~~~~~~
'''' t ~~~~..
... ?t? r~
?~~~~~~~~~~~~':...
.. ...

SCUM~~~~~~ i~~~~~~~~~-.
-. ."
-.I

?
i ,~~~~~~~~~~~~~~~~~~~~~~~~~~~~ '

?r~~~~~~~~~ru~~~~~ I ......
' ' I

" :

' ld~a~l
~ '

~ i
~ ~I .
.

.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~;
Map 9. Piu (crls;Pd?lsaa(rage)
enelani
'I"~~~ ....... ~~
~~~~~~~~~N'
' r i

rrrar ~~i... .. . . . . ?'''

(triangles).
Map9. [Link](circles);P:[Link]

US); in Sierra Madre SW of Colonia Judrez, Dec 1906, Fisher 44236 (MO, NY); along rd. from Durango to El
Goodding 2108 (A, GH, MO), Dec 1906, Goodding 2109 Salto, nearjct. to Otinapa.25 Mar 1978, Klaus s.n. (WU);
(UC); W of Chuhuichupa,26 Aug 1936, LeSueur551 (UC); La Punta,Nov 1940. M. Martinez2346 (US); near El Salto,
Sierra San Luis, I I Jul 1892. Mearns 424 (US); Basuchil, Nov 1941, M. Martinez 2363 (NY); Cerro Prieto, 30 km
near Cerro de la Ventana, 16 km NW of Miifaca, 10 May W of Otinapa, 10 Jul 1950, Maysilles 7376 (MICH); 10
1929, Mexia 2515 (A, E, MICH, MO, NY, P. UC); Arroyo km W of Durango, 5 Jan 1983, Neill 5314 (MICH. MO):
Ancho, 7 Jun 1929, Mexia 2608 (A, BM, MICH, MO, NY, El Salto, 12 Jul 1898, Nelson 4568 (A, K, US); near
UC); Canon Huhuatan,13 km SE of Madera,23 Sep 1939, Durango at Santiago Papasquiaro,Apr 1896. Palmer 776
Mueller 3436 (GH, MICH, MO, UC); Guadalupe, 12 Sep (A, B, F, K, MO, NY); E slope of Sierra de la Ulama,
1898, Nelson 5013 (K); vic. of Madera, May-Jun 1908, Cienega de los Canelos, 1911, Patoni 22 (MEXU): Sandia
Palmer330 (GH, MO, NY); "gravelly plains at the base of railway sta.. 13 Oct 1905, Pringle 13614 (MICH. US);
the Sierra Madre." Sep 1887, Pringle 1448 (A, MEXU, Agua Bendita, 1932, RamirezL. 22 (MEXU); 37 km SW
MO, NY, US); Sierra Madre, 15 Oct 1888, Pringle 1673 from Durango, 2 Apr 1991. Styles. Favela et al. 3 (FHO).
(P, S, UC); ca. 43 km S of Creel along rd. to La Bufa, 6 NUEVOLE6N: Near Galeana, 1 km N of 18 de Marzo. 29
Jul 1969, Soule 513 (MO); Cascada Basaseachic, 11 Nov Oct 1984, Rushforth495 (E); Pablillo, 8 Aug 1936, Tarlor
1986, Spellenherg et al. 10069 (FHO, NMC); Colonia 104 (MO). SINALOA: SierraMonterrey,II Mar 1940. Gen-
Garcia, 26 Sep 1899. Townsend & Barber 356 (FHO, tr' 5904 (NY). SONORA:Cailon de Santa Rosa, E of El
MICH,MO, NY, P); Majalca(Pilares), 11 Aug 1939, White Tigre, 16 Jul 1938, White 528 (MICH); Cation de los
2344 (MICH). COAHUILA: Sierra de la Madera, Canon Metates, 24 Jun 1940, White2829 (GH, MICH);Puertode
Charretera,I I Sep 1941, Johnston 8935 (MO). DURANGO: los Asseraderos, 4 Aug 1940, White 3175 (GH, MICH);
43 km W of Durango, along Hwy. 40, 24 Jul 1958, Cor- Rancho de la Nacha, 15 Aug 1941, White 3943 (MICH,
rell & Johnston 20120 (NY); Tepehuanes, 30 Jul 1944, NY); Rancho del Roble. NE of El Tigre, Sep 1941. White

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116 Flora Neotropica

4266 (MICH, NY); summit of pass between Imuris and either are inconsistent, with the leaves "por lo general
Cananea,8 May 1948, Wiggins11684 (MICH. UC, US). algo menas robustas" but still 2 mm wide (the maxi-
mum found in the species), or fall within the range
Uses. Harvested but apparently not specifically of character states found in material from widely
selected as a timbertree in most of its range together separated locations, such as cone colour and shape
with other pines; in some areas depletion of larger of seed scales and apophysis. We agree with Perry
trees has been observed (by B. T. Styles). (1991) and Carvajal and McVaugh (1992) that Marti-
Engelmann (1848) described a new species in nez's variety is insufficiently distinct for taxonomic
Pinus from "the higher mountains of Chihuahua" recognition.
with the binomial Pinus macrophylla,a later hom-
onym of P macrophylla Lindley (1839). Carriere 13. Pinus hartwegii Lindley, Edwards's Bot. Reg.
(1854) renamedit and repeatedEngelmann'sdescrip- 25: 62. Aug 1839 [& Allg. Gartenzeitung 7: 324.
tion. The type specimen (Wislizenus233, MO) was 18391. Fig. 29
collected in 1846. Sheet MO 3377671, with a label
Pinus nontezunaeA. B. [Link](Lindley)
written by Engelmann, was considered to be the G. R. Shaw, [Pines Mexico] Publ. Arnold Arbor. 1: 23.
holotype (Carvajal & McVaugh, 1992). Two more 1909. Type. Mexico. Michoacan: "Campanario,in the
sheets (MO 3377672 and 3377673), bearing Engel- mountainsof Angangueo," 1838, Hartweg s.n. (lecto-
mann's and Wislizenus's labels, respectively, and a type, P, here designated;isolectotype, W).
Pinus ehrenbergiiEndlicher,Syn. Conif. 151. 1847; Pinus
very fragmentedcone (MO 3534934) can be consid- decandolleana Roezl var. ehrenhergii(Endlicher) Car-
ered isolectotypes, as they are likely to belong to the riere, Trait6 Gen. Conif., ed. 2, 2: 570. 1867. Type.
same [Link] cone is now entirely broken up Mexico. Hidalgo: Mineral del Monte, Dec 1837, Eh-
but the scales are typically thick woody and have renherg 915 (holotype. W; isotype, HAL).
Pinus rudis Endlicher, Syn. Conif. 151. 1847; Pinus
prominent, recurved apophyses. Parlatore (1868),
montezulnae A. B. Lambert var. rudis (Endlicher)
who had received part of the type materialon loan, G. R. Shaw, [Pines Mexico] Publ. Arnold Arbor. 1: 22.
included neither Engelmann'sspecies nor Carriere's 1909; Pinus hartwegii Lindley var. rudis (Endlicher)
new name in the text of his [Link],Shaw Silba, Phytologia 68: 50. 1990. Type. Mexico. Locality
(1909) treated it as a variety of P ponderosa under not stated, 1837-1838, Ehrenberg 1083 (lectotype, W,
the broadest possible concept of that species, re- here designated).
Pinus northumherlandianaRoezl, Cat. Grain. Conif.
taining Engelmann'sepithet. Mexic. 13. 1857. Type. Mexico. Mexico: Mt. Popocate-
Sargent (1889), Lemmon (1894), and Sudworth petl, 1857, Roezl s.n. (lectotype, Fl, here designated;
(1897) recognized that this pine was not identical isolectotype, P).
with P ponderosa when it was encounterednorthof Pinus lowii Roezl, Cat. Grain. Conif. Mexic. 18. 1857.
the Mexican borderin Arizona,and they described it Type. Mexico. Locality not stated, 1857, Roezl s.n.
(lectotype, Fl, here designated).
underthree different names: respectively,P latifolia Pinus robustaRoezl, Cat. [Link]. Mexic. 19. 1857.
(later dropped in favour of a treatmentas a variety: Type. Mexico. Mexico: Ajusco Mts., 1857, RoezI s.n.
P ponderosa var. mayriana by Sargent, 1897), P (lectotype, Fl, here designated).
The two Pinus endlichericna Roezl, Cat. [Link]. Mexic. 19.
apacheca, and P mayriana. first binomials
1857. Type. Mexico: Ajusco Mts. Original materialnot
are based on different specimens from Arizona, and located.
the latter is a nomenclaturalsynonym for P latifolia Pinus wilsonii Roezl, Cat. [Link]. Mexic. 19. 1857;
C. S. Sargent(Little, 1944). Pinus decaisneana Roezl var. wilsonii (Roezl) Carriere,
Martinez (1944), who had no access to much of Trait6Gin. Conif., ed. 2, 2: 544. 1867. Type. Mexico.
the older Europeanliterature,still used Engelmann's Hidalgo: Pachuca, 1857, Roezl s.n. (lectotype, Fl, here
designated).
epithet for the species but correctedthis later (Marti- Pinus resinosatRoezl, Cat. [Link]. Mexic. 30. 1857,
nez, 1948) and published a new variety: P macro- non Aiton, 1789; Pinus roezlii Carriere, Traite Gen.
phylla var. blancoi (= P engelmannii var. blancoi). Conif., ed. 2, 2: 575. 1867, nom. [Link]. Mexico.
The lectotype, designated "by default" (Carvajal& Locality not stated, 1857, Roezl s.n. (lectotype, Fl, here
designated).
McVaugh,1992) is M. Martinez3450 (MEXU 2509), Pinus iztacihuatlii Roezl, Cat. Grain. Conif. Mexic. 30.
labeled "El Salto" and "[Link]." by Martinez,and 1857. Type. Mexico. Mexico: VolcanIztaccihuatl,1857.
probablycollected by C. E. Blanco in August 1940. RoezI s.n. (lectotype, FI, here designated; isolectotype,
Martinezapparentlywrote the labels afterwardand, P).
Pinus scoparia Roezl, Cat. [Link]. Mexic. 30. 1857.
because of this method (see McVaugh, 1972), ended
Type. Mexico. Locality not stated, 1857, Roezl s.n.
up with a differenttype locality (Santiago Bayacora) (lectotype, FI, here designated).
in the protologue. The minor differences described Pinus standishii Roezl, Cat. Grain. Conif. Mexic. 30.

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Systematic Treatment 117

FIG. 29. Pinus hartwegii (A-C, E, [Link] & Syles 5; D, Stead 303; F, Stead & Styles 625). A. Branch
with leaf fascicles and terminal bud. B. Leaf fascicle. C. Cross section of leaf. D. Shoot with leaf fascicles and pollen
cones. E. Shoot with ovuliferous cone. F. Ovuliferous cone. G. Seed scale (two views) and abortedseed. H. Seeds with
articulatewings. (Magnifications:A, B, D-H, x0.5; C, x 25.)

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118 Flora Neotropica

1857. Type. Mexico. Locality not stated, 1857, Roezl 10(-11) lines on the convex abaxial face and in 4-
s.n. (lectotype, Fl, here designated;isolectotype, P). 6(-7) lines on each adaxial face. Leaf anatomy:
Pious aculcensis Roezl, Cat. Grain. Conif. Mexic. 30. Cross section triangularor transverse-triangular,with
1857. Type. Mexico. Locality not stated, 1857, Roezl
s.n. (lectotype, Fl, here designated; isolectotype, P). convex abaxial side; cuticula and epidermis thick;
Pinus amecaensis Roezl, Cat. Grain. Conif. Mexic. 30. hypodermisnearly uniform,with 2-3 layers of cells;
1857. Type. Mexico. Mexico: Locality not stated, 1857, resin ducts (2-)4-10(-12), occasionally absent, me-
Roezl s.n. (lectotype, P, here designated). dial, occasionally 1-2 internal; stele oval in cross
Pinus papeleuii Roezl, Cat. Grain. Conif. Mexic. 31.
1857. Type. Mexico. "On Popocatepetlor Iztaccihuatl, section; outercell walls of endodermisnot thickened;
near 13000-14000 ft.," 1857, Roezl s.n. (lectotype, Fl, vascular bundles 2, distinct but approximateor the
here designated). xylem strands connate. Pollen cones crowded near
Pinus lindlevana G. Gordon & Glendinning, Pinetum the proximal end of a new shoot, subtended by
229. 1858 (nom. superfl.); Pinus montezlonae A. B. subulate bracts, ovoid-oblong to cylindrical, ulti-
Lambert var. lindlevana (G. Gordon & Glendinning)
Parlatorein Candolle, Prodr. 16(2): 399. 1868. Type. mately 20-25 X 6-7 mm, yellowish or pink-pur-
Mexico. Locality not stated, n.d., Hartwvegs.n. (holo- plish, maturingto brown. Seed cones subterminal,in
type, K). pairs or whorls of 3-6 on short, stout, bracteate
Pinus suffruticosaRoezi ex Carriere,Trait6Gen. Conif.,
peduncleswhich are hiddenby basal scales of mature
ed. 2, 2: 584. 1867. Type. Mexico: Original material
cones. Immaturecones ovoid-oblong, often slightly
not located.
Pittus donnell-smithiiM. T. Masters, Bot. Gaz. 16: 199. curved, purplishbrown to purplish black, maturing
[Link]. Volcande Agua,
Sacatepequez: in two years. Maturecones ovoid-oblong, with acute,
near the summit, Apr 1890, Donnell-Smith2182 (holo- curved apex when closed, (obliquely) ovoid with a
type, US; isotype, K). flattenedbase when opened, 8-12(-14) x 5-8 cm.
Seed scales ca. 150-200, partingto release the seeds,
Tree, tall, erect, stunted near tree line, height to spreadingwide, often recurvednearthe base, oblong,
25-30 m, dbh to 80-100 cm. Trunk monopodial, thin woody, flexible or more rigid, purplish brown
erect, terete, bole free of branchesfor 2/ of length, to blackish with a light brown margin below the
or retaining old branch stumps. Bark thick, scaly, apophysis abaxially, brown with lighter marks of
deeply fissured, divided into small or large, square seed wings on the adaxial side. Apophysis more or
or irregularplates, outerbarkdarkbrown,weathering less flat, (weakly) transversely keeled, sometimes
grey, on young trees and branches scaly, brown. gibbous, rhombicin outline with an angularor irreg-
Branchesof firstorderthick, often twisted, spreading ular upper margin, varying from light brown to
and assurging, often persistent; branches of higher purplishbrown with a blackishcentre. Umbo dorsal,
orders stout, spreading and assurging, forming a depressed, flat or raised, up to 4 mm wide, obtuse
dense, roundedcrown in maturetrees. Shoots thick, or with a deciduous prickle, grey or black. Seeds
rigid, scaly with persistent, decurrentpulvini, pur- broadlyovoid, slightly flattened,5-6 mm long, light
plish brown, sometimes glaucous. Cataphylls large, or darkbrown with black spots. Seed wings articulate
15-20 mm long, subulate,recurved,scarious,brown, but effective, held to the seed by two oblique claws,
weatheringblackish grey, with ciliate to erose mar- oblong with a straightand a curved side, 12-20 x
gins, usually persisting longer than leaf fascicles. 7-12 mm, browntinged with [Link]
Vegetativebuds ovoid-conical, large, terminal buds (4-)5-6(-7).
up to 30 x 15 mm, the lateralssmaller,not resinous; Distribution and ecology (Map 10). Mexico: In
the scales imbricatebut free at apex, subulate,scari- Chihuahua(Cerro Mohinora),S Coahuila, S Nuevo
ous, reddish brown with ciliate margins, giving the Leon, Durango, SW Tamaulipas, Jalisco (Nevado
buds a rugose [Link] sheaths of young de Colima), Michoacan, Mexico, Morelos, Hidalgo,
leaves long, 30-40 mm, with ca. 10 imbricate,ap- DistritoFederal,Tlaxcala, Puebla,W Veracruz,Guer-
pressed scales with ciliate margins, light brown, rero (CerroTeotepec and vicinity), Oaxaca, and Chi-
persistent, reduced in older fascicles to (10-)15-20 apas; in Guatemala in most of the SW highlands;
mm, tight, resinous,weatheringblackishgrey. Leaves in Hondurason several isolated mountainsummits;
in fascicles of (3-)4-5(-6), most commonly 5, in reported from the extreme N of El Salvador, but
dense tufts toward the ends of upturnedbranches, not confirmed. Pinus hartwegii is the typical high-
persisting2(-3) years, straightor curved, not twisted, altitude pine of Mexico and Guatemala, where it
rigid, (6-)10-17(-22) cm X (1-)1.2-1.5 mm, with often forms extensive, monotypic pine forests up to
serrulatemargins, acute-pungent,dull light green or the treelineon high, isolated volcanos or summits of
[Link] all faces of leaves, in 6- mountain ranges. In Hondurasit is rare, of limited

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Systematic Treatment 119

i I
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~~~~ % .......

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..~ ~~ ~ ~ ~ ~~~, j. J~, ,,., .......,,.. 1

? M.i?l.?.
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Map I0. Pinus [Link]'e~?uii.

extent and usually found with Abies guatemalensis, Representative specimens examined. MEXICO. CHI-
HUAHUA: Cerro Mohinora, I Sep 1898, Nelson 4916 (A.
Cupressus lusitanica, Juniperus standleyi, Quercus US). GUERRERO: Cerro Zacatonal,3 km S of CerroTeote-
spp., Dendropanaxlempirianus,Dryimisgranadensis, pec, 15 Jan 1988, Aquino 328 (FHO, MEXU); 3 km W of
a ground cover of Ericaceae, Lycopodiaceae and CerroTeotepec, 27 Sep 1988, Aquino 514 (FHO, MEXU).
epiphytic Bromeliaceae in a cool cloud forest type HIDALGO: Locality not stated, 1837, Ehrenberg648 (MO);
near Pachuca, in El Chico National Park, 5 Nov 1992,
usually between 2700-2850 m on the highest moun- Gardner & Knees 5300 Nevado de Colima.
tain [Link] forests occur in Guatemalaand 15 Mar 1974, Styles 79 (U). JALISCO:
(FHO). MExico: Nevado de Toluca,
the southern states of Mexico, but here extensive May 1827, KarMinskis.n. (M); Valle de Mexico: 1800, er
pine forests predominate,in which P hartwegii in- herb. LindleY3164 (CGE); Salazar. 1900, Nelson 4 (A.
creasinglydominateswith rising altitude. Its altitudi- US); Ajusco Mts., 4 Dec 1903, Pringle 8789 (A. E. F,
nal range in Guatemala and Mexico is similar: GOET, K, M, MEXU, MICH, MO, NY,); Ajusco Mts.,
18-19 Aug 1906, Pringle 13782 (JE, MICH, S, US).
(2300-)2500-4000(-4300) m. At lower elevations it MICHOACAN: Cerro Patamban, n.d., Nelson 6563 (US);
is often mixed with P montezumae,with which it is Cerro Tancitaro, La Bolita, n.d., Madrigal 3269 (IBUG,
closely related (Carvajal& McVaugh, 1992; Styles, MEXU). MORELOS: Near Toro Sta., 14 Nov 1907, Pringle
1993), and with other pines depending on the geo- s.n. (US). NUEVOLE6N: Sierra Infernillo, 22 km SW of
Galeana, 16 Jun 1934, Mueller & Mueller 822 (MICH, NY,
graphical area. Soils are from both volcanic and P); Cerro Potosi, 26 Jul 1934, Mueller & Mueller 1261 (F,
granitic rock, of various depths but often poor in MICH, NY, P); La Joya, near Galeana, 13 Mar 1980,
[Link] there are considerable differ- Stead & Stvles 625 (FHO); 1 km N of Mimbres, 6 Nov
ences congruent with latitude/altitudedines, with 1984, Styles, Hoogh & Favela 19 (FHO. MEXU); La
Escondido. Cerro Viejo Zaragosa, 7 Nov 1984, Styles,
heavy frost and snow during several months and
Hoogh & Favela 24 (FHO, MEXU). OAXACA:Sierra
often high winds near the treelines of the high volca- Juarez.
along Hwy. 175 from Oaxaca to Tuxtepec, 23 Apr
nos in Central Mexico. Phenology: Time of pollen 1994, Farjon & Mejia 3/7 (E, FHO, HEH. K, MEXU, U);
dispersal varies mainly with altitude and is from Cuajimoloyas,2 Oct 1973, Hill 1846 (NY). PUEBLA: Lower
March(Critchfield, 1966b) to July. slopes of Pico de Orizaba W of Chlachicomula, 21 Sep

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120 Flora Neotropica

1957, Beaman 1789 (US); Albergue PiedraGrande, 14 Jul the protologue),it is original materialand a lectotype
1960, Beaman 3642 (US); Mt. Iztaccihuatl,on S side of was chosen from it (P). The Paris sheet bears only
mtn., along rd. between Paso de Cort6sand ParajeLa Joya,
23 Sep 1961, Beaman 4520 (US). VERACRUZ: Cofre de foliage, while in W only a mature cone has been
Perote, 28 Aug 1971, Dorantes 350 (FHO, MEXU, XAL); preserved.
Pico de Orizaba, (in Natl. Park), 25 Apr 1994. Farjon & There are evidently variations in cone colour in
Mejia 321 (E, FHO, HEH, K, MEXU, U); Cofre de Perote, several populations;a form with lighter brown (not
28 Mar 1963, Little 18974 (K, MEXU, US); Cofre de
purplish or blackish) cones has often been distin-
Perote, 1 km NW of the summit, 5 Sep 1986, Nee 32987
(FHO, MO, NY); Pico de Orizaba, Rose & Hay 5780 guished as P. rudis. Endlicher(1847) described that
(A, US). UNKNOWN LOCALITY: Locality not stated, n.d.. species without any reference to cone colour. In the
Hartweg s.n. (K); locality not stated, 1857, Roezi s.n. (FI); same publication, P hartwegii was described with
locality not stated, 1857, Roezl s.n. (FI). cones of a clear greyish brown. Martinez (1948)
GUATEMALA. CHIMALTENANGO: Volcin Acatenango.
16 Feb 1906, Kellerman5840 (US), 7 Feb 1907, Kellerman emphasized the colour differences and designated
6073 (US). GUATEMALA: Chinantla, May 1892, Donnell-
the name P hartwegii for the high-altitude pine
Smith 2633B (K). HUEHUETENANGO: Sierra los Cuchuma- with blackish cones and P rudis for those with
tanes, along rd. between Huehuetenangoand Concepci6n. brown cones. With Carvajal and McVaugh (1992)
27 Aug 1976, Boeke 99 (NY, US). [Link]: we agree that such distinctions mean little. A
Volcan Santa Maria, Sep 1876, Bernoulli & Cario 1089
(GOET); along rd. from Quetzaltenango,40 km toward morphological assessment (Matos, 1995) using 25
Guatemala City, I Oct 1983, McCarter & Hughes 41 characters of samples taken from two elevational
(FHO); "from mountains near Hacienda Chaucol," 1900, transects through populations in Mexico reported
Nelson 3729 (US). SACATEPEQUEZ: Volcan de Agua. 31 to include both taxa was unable to separate two
Jul 1959, Beaman 2960 (US), 15 Feb 1905, Kellermtan
4580 (US); Volcan de Agua, on crater rim, 16 Feb 1983, morphological entities. We also found that these
[Link] & Styles 5 (FHO), June 1892, Shannon charactersare far fromconsistent,and our conclusion
3622 (A, K, NY, S, US). SOLOLA:Mtn. ridge 1.5 km W is likewise to sink P rudis into synonymy of P
of Hwy. CA I, at Km 168, 18 Apr 1994, Farjon & Mejia hartwegii. Lower-altitude collections with brown
306 (E, FHO, HEH, K, MEXU, U); VolcanAtitlan, 16 Feb cones from W Mexico by J. N. Rose (Rose 2376,
1906, Kellermans.n. (US). TOTONICAPAN: Near Totonica-
2379, 3004), cited by Shaw (1909) underP. montezu-
pan at Chuapechec,6 Mar 1993, Higman. Padilla & Styles
4 (EAP, FHO, HEH, MO); 18 km from Nahuali toward mae var. rudis, have been treated under P dur-
San Crist6balTotonicapdn,I1 May 1979, Stead 303 (FHO). angensis by Carvajal and McVaugh (1992) and in
HONDURAS. CORTES:Ridge between Cerro Cantiles this volume.
and Cerro Jilinco, 20 km W of San Pedro Sula, in Cusuco
Natl. Park, 19 Mar 1993, Mejia 343 (EAP, FHO, HEH, Perry(1991), who recognizes P rutdisand P don-
MO, TEFH), 22 Mar 1993, D. Mejia 372 (EAP, FHO,
nell-smithii as full species close to P hartwegii,
HEH, MO, TEFH). LEMPIRA: El Moj6n Peak, 14 km SW presents a number of "distinguishing" character
of Gracias, in Celaque Natl. Park, 15 Apr 1994, Farjon & states for all three in a table, which at close inspec-
Mejia 300 (E, FHO, HEH, K, MEXU, U), 17 May 1992, tion appearto be overlappingor even similar. Pinus
Mejia 107 (EAP, FHO, HEH, MO, TEFH); Mt. El Cascal,
7.3 km NW of San Manuel Colohete, 17 Feb 1993, Mejia donnell-smlithii was described by Masters (1891)
313 (EAP, FHO, HEH, MO, TEFH). OLANCHO:13 km from materialcollected on Volcande Agua in Guate-
NW of Catacamas, 2 Jun 1992, Mejia 154 (EAP, FHO. mala, near the summit at ca. 3750 m (Dolnnell-Smith
HEH. MO, TEFH). 2182, holotype, US, isotype K). Like elsewhere,
it forms pure forest at this high altitude. Further
Uses. Pinus hartwegii has fairly dense but resinous collections and fieldworkby McCarter,Hughes, and
wood. It is exploited as a timber tree where stands Styles (FHO) revealed that the numberof leaves per
are extensive and accessible, but due to high altitude fascicle from that location is consistently 4-5, not
and consequent lack of infrastructure (access roads), 5-6(-7) as given by Perry (1991) for P donn1ell-
many forests remain virtually untouched today. smithii. There is only one species of Pinus present
Based on a collection by C. T. Hartweg ("Campa- around the crater rim of this stratovolcano;the cor-
nario, in the mountains of Angangueo, Hartweg"), rect name for it is P hartwegii. Leaf numberseems
Lindley (1839) described this species as a 4-leaved to decrease furthernorth and may be predominantly
pine with very stout branches. There is no original 3 in places in CentralMexico (Carvajal& McVaugh,
material at CGE (Lindley's herbarium); a specimen 1992), but 4-5 occurs throughoutthe range of P
at K ("ex herb. Gordon, Mexico, Hartweg") bears hartwegii.
no indication that Lindley saw it. In P and W, The numerous"new" species described by Roezl
material was found with Lindley's original labels; (1857) have here been lectotypifiedfor the first time
although no locality was stated (this is cited from by materialpreservedmainly in Fl and P. As Parla-

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SystematicTreatment 121

tore (1868) and Shaw (1909) alreadyobserved, none which remain with a few cone scales on the branch
of them are even worthtaxonomicconsideration,but when the cone has fallen. Immaturecones ovoid,
theirtrueidentitycould (in most cases) be established 15-20 X 10-12 mm, with small, spreading spines,
only throughtypification. More details about Roezl purplish, soon pinkish brown, sometimes glaucous,
and his catalogue of pines are presented under P maturingin two seasons. Maturecones ovoid-oblong
rmontezuniae. to obliquely ovoid when closed, sometimes ovoid-
attenuate,more or less asymmetrical,often curved
14. Pinus pseudostrobus Lindley, Edwards's Bot. at base, ovoid-oblong to broadlyovoid when opened,
X 6-13 cm. Seed scales ca. 140-190,
Reg. 25: 63. Aug 1839 [& Allg. Gartenzeitung7: then 7-16
325. 1839]. parting to release the seeds except the proximal
infertilescales, usually thick woody, oblong, straight
Tree, medium to large, height to 20-40(-45) m, or slightly curved, somewhatlargerand more curved
dbh to 80-100 cm. Trunkmonopodial,erect, terete, on one side of the cone, reddish brown to dark
the bole usually for %A-/2of the total height free of purplish brown, with light brown marks of seed
branches. Bark thick, scaly, with elongated plates wings on the adaxialside. Apophysisextremely vari-
and deep, longitudinalfissures, dark brown to grey- able, from nearly flat to elongated, more so on one
brown, on young trees and branchessmooth, reddish side of the cone and toward the base, transversely
brown or [Link] of first order long, keeled, taperingto an obtuse umbo (or umbo mucro-
spreading or ascending, the lower branches curved nate),rhombicor pentagonalin outline, uppermargin
down, often distinctly whorled; branches of higher angular,irregularlyundulate,or rounded,colour in
orders slender, spreading, curved down and then various hues of brown. Umbo dorsal, variable,from
upwards,or ascending. Shoots slender,smooth, with obtuse to prominent,3-15 mm long, 5-10 mm wide
(short)decurrentpulvini,glaucous or [Link]- at base, without a prickle or prickle deciduous, usu-
phylls 10-15 mm long, subulate-caudate,soon really darker than the apophysis. Seeds obliquely
curved, with ciliate margins, reddish brown to dark ovoid, slightly flattened,5-7 X 3-4.5 mm, ochra-
brown, falling shortly after the fascicles. Vegetative ceous to grey-brown, with or without dark spots.
buds ovoid-conical, acute, the terminal buds 15-20 Seed wings articulate,effective, held to the seed by
X 10-15 mm, the laterals smaller, not resinous; two claws, thinly covering part of the seed on one
the scales imbricate, with free, spreading apices, side, obliquely-ovate, with a straightside, 20-25 X
subulate-linear,with ciliate margins, reddish brown. 7-10 mm, yellowish brown, translucent,with darker
Fascicle sheaths (15-)20-30(-35) mm long, persis- tinge distally. Seedlings: The hypocotyl of this
tent, consisting of ca. 10 initially distinct, imbricate, species is consistently longer (mean 31.8 mm) than
subulatescales, lustrousred-brown,weatheringgrey- that of P montezumnae (mean 17.5 mm), according
brown. Leaves in fascicles of 5, rarely4 or 6, in lax to a study by Caballero (1967). Cotyledons (7-)8-
tufts at the ends of upturnedbranches,persisting2- 12(-13), but given as a mean of 7.2 by Caballero
3 years, slender, straight, spreading or drooping, (1967).
rarely more or less rigid, (18-)20-30(-35) cm x Distribution (Maps 4, 11, 12). Mexico: Sinaloa-
0.8-1.3 mm, with serrulate margins, acute, (light) Durangoborder,Nuevo Le6n, SE Coahuila, E Gua-
green or glaucous-green. Stomata on all faces of najuato(?),Jalisco, Michoacan,Mexico, DistritoFed-
leaves, in (2-)3-7 lines on the convex abaxial face eral, Morelos, Hidalgo, Puebla,Tlaxcala, W Central
and 2-4(-5) lines on each adaxial face. Leaf anat- Veracruz, Guerrero, Oaxaca, Chiapas; Guatemala
omly: Cross section triangular;hypodermis multi- (highlands);W Honduras;N El Salvador.A consider-
layered,not or scarcely intrudinginto the mesophyll; able east-west gap exists at the Isthmus of Tehuan-
resin ducts (2-)3-4(-6), the 3 marginalducts usually tepec, separatingthe Mesoamericanpopulationsfrom
present,medial, rarely 1-2 internalor external;stele the Mexican populations by ca. 250 km. The NW
more or less terete;the outer cell walls of the endo- (Sinaloa-Durango)and NE (Nuevo Le6n-Coahuila)
dermis thickened; vascular bundles 2, distinct, the disjunctionsare separatedfrom the main distribution
xylem strands connate. Pollen cones crowded near in central Mexico by similar distances.
the proximalend of a new shoot, numerous,spread-
ing, ovoid-oblong to cylindrical, 20-35 x 5-7 mm,
purplish or pinkish yellow, turning brown. Seed Key to the varieties of Pinus pseudostrobus
cones subterminal,solitary or in pairs, more rarely I. Seed scales with slightly to prominentlyraised or
in whorls of 3-4 on very short, stout peduncles elongated apophyses; umbo obtuse or elongated,

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122 Flora Neotropica

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.
...'
.

~
,'

i '
...

: .
.

. .
.

. . . . . .
!i!,

,
..-..... ......
,cm'':*....
? ..."'''"

.. ,, : :.......o..r .... .' ? ," :? .... 1~'

??~I.?
Pi ~ n ~~a u
uotoua.p
obu.. s s d

more or less continuing the line of the transverse Pinus orizabae G. Gordon, J. Hort. Soc. London 1: 79.
keel on the apophysis .................................................... 2 1846. Type. Mexico. Puebla: [Vic. of] Pico de Orizaba,
I. Seed scales with nearly flat apophyses;umbo acut- 1838?, Hartwegs.n. (original materialnot located) (lec-
ish, with a distinct angle to the transversekeel of totype, fig. in J. Hort. Soc. London 1: 238, here desig-
the apophysis .......................................................... nated).
........................ 14a2. var. pseudostrobusfo. protuberans Pinus protuberans Roezl, Cat. Grain. Conif. Mexic. 27.
2. Seed scales with slightly to prominentlyraised 1857. Type. Mexico. Mexico: Contreras, 1857, Roezl
but not elongated apophyses; umbo not elon- s.n. (lectotype, FI, here designated; isolectotype, BM).
gated, obtuse. ........................................ Pinus angulata Roezl, Cat. Grain. Conif. Mexic. 27.
............... 14al. var. pseudostrobusfo. pseudostrobus 1857; Pinus protuberans Roezl var. angulata (Roezl)
2. Seed scales with prominently raised and, at Carriere,Traite Gen. Conif., ed. 2, 2: 523. 1867. Type.
least in the lower half of one side of the cone, Mexico, n.d., Roezl s.n.-n.v.
elongated apophyses;umbo on these scales also Pinus regeliana Roezl, Cat. Grain. Conif. Mexic. 29.
elongated .................................... 14b. var. apulcensis 1857. Type. Mexico. Mexico: Volcan Iztaccihuatl, on
SW side of volcano, near Hacienda de Zavaleta, 1857,
14al. Pinus pseudostrobus Lindley var. pseu- Roezl s.n. (lectotype, Fl, here designated).
dostrobus fo. pseudostrobus. Type. Mexico. Mi- Pinus pseudostrobusLindley var. estevezii Martinez,An-
choacan: Near Angangueo, 1838, Hartweg s.n. (lec- ales Inst. Biol. Univ. Nac. Mexico 16: 188. 1945; Pinus
estevezii (Martinez) J. P. Perry, J. Arnold Arbor. 63:
totype, P, sheet with foliage, here designated; 187. 1982. Type. Mexico. Nuevo Le6n: Caion de Miel-
isolectotype, MO, cone scale and seeds). eras, Jun 1941, M. Mart'nez 3433 (holotype, MEXU;
Fig. 30-31 isotypes. A. NY).

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Systematic Treatment 123

i : :
~???i?~ - .. ......... .

MON.

--":t

II I-,

Map 12. Pinus pseudostrobusfo. protuberans(circles); [Link]. gordoniana (triangles).

PinuspseudostrobusLindley [Link], in various hues of brown. Umbo dorsal, obtuse, 3-6

Anales Inst. Biol. Univ. Nac. Mexico 16: 187. 1945. mm long, 4-6 mm wide at base, without a prickle or
Type. Mexico. Veracruz: "Cerca de Coatepec," Jun prickledeciduous, usually darkerthan the apophysis.
1940, M. Martinez3432 (holotype, MEXU).
Pinus pseudostrobus Lindley var. apulcensis (Lindley) Distribution and ecology. Mexico: In the Sina-
Martinez,Anales Inst. Biol. Univ. Nac. Mexico 16: 192. loa-Durango borderarea, in Nuevo Le6n, SE Coahu-
1945, non Shaw, 1909; Pinus pseudostrobus Lindley ila, E Guanajuato(?),Jalisco, Michoacan, Mexico,
subsp. apulcensis (Lindley) sensu Stead, excl. basion. & DistritoFederal,Morelos, Hidalgo, Puebla,Tlaxcala,
type, Bot. J. Linn. Soc. 89: 269. 1984. Type. Mexico. W CentralVeracruz,Guerrero,Oaxaca, and Chiapas.
Hidalgo: Apulco, Sep 1941, M. Martinez 3434 (holo-
type, MEXU).
Further south in Guatemala, W Honduras, and N
Pinus pseudostrobus Lindley forma megacarpa Loock, El Salvador. Of the varieties here recognized, var.
[pines Mexico Brit. Honduras] S. Afr. Dept. Forest. pseudostrobus is the most widely distributed, oc-
Bull. No. 35: 153. 1950. Type. Mexico. Michoacan:
Ciudad Hidalgo, Loock 10618 (holotype, PRF). curring throughoutthe entire range of the species.
Pinus nubicola J. P. Perry, J. Arnold Arbor. 68: 447. The other two varieties are often [Link] is
1987. Type. Guatemala. Guatemala:40 km E of San a widely distributedand in many places common or
Jose Pinula along rd. to Mataquescuintla,25 Feb 1979, abundantpine of montane to high montane habitat
Pern' GUA-32-79 (holotype, GH; isotypes, CHAP, E, in cold-temperateto warm-temperatezones. Its alti-
K, MEXU).
Pinus pseudostrobusLindley var. laubenfelsii Silba, Phy-
tudinal range is considerable but differs little along
tologia 68: 60. 1990. Type. Mexico. Sinaloa: In the
the entire 2500 km of its range: (850-)1900-3000
mtns. E of Mazatlan,Aug 1968, Laubenfels663 (holo- (-3250) m; in Mesoamerica and in N Mexico its
type, SYR-n.v.; isotype, NY). main distributionfalls within a 1900-2400 m zone.
Annual precipitation is also variable, but with a
Apophysis variable, from nearly flat to prominently minimumof ca. 800 mm; in Guatemalaand fondu-
raised, more so on one side of the cone and toward ras it can be >2000 mm. Throughoutits range it is
the base, transversely keeled, tapering to an obtuse an importantconstituent of mixed conifer pine and
umbo, rhombic or pentagonal in outline, upper mar- pine-oak forests, occasionally associated with Liq-
gin angular, irregularly undulate, or rounded, colour uidambaras on wet E slopes on the Gulf Coast side

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124 Flora Neotropica

L I I

?il /

?,, 30~~~~~.'4
?
~~~~~~~~~~~~-~~~~~~~~~.~~~~~~~~~~~~~$.,,,,,...,....??I
.
., . . -.
c~~~~~~,~ -, K-"~/ :i:
?,

FIG. 30. A-C, G, H. Pinus pseudostrobus var. apulcensis (Hignan. PcidillaS& StYles 26). D-F. P pseulostrobus var.
pseludostrobus(McCcarter572). A. Branchwith leaf fascicles and pollen cones. B. Leaf fascicle. C. Cross section of leaf.
D, G. Ovuliferous cones. E. Seed scale (three views). F, H. Seeds (two views). (Magnifications:A. B. D-H, XO.5; C.
x 40.)

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Systematic Treatment 125

FIG. 31. A. P pseudostrobusvar. pseudostrobus(Farjon & D. Mejit 329). Ovuliferous cone and scale (two views).
B-E. Pinus pseudostrobihs var. upulcensis (B, Hartwegs..t. lectotype: C. Higman. Pxdilla & Styles 26; D, Slell & Styles
446; E, E. [Link] 985. holotype of P. (xuricana). B. Closed cone. C. Cone. lateral(top) and proximal(bottom) views.
scale (two views). D. Cone, scale (three views). E. Upperpartof scale (two views). F. P pseclostrobus [Link]
fo. protuberans(Pringle 8788). Scale (two views). (Magnifications X 0.5.)

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126 Flora Neotropica

of the mountains of central and southern Mexico. In Tualor29478 (MO, NY); "nearJalapa,4-7000 ft.," 3 Mar
the many disturbed forests it may survive as scattered 1899. Pringle 8090 (A, F, GOET,JE, M, MEXU. MO, NY,
US, W); Municipio Perote. nearCoatepeque,22 Mar 1980.
groves or as individual trees, often with an understo- Stead & Styles 688 (FHO. P).
rey of, e.g., Gaultheria, Cassia, or, when associated (UATEMALA. CHIMALTENANGO: Tecpan.5 Feb 1972,
with fire, grasses and/or Pteridium aquilinum. In its Kemip138 (FHO). EL QUICHE:Chichicastenango, 7 Jul
driest habitat in central and northeastern Mexico 1951, Schwerdtfeger47 (M); Cunun, 4 Apr 1979, Stead &
it occurs with P cembroides, Juniperus flaccila, Styles 337 (ENCB, FHO). GUATEMALA: Along rd. between
San Jose Pinula and Mataquescuintla,ca. 20 km from San
Quercus, and an understorey with, e.g., Agave, Bud- Jose Pinula. 17 Apr 1994, Farjonl& Mejia 302, 303 (E,
dleja, Opuntia, and Salvia, probably mostly in sec- FHO, HEH, K, MEXU, U). HUEHUETENANGO: N of Chian-
ondary forest. Phenology: Pollen dispersal occurs in tla, CuchumatanesMtns., 4 Apr 1979, Stead & Styles 336
February-April (Critchfield, 1966b; and in speci- (FHO);Aguacatin, 4 Apr 1979, Stead & Styles 338 (FHO).
mens examined), dependent on location and altitude. JALAPA:La Lagunilla, II Sep 1976. Mittak 8299 (FHO.
M). SACATEPEQUEZ: Antigua, 3 Jul 1951, Schwerdtf,ser
43 (F). SOLOLA:Argueta, Sep 1870, Ber,noulli& Cario
Representative specimens examined. MEXICO. CHIA-
PAS:Las Rosas, 12 Mar 1993, Higman, Padilla & Styles 15 1081 (GOET); San Antonio Palopo, along shore of Lake
(EAP, FHO. HEH, MO); Arcotete. 15 Mar 1993. Higman. Atitlan, 4 Apr 1993, Higmlan,Padilla & Styles 66 (EAP,
Padilla & Styles 29 (EAP, FHO. HEH, MO). DURAN(;O: FHO, HEH, MO); San Andres Sementajab,along rd. from
Los Negros, 5 Apr 1991, Styles. Favela et al. 16 (FHO). Panajachelto Godinez, 19 Apr 1979, Stead & Styles 372
between Solola and Santa Lucia Utatlan ("Santa
GUERRERO: Chilpancingo,at Soxamanatlin. 25 Mar 1993, (FHO):
Higmnan,Padilla & Styles 61 (EAP. FHO. HEH, MO); Lucia Uspantan"),20 Apr 1979, Stead & Styles 376 (FHO).
Cerro Teotepec. 12 km NE of Puerto del Gallo. 19 Oct TOTONICAPAN:Near Totonicapin, 6 Mar 1993, Higtman,
1983, Soto N. et al. 5857 (MEXU, MO); Puntecillas. 10 Padilla & Styles 2 (EAP, FHO, HEH, MO); near jet. of
Nov 1965, May Nab AM237 (ENCB, INIF). HIDALG(O: rds. to San Francisco El Alto and Momostenango. 6 Mar
Along Hwy. 85 NE of Zimapin, near hamlet of Maguey 1993. Higm,tan. Padilla & Styles 5 (EAP. FHO, HEH. MO);
Verde, 28 Apr 1994, Farjon & Mejia 329 (E, FHO, HEH. San Crist6bal, 3 Apr 1993, Higman. Padilla & Styles 63
K. MEXU, U); (near)Apulco, 17 Mar 1980, Stead & Styles (EAP, FHO, HEH. MO).
641 (FHO. MEXU). JALISCO:G6mez Farias,21 Feb 1980, HONDURAS. COPAN:Dulce Nombre, May 1977,
Stead & Styles 506 (EAP, FHO). MEXIco: Ajusco Mtns.. Chaplin 367 (FHO). LA PAZ:On mtn. slope near small
1905, Lemt1on 241 (UC); Valle de Mexico, n.d., e. herb. hamlet of El Pinar, 13 Apr 1994, Faijon & Mejia 295 (E.
LindIle 3178 (CGE); Vallede Mexico, Eslava, 31 Jan 1899, FHO, HEH, K, MEXU, U); 12 km S of San Pedro Tutule
towardGuajicirro,MontanaPacayal,28 Jun 1982,
Pringle 8054 (A. BM, E, F, GOET. JE. K, M, MEXU, along rd.
MO). MICHOACAN: Ca. 29 km S of Patzcuaro,20 Nov HSglhes222 (FHO, K). LEMPIRA: Mt. Celaque. 20 Nov
1961, King & Soderstromi5168 (MEXU, MICH, MS, NY, 1974. Hiazlett2277 (HEH); Cerro de las Delicias, 5 Mar
US); along rd. from Ocampo to San Felipe, I kin from San 1988. McCarter572 (EAP, FHO); on ridge N of Arcagual
13 km SW of Gracias, Mt. Celaque National Park,
Felipe, I Mar 1980, Stead & Styles 586 (FHO); near camp,
8 km N main rd. to 22 Feb 1974. 15 May 1992, Mejia 106 (EAP, FHO, HEH, MO. TEFH).
Uruap:in, along Carapin,
Styles 3 (FHO); 10 km along rd. from Uruapanto Tancitiro. EL SALVADOR. SANTAANA:On the upper S-facing
23 Feb 1974. Styles 6 (FHO). NuEvo LE6N: Iturbide, 14 slopes of Cerro Montecristo, within the Montecristo Na-
Nov 1991. Frankis 159 (E); 5 km W of Iturbide,30 Mar tional [Link]. 16 km N of Metap6in,23 Feb 1989. Hulghes
1963, Little 18981 (IFGP. K, US); "Sierra Madre near 1262 (FHO. K. LAGU).
Monterrey,"9 Jul 1888, Pringle 1965 (E, MO. S. UC,
US. W); ParqueNac. Chipinque Monterrey,I) Mar 1980, 14a2. Pinus pseudostrobus Lindley var. pseu-
Stead & Styles 600 (FHO, MEXU); 8 km from Iturbide dostrobus forma protuberans Martinez, Pinos
along rd. from Linares to Iturbide. 12 Mar 1980. Stead &
Styles 607 (FHO, M, P), 12 Mar 1980, Stead & Styles 607h Mexic., ed. 2, 192, f. 154. 1948. Type. Mexico.
(FHO. P). OAXACA: At La Cumbre,along dirt rd. to Yurila, Distrito Federal: Eslava, Jun 1940, M. Martihez
3 km E of Hwy. 175, 24 Apr 1986, Gereau & Martin 1927 3431 (lectotype, MEXU, designated here).
(FHO, MO); Sierra Benito Juirez, 16 Nov 1984, Lepez et Fig. 31F
al. 248 (UC); roadside between Suchistepec and Miahuat-
ain.22 Mar 1895, Nelsonl 2521 (K, US). PUEBLA:Above Apophysis nearly flat, or slightly raised only on
Coxcatlin between Apala and top of Cerro Chichiltepec. one side of the cone and toward the base,
transversely
17 Jul 1961, C. E. Smithet al. 3850 (F. MEXU). SINAIOA:
keeled, rhombic or pentagonal in outline, upper mar-
17 km W of Sinaloa borderalong Hwy. 40. 22 Jul 1975.
Dunn et al. 21800 (NY); in mtns. E of [Link] gin angular, irregularly undulate, or rounded, colour
1968, Laithlfenfls 665 (NY). TAMAULIPAS:"Between Los in various hues of brown. Umbo dorsal, prominent,
Caballos and Agua de Vacas," May 1970. Gonsiles M. 4-6 mm long, 3-4 mm wide at base, without a
3058 (MEXU). VERACRUZ: Along Hwy. 140, 5 km SE of darker than the apophysis.
Las Vigas. 10 Mar 1983. Nee & 7Taylor25879 (F, FHO. prickle, usually
Distribution and ecology. Mexico. This form has
MICH, NY); Cerro de Tecomal, 5 km S of Los Altos
Veracruz. 12 Mar 1983, Nee & Tavlor 25936 (F, FHO, been first observed in the state of Mexico and in the
MEXU. MO, NY. XAL); Zongolica. 8 Feb 1984. Nee & Ajusco Mountains near Mexico City. The much

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Systematic Treatment 127

wider distribution given by Martinez (1948) and Padilla & Styles 28 (EAP. FHO. HEH. MO); near San
Crist6balde las Casas. 16 Oct 1960. Little 17963 (IFGP.
Perry(1991) includesS Jalisco, Michoacan,Morelos,
K, US); Mun. Tenejapa.20 Jan 1966, ShilornTon 650 (A,
Puebla, and one locality in Oaxaca. This taxon was
NY); between Comitanand San Crist6balde las Casas. ca.
not recognizedby Stead and Styles, and the (original) 7 km from Comitan. 26 Mar 1979, Stead & Styles 293
identificationsas this taxon, if present,on collections (FHO, US). GUERRERO:Along rd. from Chilpancingo to
seen by them were not recordedin the database. San Vicente, near Chilpancingo, 25 Mar 1993, Higman.
Padilla & St les 59 (EAP. FHO, HEH. MO); along rd.
Representative [Link]- from Xochipala to Filo de Caballo, 19 Apr 1985. Soto
Ico: Santa Rosa, Oct 1940, M. Martinez3505 (US). Mix-
N. & Aureoles C. 8261 (FHO); "Mpio. V. [Link]
ICO D.l.: Eslava, at base of Sierra de Ajusco at Eslava
Bernardino,La Laguna sobre terraceriaAzumbella," 27
19 Nov 1903,Pringle8788 (A, BM,F, GOET,
("Eslaba"), Feb 1982. Lott & Wendt133 (MEXU); between Filo de
K, MEXU,NY,P, S, U, UC);Eslava,n.d.,Pringle13262 Caballo and Colonia Eliodora de Castilla, 8 Feb 1981,
([Link]). Stead et al. 734 (FHO). HIDALGO: Apulco, n.d., (Hartweg?)
[Link]. Gordons.r. (FI). OAXACA: La Carbonera,18 Mar
1993, Higman, Padilla & Styles 36 (EAP. FHO, HEH.
14b. Pinus pseudostrobus Lindley var. apulcensis MO); NE of Guelatao. ca. 50 km NE of Oaxaca. 13 Oct
(Lindley) G. R. Shaw, [Pines Mexico] [Link] 1960, Little 17942 (IFGP,K. US); Mun. Ixtepeji, 19 km N
Arbor. 1: 19, t. 12, figs. 6-8. 1909. Figs. 30-31 of Hwy. 190 along Hwy. 175. 13 Feb 1981. Martin 320
(ENCB, MEXU); Teotitlandel Valle, Rancho Tablas, Nov
Pinusapulcensis Lindley,Edwards'sBot. Reg. 25: 63. 1940, M. Martinez 3435 (MEXU); Lachatao, Jan 1941.
Aug 1839[& Allg. Gartenzeitung 7: 325. [Link]. M. Martin. 4102 (US); "Miahuatlin" [= Santa Lucia
[Link]:RavinesnearApulco,1838,Hartweg Miahuatlan?].1895, Nelson 2539 (US); San Pablo Macuilti-
s.n. (lectotype,W, 2 seed cones with labels:P aptpl-
anguis. "cerco del panteon viejo." 3 May 1980. Pdrez
censis [Link] Lindley'shandwriting and phialwith C. 182, 183 (MEXU); distr. Coixtlahuaca,4 km SW of
seedsandscales,designated by Farjon(1995);isolecto- Magdalena Jicotlan. 17 May 1968. Rzedowski 25712
type,GH,seedconeandsomeleaves). (ENCB); Nochixtlin. along rd. to Jaltepec. ca. 3 km from
Pinuls pseudostrobus Lindley var. oaxaicana Martinez. Hwy. 190. 29 Jul 1970, C. E. Smith & Kitchen5034 (US);
Anales Inst. Biol. Univ. Nac. Mexico 16: 195, figs. ca. 13 km S of Miahuatlan,along rd. to Puerto Angel. 2
164-168. 1945, nom. inval.. sine lat. diagn.;Pinuis Feb 1980. Stead & Styles 423 (FHO, US). 2 Feb 1980.
[Link] Mirov, Madrofo 14: 145. 1958; Pinus pselu- Stead & Styles 434 (FHO. P), 2 Feb 1980. Stead & Styles
[Link](Mirov)S. G. Har-
cdostrohus 444-447. 449 (FHO); 10 km along rd. from Oaxaca to
rison,Taxon 14: 247. 1965. Type. [Link]: Tuxtepec. 26 Mar 1974. Styles 101 (FHO); 6 km SW of
"NearLa Parada,"18Aug 1894,Nelson985 (holotype,
Cuquila, along rd. from Tlaxiaco to Putla. 8 Jun 1985,
US; isotype,A). TorresC. 6693 (MEXU); 24 km NE of Ojite, nearTlaxiaco,
9 Aug 1985, TorresC. et al. 7144 (MEXU). PUEBLA: San
Apophysis variable,prominentlyraised and partly Pedro Chapulco, 18 km NE of Tehuacin. 20 Mar 1989,
elongated, especially on one side of the cone and Hughes 1322 (FHO. K. MEXU); 6 km S of San Salvador
toward the base, transversely keeled, rhombic or (El Seco). km 204. 28 Mar 1963. Little 18966 (IFGP,INIF.
pentagonalin outline, upper marginangular,irregu- K, US); Carpinteros.3 Jan 1974, Magana 900 (MEXU); I
larly undulate,or rounded,colour in various hues of km S of Carpinteros.3 Jan 1974, Magana 990 (MEXU):
brown. Umbodorsal,prominentand/orelongated, 5- Zacatepec, Jul 1986, Nee 33008 (NY). VERACRUZ: Cofre
de Perote, Jul 1866, Hahn s.n. (P); Saltillo de la Fragua.
15 mm long, 5-10 mm wide at base, without a Nov 1979, Klaus s.n. (WU).
prickle, usually darkerthan the apophysis. GUATEMALA. SAN MARCOS:Location given as
Distribution and ecology. Mexico: Mexico, Hi- 14?58'N.91?43'W, 7 Apr 1979, Stead & Styles 346 (FHO.
NY). 7 Apr 1979. Stead & Styles 349 (FHO). SOLOLA:
dalgo, Puebla, Tlaxcala, W central Veracruz,Guer- Mtn. ridge W of Hwy. CA 1. at Km 168, 1.5 km W of
rero, Oaxaca, and Chiapas. Also in the Guatemala
hwy.. 18 Apr 1994. Farjon & Mejia 307 (E. FHO. HEH.
highlands and N El [Link] K. MEXU. U).
to the SE part of the range of the species, but most EL SALVADOR. CHALATENANGO: Los Esesmiles, 6
common in Oaxaca, where it is perhapseven more May 1959, Allen & van Severe, 7324 (A. EAP, F, NY.
common than [Link] is often sympat- US); 4 km SSE of La Palma, 11 Jun 1970, Davidse & Pohl
2076A (FHO, MO).
ric and occupies much the same habitat but seems
absent from the driest locations. Uses. Pinus pseudlcstrobus is one of the most
[Link]- common and important "hard pines" in the southern
PAS:10 km E of Las Ollas, near hamlet of Cruzchen,E of half of Mexico and the highlands of Guatemala and
SanCrist6balde las Casas,20 Apr 1994,Farjon& Mejfia of Honduras. Its wood is light yellowish in
311 (E, FHO,HEH,K, MEXU,U); Las Rosas, 12 Mar parts
colour, lightweight (specific gravity -0.45), strong,
1993, Higman,Padilla& Styles 14 (EAP,FHO, HEH,
MO); Jitotol, 14 Mar 1993, Higman, Padilla & Styles 26 with long intervals of knot-free wood that is only
(EAP,FHO,[Link]);Arcotete.14 [Link],ntn, slightly resinous. Exploitation for timber is wide-

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128 Flora Neotropica

spread and has led to regional depletion of the best under P imontezumnae,the great majority of his names
stands. In Mexico, it is also used as a source of now have here been typified. One of them is P
resin. protulberansRoezl, first cited in synonymy with P
Pinus pseudostrobus was described by Lindley pseudostrobusby Parlatore(1868), who had indeed
(1839). based on materialcollected by Hartwegnear seen Roezl's materialat Florence (Fl).
Angangueo in Michoacan,Mexico. The only original Martinez(1948) recognized a numberof infraspe-
materialfound so far is a sheet in P without a cone, cific taxa under P [Link] of these is P
which refers to "Benth. pi. Hartw.p. 58. No. 443*" pseudostrobusf. [Link] characters
under P pseudostrobusLindl. and with a Paris label are describedabove. Stead and Styles (1984) synony-
identifying it as a Hartweg collection. Number 443 mized it under P pseudostrobus(subsp. psetdostro-
is not Hartweg'scollection numberfor this specimen. bus) as [Link]. prottuberais(Roezl) Mar-
The specimen was annotatedas "type collection" by tinez, but Martinez did not refer to Roezl. In an
E. L. Little in 1969. In BM a single cone with a earlier edition (Martinez, 1945: 184) he referred
label-"Pinus pseudostrobusLindl. Mexico, Angan- to Pringle 8788 (MEXU!) as "formaprotuberance
guco-Lambert's Sale [XXVI.55] 68"-is annotated Roezl," but this is Pringle's determination, not
as "type specimen" by E. L. Little. Apart from Roezl's type, which is somethingaltogetherdifferent.
the location, this specimen and its label profoundly We must simply consider this as a coincidence of
disagree with the protologue and belong with P homonymy, but at different ranks, and not read too
monteznumae; it is not accepted here as a type speci- much into the case.
men for Pifills [Link] Pinluspseudostrobus var. coatepecensis Martinez
original cone has still to be found; Stead and Styles is based on Martbiez 3432 from the vicinity of
(1984) reportedthat they did not see any type mate- Coatepec ("la region vecina a Coatepec") in Vera-
rial. The species was described as a 5-leaved pine, cruz, Mexico. It has a rather small, broad cone,
the leaves very slender and glaucous, "like those of which was taken as a distinction by Martinez,but in
the Wymouth pine." The cone scales are described all other charactersit is similar to P pseudostrobhts
as having a rhomboid,pyramidalapophysis("apex"), [Link] and Styles (1984) synony-
with an elevated, transverseline (i.e., keeled trans- mized it with their subsp. apulcensis, which was
versely); the length of the cone is given as ca. 10 defined by its relatively large cones, disregardingthe
cm. type but basing this opinion on their "population
As was customaryand understandableat the time, collection" gathered from 23 trees at Coatepec,
such a descriptionwas based on very scanty material, which was presumedto be the type locality (Stead.
often a single specimen. Little was known about the 1983).The inherentuncertaintiesof this methodhave
diversity of species in Mexico, and the material been discussed elsewhere (Farjon, 1995). Size and
was often comparedwith specimens of species from shape of cones are variable quantitativetraits in P
relatively high latitudes, which are much less vari- pseudostrobus, and the morphometric statistics in
able. As Stead and Styles (1984) pointed out, Lin- Stead (1983) fail to produce evidence for discrete
dley's description,in the light of today's knowledge, charactersbased on these measurements.
could apply to numerousspecies. However,any sub- Pinus pseludostrobuis var. estevezii Martinez,based
sequent application of such names can be correct on materialfrom Canionde las Mieleras, near Santa
only if it definitely includes its type as well as the Catarina,Nuevo Le6n, was also distinguished from
charactersgiven in the protologue. P pseutdostrobusvar. pseudostrobus by the cones,
There is an extensive synonymy for this species, which were describedto be larger,asymmetrical,and
in part discussed in Stead & Styles, 1984. Because "harder,"with elevated apophyses and a persistent
the treatment of P pseudostroblusLindley here- spine on the umbo. Differences in the leaves pertain
mainly due to the more recent typification of Lin- to a thicker hypodermis and an endodermis with
dley's two relevant species, P apulcensis (Farjon, thickeroutercell walls. Gaussen (1960) elevated this
1995) and P pseudostrobus--differs considerably variety to species but failed to cite the basionym,
from the one presented by Stead and Styles (1984), resulting in an invalidly published combination.
emphasis in the discussion will be on some taxa Perry (1982) elevated it once more as a distinct
treateddifferently,or published since 1984. species, based on his furthercollections, e.g., W of
Roezl (1857) published a numberof species that the town of Iturbide,Nuevo Le6n, where Zobel and
were tentatively synonymized, but not typified, in Cech (1957) and Stead (1983) also collected it.
recent treatments of Mexican pines. As discussed Although more or less confirming Martinez's mor-

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Systematic Treatment129

phological differences and adding a few from his Carvajal& McVaugh,1992), whose genetic diversity
own field observations(mainly habitand bark),Perry apparently indicates complex relationships among
based his main argumenton the chemical composi- [Link] con-
tion of the turpentine, with high levels of light tributed inconsistent classifications, which in turn
hydrocarbonsin P estevezii and low levels in P have been shown to use one of Lindley's binomials
pseudostrobis. Trees with morphological character contraryto its type. The treatmentpresentedhere is
states ascribed to this taxon can be found in many more conservative. New evidence, preferablybased
other places, far beyond the region Perry (1991) on moleculardata, could perhapsshow the existence
defined for it. None of these populations has been of distinct, taxonomicallymeaningfulcharacters.
found to be truly distinct.
Farjon (1995) published a full discussion of the 15. Pinus montezumae A. B. Lambert,[Link],
identity, including its typification, of P. [Link] ed. 8?, 1: 39, t. 22. 1832.
Lindley, and he showed that Shaw (1909) correctly
cited the illustrationin Loudon (1842) as represent- Tree, medium to large, height to 20-30 m, dbh to
ing Hartweg'soriginal [Link] original use of 100 cm. Truik monopodial,erect, terete, massive in
the name as to type, confirmed by Shaw, has been solitary trees, with persistent branches. Bark thick,
restored against the alternative of rejecting it in scaly, breakinginto numerous,relativelysmall, irreg-
favour of the usage by Martinez (1948) and Stead ular, slightly elongated plates divided by shallow
and Styles (1984), with a conserved type, because fissures, dark brown, weathering grey-black, on
there is no case of consistent use in a differentsense young trees and branches rough and scaly, reddish
since its publicationand the taxon is now recognized brown or greyish brown. Branches of first order
as a variety(subspecies)only. Pinlusoaxaccaa Mirov long, often massive, spreading wide; branches of
is a taxonomic synonym of P apulcensis. higher orders spreading to ascending in the upper
From Guatemala,ca. 40 km from San Jos6 Pinula partof the tree, assurgenttoward the ends, forming
E of GuatemalaCity, Perry(1987) describedanother a dense, usually broad,roundedcrown. Shoots thick,
new species, P nubicola. He also reported it from rigid, or more slender, very rough and scaly, with
other localities in Guatemala,Honduras,El Salvador, persistent,decurrentpulvini, brown,sometimesglau-
Chiapas, and a location in Veracruz. Virtually all cous. Cataplhlls ca. 15 mm long, subulate,recurved
collections are in Perry's personal herbarium(Hert- or reflexed,scarious, with erose-ciliatemargins,dark
ford, North Carolina,United States, but not seen on brown, weatheringgrey, persisting somewhat longer
a visit March 1994 by A. Farjon);only type speci- than the leaf fascicles. Vegetativebuds (broadly)
mens were distributedto several institutionalherb- ovoid, the terminalbuds 15-30 mm long, the lateral
aria. This makes evaluation rathercomplicated; the buds smaller,not resinous;the scales imbricate,with
only other accessible collection cited by Perry was free, spreading apex, subulate-linear,with ciliate
Mittak 8299, of which the FHO specimen consists margins, reddish brown to orange-brown,the ciliae
merely of a few seed scales. Collections from (the 3-5 mm long, [Link] sheaths (20-)
vicinity of) some of Perry's locations were made in 25-35 X 1.5-2.5 mm, persistent,consisting of ca. 10
1994. It was found that besides Perry'sform, several imbricate,orange-brownscales with ciliate margins.
others, both in regard to foliage and cones, were remainingtight, weatheringgrey to black, sometimes
easily gatheredwithin a few hundredyards of each resinous. Leaves in fascicles of (4-)5 (rarely3 or 6),
other, e.g., at the village of Las Trancas, Dept. La in dense tufts towardthe ends of upturnedbranches.
Paz, [Link] a larger geographical scale, the persisting 2-3 years, slender, flexible and drooping,
cone of Stead & Styles 641 (MEXU), part of a or lax to rigid and spreading,straight,(15-)20-35(-
"populationcollection" of P. pseudostrobus subsp. 40) cm X 0.8-1.3 mm, with serrulatemargins,acute
apulcensis sensu Stead & Styles at Apulco, Hidalgo, or pungent,(light) green to [Link]
Mexico, matches the cone of Perry GUA-32-79 on all faces of leaves, in 4-7 lines on the convex
(MEXU!), an isotype of P. nubicola J. P. Perry abaxial face and 3-6 lines on each adaxial face,
from [Link] (1987) reportedhaving seen weakly correlated with leaf width. Leaf anatoty:
intermediatesbetween P nubicola and P pseudostro- Triangularin cross section, with a convex abaxial
bus and assumed hybridization and back-crossing side; hypodermis sometimes thin and uni-layered,
were responsible. but mostly multi-layered,especially in the marginal
The species Pitnus pseudostrobus is a variable regions, with slight intrusions into the mesophyll:
and widely distributedtaxon (Stead & Styles, 1984; resin ducts (2-)4-5(-6). medial or occasionally 1-2

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130 Flora Neotropica

internal; stele terete; the outer cell walls of the ovoid-oblong. apophysis of scales flat, especially
endodermis thickened; vascular bundles 2, separate towardthe base of the cone, with a weak transverse
but close to each other, or the xylem strandssome- line.................................................. 15b. var. gordoniana
times connate. Pollen cones crowded near the proxi-
mal end of a new shoot, subtended by subulate to 15a. Pinus montezumae A. B. [Link]-
lanceolate bracts, cylindrical, 20-40 x 6-8 mm, umae. Type. Mexico. Veracruz:"InterCruz Blanca
initially pink or purplish, turning reddish brown. et Jalacingo, reg. frig. Nov. 28," Schiede & Deppe
Seed cones subterminal,solitary or in whorls of 3- s.n. (lectotype, BM, designated by McVaugh in
6, spreading or recurved on short, stout peduncles. Carvajal & McVaugh, 1992: 79; isolectotypes,
Immature cones ovoid, often curved, purplish to HAL, W). Fig. 32
purplish brown, maturing in two seasons. Mature Pinulf occidenltalis Kunth in Humboldt, Bonpland &
cones variable,from (broadly)ovoid or symmetrical Kunth, Nov. Gen. & Sp. PI. 2: 4. 1817, non Swartz.
ovoid-oblong to ovoid-attenuate,often curved, with 1788. Type. Mexico. Locality not stated, 1806. Bonp-
a more or less flattened base when opened, usually la(nd & Humboldt 3913 (holotype. P-Bonpl.; isotype,
widest near the base, 8-20 x 5-10 cm when open. P).
Pinu,. russelliana Lindley, Edwards's Bot. Reg. 25: 63.
Seed scales ca. 175-250, partingto release the seeds, Aug 1839 l& Allg. Gartenzeitung7: 325. 1839] Type.
the proximal,infertilescales remainingon the branch Mexico. Hidalgo:Near Mineraldel Monte. "lalong] the
when the cone falls, thin or thick woody, oblong, road from San Pedroto San Pablo," 1838, Hartwegs.n.
straightor curved, with thin margins,reddishbrown (lectotype, W, cones only, here designated).
Piltns monzttez:umae A. B. Lambert var. liidlevi J. C.
to purplishblack with a light bandunderthe apophy- Loudon. Encycl. Trees Shrubs 1004, figs. 1882-1883.
sis abaxially,(light) brown, with faint marksof seed 1842. Type. Mexico. Locality not stated, n.d., Hatrtweg
wings on the adaxial side. Apophysis flat or raised 31(?) (lectotype. W (ex Hort. Soc. London. 2 cones
especially on proximal scales, if raised with a trans- only), here designated).
Pinl.s teiinltgaensis Roezl. Cat. Grain. Conif. Mexic. 10.
verse keel, (transverse-)rhombicin outline, some-
1857; PiinuspseudostrobusG. Gordon & Glendinning,
times irregular,up to 20 mm wide, often rugose or Pinetum 237. 1858, non Lindley, 1839. Type. Mexico.
radially striate, in various shades of ochraceous to Mexico: Mt. Ajusco, 1857, Roezl s.n. (lectotype. Fl.
brown. Umbodorsal, depressed,flat or raised,obtuse here designated;isolectotypes, CGE. P).
or terminatingin a small prickle, weathering grey. Piinus grandis Roezl, Cat. [Link]. Mexic. 11. 1857.
non D. Don, 1832. Type. Mexico. Between the volcanos
Seeds obliquely ovoid, slightly flattened,5-7 x 4-
Popocatepetland Iztaccihuatl,at <9000 ft, 1857. Roe.l
5 mm, light brown,often with darkspots. Seed wings s.n. (lectotype, P. here designated).
articulate, effective, held to the seed by two thin Pirius imonstro.s Roezl, Cat. Grain. Conif. Mexic. II.
claws, covering the seed partlyon one side, obliquely 1857. Type. Mexico. Roezl s... photograph of seed
cone from Roezl's collection at P in G. R. Shaw's pine
oblong, 18-28 x 7-12 mm, light brown with dark collection; original material not located (lectotype, A.
stripes. Seedlings: Hypocotyl consistently longer here designated).
(29-37 mm) than in P pseudostrobus(14-28 mm); Pinus souliangeanaRoezl, Cat. Grain. Conif. Mexic. 11.
mean number of cotyledons 7.2 (Caballero, 1967), 1857. Type. Mexico. Mexico: Mt. lztaccihuatl,on SW
range 7-10 according to Mirov (1967). slope of volcano, 1857, RoezI s.n. (lectotype, Fl, here
Distribution (Maps 12, 13). Mexico: In Nuevo designated:isolectotype, P).
[Link],ltRoezl, Cat. [Link]. Mexic. II.
Le6n, SW Tamaulipas,Nayarit,S Zacatecas,Jalisco, 1857. Type. Mexico. Mexico: Mt. Iztaccihuatl,on SW
Michoacan,Mexico, Distrito Federal,Quer6taro,Hi- slope of volcano, 1857, Roe z s.n. (lectotype, Fl. here
dalgo, Morelos, Tlaxcala, Puebla, central Veracruz, designated).
Piirl.s hoothiIna Roezl, Cat. Grain. Conif. Mexic. 12.
Guerrero,Oaxaca, and Chiapas;in Guatemalain the 1857. Type. Mexico. Locality not stated, 1857, Roezl
highlanddepartments. s.n. (lectotype, Fl, here designated).
[Link] Roezl. Cat. Grain. Conif. Mexic. 12.
1857. Type. Mexico. Puebla:Mt. Iztaccihuatl,on W side
Key to the varieties of Pinus monteztumae of volcano, nearAmeca, 1857, Roezl s.. (lectotype. Fl.
I. Leaves spreading or drooping, 1-1.3 mm wide, here designated).
with (4-)5-7 lines of stomata on the abaxial face Pin,is elegans Roezl, Cat. [Link]. Mexic. 12. 1857.
and 4-6 lines on each adaxial face; mature seed Type. Mexico. Locality not stated, 1857, Roezl s.I.
cones variablein shape, apophysisof scales raised, (lectotype, Fl, here designated).
especially towards the base of the cone, with a Pinus runielianu Roezl, Cat. Grain. Conif. Mexic. 12.
transversekeel ............................. 15a. var. mtonte-uuneue 1857. Type. Mexico. Roezl s.n., original material not
1. Leaves slender, drooping, 0.8-1 mm wide, with located; photographof seed cone from Roezl's collec-
4-5 lines of stomata on the abaxial face and 3-4 tion at P in G. R. Shaw's pine collection (lectotype, A,
lines on each adaxial face; mature seed cones here designated).

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Systematic Treatment 131

I" -.- --4.-n

t-- ,, i f ??-- -? --? .. . . .

: i .....

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I i
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. ......r ...
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';'"" " .... ""
'.t~
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? ' i1'
-4~~~~~~~~~~~~~~~~~~~~
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.,+ ; ?
:...
.. -I .-.. . I

a'"rMap13 Pmnusmonezumne n.o..z u . .

Pinus spinosa Roezl, Cat. [Link]. Mexic. 12. 1857. Pinus paxtonii Roezl, Cat. [Link]. Mexic. 14. 1857.
Type. Mexico. Roezl s.n., original materialnot located. Type. Mexico. Mexico: Hacienda Tomacoco, 1857,
Pinus keteleerii Roezl, Cat. Grain. Conif. Mexic. 13. Roezl s.n. (lectotype, Fl, here designated).
1857. Type. Mexico. M6xico: Toluca, 1857, Roezl s.n. Pinus vanhouttei Roezl, Cat. Grain. Conif. Mexic. 14.
(lectotype, P, here designated; isolectotype, FI). 1857. Type. Mexico. Mexico: Mt. Ajusco. 1857, Roezl
Pinus san-rafaeliana Roezi, Cat. Grain. Conif. Mexic. s.n. (lectotype, FI, here designated; isolectotype, P).
13. 1857. Type. Mexico. Mexico: On the slope from Pinus haageana Roezl, Cat. Grain. Conif. Mexic. 15.
Acoculco to San Rafael, 1857, Roezl s.n. (lectotype, Fl, 1857; Pinus grandis Roezl var. haageana (Roezl) Car-
here designated). riere, Traite Gin. Conif., ed. 2, 2: 535. 1867. Type.
Pinus vangeertii RoezI, Cat. Grain. Conif. Mexic. 13. Mexico. Mexico: San Rafael, 1857, Roezl s.n. (lecto-
1857. Type. Mexico. Mexico: Hacienda Tomacoco, type, P, here designated; isolectotype, Fl).
1857, Roezl s.n. (lectotype, Fl, here designated). Pinus hendersonii Roezl, Cat. Grain. Conif. Mexic. 15.
Pinus antoineana Roezl, Cat. Grain. Conif. Mexic. 14. 1857; Pinus monstrosa Roezl var. hendersonii (Roezl)
1857. Type. Mexico. Mexico: Near HaciendaLa Valeta, Carribre,Trait6G6n. Conif., ed. 2, 2: 526. 1867. Type.
1857, Roezl s.n. (lectotype, P, here designated;isolecto- Mexico. Mexico: Rfo Frio, 1857, Roezl s.n. (lectotype,
type, FI). P, here designated; isolectotype, Fl).
Pinus doelleriana Roezl, Cat. Grain. Conif. Mexic. 14. Pinus jostii Roezl, Cat. Grain. Conif. Mexic. 15. 1857.
1857; Pinus grandis Roezl var. doelleriana (Roezl) Type. Mexico. Locality not stated, 1857, Roezl s.n.
Carritre,TraitdGen. Conif., ed. 2, 2: 532. 1867. Type. (lectotype, CGE (ex herb. Lindley 3177), here desig-
Mexico. Mexico: Toluca, 1857, Roezl s.n. (lectotype, P, nated; isolectotype, Fl).
here designated; isolectotype, Fl). Pinus thibaudiana Roezl, Cat. Grain. Conif. Mexic. 15.

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132 Flora Neotropica

/ I '. /

f' '"I". f.

iiIi .
iii .
'?I?
Viit 'I'Iru1.

'F.. .&

' ' "

~,. t., '
I'" ,i.,~ ' .
~ij~E~

? ':,.
A~~~~~~~~~~'.

FIG. 32. A-F, H. Pinns 01monttez.uma1evar. monte:.-unae(A-C, Stead & StYles536f D-F, H, Stead & StYles637). G. P
itionitezitmae var. gorloniianua (M. Martinez 3438). A. Branch with leaf fascicles and flushing terminal bud. B. Leaf
tfascicle.C. Cross-section of leaf. D. Ovuliferous cone. E. Seed scale (three views). F. Branchwith remnantcone scales.
G. Ovuliferous cone. H. Seed with articulatewing. (Magnifications:A, B. D-H. x0.5; C. x35.)

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SystematicTreatment 133

1857. Type. Mexico. Mexico: Mt. Popocatepetl,on N [Link]:Localitynot stated,1857.

side of volcano, 1857, Roezl s.n. (lectotype, Fl, here Roezls.n. (lectotype,Fl, heredesignated;
isolectotype.
designated;isolectotype, P). A).
Pinus Iullata Roezl, Cat. [Link]. Mexic. 16. 1857. Pinus rohanii Roezl, Cat. [Link]. Mexic. 25. 1857.
Type. Mexico. Mexico: Near village of San Mateo, Type. Mexico. Mexico: San Rafael. 1857, Roezl s.n.
1857, Roezi s.n. (lectotype, P, here designated;isolecto- (lectotype, P, here designated).
type, Fl). Pinus aztecaensis Roezl, Cat. Grain. Conif. Mexic. 26.
Pinus coarctata Roezl, Cat. Grain. Conif. Mexic. 16. 1857. Type. Mexico. Roezl s.n., original material not
1857. Type. Mexico. Mexico: Mt. Tzompoli, 1857, located;photograph of seed cone fromRoezl'scollec-
Roezl s.n. (lectotype, P, here designated). tionat P in G. R. Shaw'spinecollection(lectotype.A.
Pinus nesselrodianaRoezl, Cat. [Link]. Mexic. 16. heredesignated).
1857; Pinus monstrosaRoezl [Link](Roezl) Pinus zacatlanae Roezl, Cat. Grain. Conif. Mexic. 26.
Carriere,Trait6G6n. Conif., ed. 2, 2: 526. 1867. Type. [Link].n.. originalmaterialnot located.
Mexico. Between the volcanos Popocatepetland Iztac- Pinus exserta Roezl, Cat. [Link]. Mexic. 27. 1857;
cihuatl, 1857, Roezi s.n. (lectotype, P, here designated; Pinus protuberansRoezl var. exserta (Roezi) Carriere.
isolectotype, Fl). [Link].,ed. 2, 2: 524. [Link].
Pinus valida Roezl, Cat. Grain. Conif. Mexic. 16. 1857. Mexico:Guarda, [Link] Guerna-
Type. Mexico. Roezl s.n., original materialnot located; vaca, 1857,Roezls.n. (lectotype,FI, heredesignated;
photographof seed cone from Roezi's collection at P isolectotype,BM).
in G. R. Shaw's pine collection (lectotype, A, here Pinus cornea Roezl ex G. Gordon & Glendinning,Pine-
designated). tum, Suppl. 74. 1862. Type. Mexico. Localitynot stated.
Pinus thelemannii Roezl, Cat. Grain. Conif. Mexic. 16. 1857-1858, Roezi s.n. (lectotype, CGE (ex herb. Lin-
1857; Pinus grandis Roezl var. thelemannii (Roezl) dley 3187), here designated).
Carriere,Trait6G6n. Conif., ed. 2, 2: 532. 1867. Type. Pinus tenangaensis Roezl var. longifolia Carriere,Trait6
Mexico. Mexico: Mt. Tzompoli, on N side of mtn., Gen. Conif., ed. 2, 2: 520. 1867. Type. Roezl s.n..
1857, RoezI s.n. (lectotype, P, here designated). original materialnot located.
Pinus horizontalis Roezl, Cat. Grain. Conif. Mexic. 17. Pinus pescatorei Roezl ex Carriere,Trait6 Gen. Conif.,
1857. Type. Mexico. Roezi s.n., original material not ed. 2, 2: 555. 1867. Type. Mexico. Locality not stated.
located; photographof seed cone from Roezl's collec- 1858, Roezl s.n. (lectotype, P, here designated).
tion at P in G. R. Shaw's pine collection (lectotype, A, Pinus montezumnaeA. B. Lambert forma macrocarpa
here designated). Martinez, Pinos Mexic., ed. 2, 219. fig. 177. 1948.
Pinus rubescens Roezl, Cat. Grain. Conif. Mexic. 17. Type. Mexico. Morelos: Along rd. from Mexico City
1857. Type. Mexico. Mexico: San Augustin, 1857, to Cuernavaca, near Km 58, Jun 1940. M. Martinez
Roezi s.n. (lectotype, Fl, here designated; isolectotype, 3437 (lectotype, MEXU, designated by Carvajal &
P). McVaugh, 1992: 79; isolectotype, MICH).
Pinus carrierei Roezl, Cat. Grain. Conif. Mexic. 18. Pinus ntontezumaeA. B. Lambertvar. mezamnbrana Car-
1857; Pinus grandis Roezl var. carrierei (Roezl) Car- vajal, Phytologia59: 138. 1986 ("'m'ezenmiranuls").
Type.
riere, Trait6 G6n. Conif., ed. 2, 2: 533. 1867. Type. Mexico. Jalisco: La Media Luna, 25 Sep 1981. Carva-
Mexico. Hidalgo: Tulancingo, 1857, Roezl s.n. (lecto- jt(l & Luce 3411 (holotype, CREG).
type, P, here designated).
Pinus planchonii Roezl, Cat. Grain. Conif. Mexic. 18. Leaves in fascicles of (4-)5 (rarely 3 or 6), vari-
1857; Pinus grandis Roezl [Link] (Roezl) Car- able, slender and droopingor more rigid, 1-1.3 mm
riere, Trait6 Gen. Conif., ed. 2, 2: 534. 1867. Type. wide; stomata in (4-)5-7 lines on the abaxial face
Mexico. Hidalgo: Tulancingo, 1857, Roezl s.n. (lecto-
and in 4-6 lines on each adaxial face. Seed con)es
type, FI, here designated;isolectotype, P).
Pinus richardiana Roezl, Cat. Grain. Conif. Mexic. 18. variable in shape and size, broadly ovoid, ovoid-
1857; Pinus endlicheriana Roezl var. richardiana oblong or ovoid-attenuate,8-20 x 5-10 cm when
(Roezl) Carriere,Trait6Gen. Conif., ed. 2, 2: 542. 1867. open. Apophysis raised, especially toward the base
Type. Mexico. Mexico: Mt. Ajusco, 1857, Roezl s.n. of the cone, transverselykeeled, often rugose. Umbo
(lectotype, Fl, here designated;isolectotype, P).
Pinus decaisneana Roezl, Cat. Grain. Conif. Mexic. 19. raised, sometimes with a small prickle.
1857. Type. Mexico. Hidalgo: Pachuca, 1857, Roezl Note. Although Carvajal and McVaugh (1992)
s.n. (lectotype, Fl, here designated). reporteda maximumleaf width of 1.6 mm, but when
Pinus inflexa Roezl, Cat. [Link]. Mexic. 19. 1857; material associated with "doubtless montezumae"
Pinus rohustaRoezl var. inflexa(Roezl) Carriere,Traite
cones was measured,no width greaterthan 1.3 mm
Gen. Conif., ed. 2, 2: 541. 1867. Type. Mexico. Mexico:
Mt. Ajusco, 1857, Roelz s.n. (lectotype, Fl, here desig- could be found in the collections cited here. We
nated). assume that the thicker leaves have to be referredto
Pinus ortgiesiana Roezl, Cat. Grain. Conif. Mexic. 25. P devoniana instead.
1857; Pinus tenangaensisRoezl [Link] (Roez!) Distribution and ecology. Mexico: In Nuevo
Carriere,[Link]., ed. 2, 2: 520. 1867. Type.
Mexico. Mexico: San Rafael, 1857, Roezl s.n. (lecto- Le6n, SW Tamaulipas,Nayarit,S Zacatecas,Jalisco,
type, P, here designated). Michoacan,Mdxico, Distrito Federal,Quer6taro,Hi-
Pinus rinzii Roezl, Cat. Grain. Conif. Mexic. 25. 1857. dalgo, Morelos, Tlaxcala, Puebla, central Veracruz,

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134 Flora Neotropica

Guerrero, Oaxaca, and Chiapas. Guatemala: In the Pringle 10170 (F, GOET, M, MEXU, MICH, MO, NY,
highland provinces. Pinus montezumae occurs in W); I km N of 18 de Marzo, 29 Oct 1984, Rushforth497
a wide range of edaphic and climatic conditions (E): La Encantada,
Dec 1955,Zobel 10 (UC);Chipinque,
Dec 1955, Zobel 23 (UC). OAXACA: San Jer6nimoSosola,
throughout the mountainous regions of central and Oct 1940, M. Martinez3436 (MEXU); 4.5 km SW of La
southern Mexico and the Guatemalan highlands. In Uni6n, 9 Nov 1984, Manzanero M. 508 (MEXU, MO).
Nuevo Le6n it reaches from semi-arid pinyon-juniper PUEBLA:Miguel Hidalgo, 17 Mar 1983, Nee & Tavlor
woodland up to cold-temperate mixed conifer forest. 25982 (F. FHO, MEXU, MO, NY, XAL). TAMAULIPAS:
Acahuales, along Hwy. 70 between Ocampo and Tula, 29
Its altitudinal range is also great-(1200-)2000-
Apr 1994, Farjont& Mejia 332 (E, FHO, HEH, K, MEXU,
3200(-3500) m overall, with occurrences below 2000 U). VERACRUZ: 2 km SE of Palo Bendito, 26 Apr 1983,
m mainly in Nayarit, Nuevo Le6n, and Tamaulipas. Nee & Taylor26760 (F, FHO, INIF,MO, NY, XAL); "inter
It is most abundant and best developed in the temper- Cruz Blanca et Jalacingo,"Nov 1828, Schiede s.n. (HAL).
ate zone at ca. 2400-2800 m, with annual precipita-
UNKNOWN LOCALITY: Locality not stated, 1857, Roezl s.n.
(Fl); locality not stated, 1857, Roezl s.n. (FI).
tion exceeding 800 mm, most of it falling from June GUATEMALA. HUEHUETENANGO:5 km SW of San
through September. Throughout its wide geographi- Mateo Ixlatan, Sierra de los Cuchumatanes,6 Feb 1965,
cal range, P mrontezumae occurs together with many Breedlove 8643 (F). SOLOLA:Along rd. between Solola
other tree species, mainly in pine-oak and mixed pine and Santa Lucia Uspantan, 20 Apr 1979, Stead & Styles
377 (FHO). TOTONICAPAN: Alasca, 24 May 1979, Mittak
forests, but also with Abies religiosa, A. guatema- 8894, 8895 (M).
lensis, and Cupressus lusitanica. Many of these for-
ests have been depleted or even turned into small-
15b. Pinus montezumae A. B. Lambert var. gor-
scale farms with groups of scattered trees, among
doniana (K. T. Hartweg ex G. Gordon) Silba,
which P montezumiae is often present. Phenology:
Phytologia 68: 55. 1990. Fig. 32G
Time of pollen dispersal appears to be March-April
(material from Oaxaca and Veracruz). Pinusgordonicana K. T. Hartwegex G. Gordon,J. Hort.
Soc. London2: 79. [Link] San
Representative specimens examined. MEXICO. CHIA- Juan,nearTepic,n.d., Hartwegs.n., originalmaterial
PAS:10 km NE of Las Margaritas,17 Feb 1973, Breedlove not located(lectotype,J. [Link]. 2: 80, fig. (1847),
33414 (MEXU, MO, NY). DISTRITOFEDERAL:2 km S of designatedby Carvajal& McVaugh,1992:79).
San Miguel Topilejo. along rd. to Guernavaca, 27 Feb
1981, Mendez G. s.n. (MEXU). GUERRERO:Los Morros, Leaves in fascicles of (4-)5, slender and drooping,
between Xochipala and Filo de Caballos, 14 Feb 1982, 0.8-1 mm wide; stomata in 4-5 lines on the abaxial
Fonseca 193 (FHO, MEXU); Chilpancingo,25 Mar 1993, face and in 3-4 lines on each adaxial face. Seed
Higman, Padilla & Styles 57 (EAP, FHO, HEH, MO).
HIDALGO: Mineraldel Monte, "on the road from San Pablo cones ovoid-oblong or ovoid-attenuate,10-15 X 5-
to San Pedro, near Real del Monte," 1838, Hartweg s.n. 7 cm when open. Apophysis flat, especially toward
(W), Dec 1839, Schiede 915 (HAL); La Majada, 20 km the base of the cone, with a weak transverse line,
NE of Zimapin, 6 Nov 1979, HertcindezM. 3863 (MEXU); often smooth. Umhboflat or obtuse.
Ocotillos, Mar 1939, M. Martinez3625 (US); Apulco, 17
Mar 1980, Stead & Styles 637 (FHO, MEXU). MEXIco:
Distribution and ecology. Mexico: Scattered in
DesertaVieja. 1865, Bourgeau902 (P); Las Cruces, "Cerca the Sierra Madre del Sur N to Nayarit; also found
de Las Cruces, Valle de M6xico," 19 Dec 1950, Matuda along the "Eje Volcanico Transversal."Its distribu-
20905 (MEXU); HaciendaTomacoco 1857, Roezl s.n. (FI); tion as given here is probably incomplete, but it
Mt. Iztaccihuatl, 1857, Roezl s.n. (P); between San Felipe
and Jala, 60 km along rd. from Toluca to Zitacuaro,2 Mar appears to be sympatric with var. montezumae in
1980, Stead & Styles 592 (FHO, MEXU); 8 km S of Rio several locations. Its ecology is generally similar,
Frio. II Dec 1978, Vega A. 576 (MEXU). MICHOACAN: too, but it has not been found in the drier zones to
"[Link] 116.5 on road from Toluca to the northeast. Neither have we seen conclusive mate-
Morelia,"21 Jun 1988, Conzales M. 17002 (MEXU); Cerro rial from Mesoamericanlocations.
San Miguel, Pomocuaran,9 Apr 1979, Madrigal S. 3255
(MEXU, MS), 25 Feb 1980, Stead & Styles 536 (FHO, Representative specimens examined. MEXICO.
MEXU); along rd. from Ocampoto San Felipe, nearZitacu- GUERRERO: NE slope of Sierra Madredel Sur, near Chil-
aro, I Mar 1980, Stead & Styles 590 (FHO); Angangueo, pancingo, 24 Dec 1894, Nelson 2180 (A); Filo de Caballo,
1838, sin coll., ex herb.LambertXXVI55 (BM). MORELOS: 5 km SE of Fili de los Caballos,21 Apr 1985,SotoN. &
Toro, 14 Nov 1907, Pringle s.n. (MICH). NUEVOLE6N: Aureoles C. 8322 (FHO). JALISCO: Nevado de Colima, Jan
Locality not stated, 5 Jul 1935, Mueller 2125 (A. MEXU, 1942, M. Martitnez2212 (NY), Jul 1941, M. Martinez.3438
MICH, MO); 22 km SW of Pueblo Galeana,Sierra Madre (MEXU). MICHOACAN: On N slope of Mt. Patamban,Jan
Oriental, 18 May 1934, Mueller & Mueller 439 (MICH); 1903, E. W. Nelson 6599 (A). NAYARIT: Cerro San Juan.
10 km NE of Laguna de Sanchez, Sierra Madre Oriental, 1846, Hartweg s.n. (K, fragments), 1847, Hartwegs.n. (W,
3 May 1983, Marshall 83-26 (FHO); near Monterrey,23 cone). OAXACA:"Mountains W of Oaxaca," Sep 1894,
Jun 1888, Pringle 1964 (A, B, E, K, M, MO, US); "Sierra Nelson 1486 (A). PUEBLA:Buenavista. Huejotzingo. Jul
Madre above Monterrey,alt. 2500-4000 ft.," 2 Apr 1906. 1941, M. Martine:z2002 (A).

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Systematic Treatment 135

Uses. Pinus montezumaeis exploited as a timber nurserymenand estate owners of the time who were
tree throughoutits range. The timber is heavy and likely to buy "novelties") also guaranteed a wide
strong and in demand for construction purposes. dispersalof the [Link] publicationswere of
Locally, especially in the southernpart of its range, an ephemeral nature and are now difficult to find
this (and other) species, growing often near habita- in libraries. This perhaps influenced Schlechtendal
tions among fields as remnantsof more contiguous (1857-1858) to reprintall of Roezl's names in Lini-
forests, has its branches cut off due to the rural naea and translatetheir descriptionsinto Latin, with
population's incessant demand for firewood. Since only a slight doubt expressed as to the question of
pines rarelyregenerate(only one Mexican species is whether there really could be so many more new
known to do so) and none can be coppiced or pines in Mexico thanthose which Schiede and Deppe
pollarded indefinitely like many temperate broad- had previouslybroughtto his [Link]'s cata-
leaved trees, this use will be detrimentaland lead to logue of pines was furtherdistributedin France by
the demise of the species in these areas. Vilmorin-Andrieux& Cie. and also translatedinto
As Carvajal and McVaugh (1992) have pointed Germanfor distributionin [Link]
out, Pinus montezumae was described as a new much of his seeds and other materialcame to Eng-
species, not as an avowed substitute for the later land, we have not come across an English version
homonym P. occidentalis [Link] lectotypified of the catalogue.
Lambert'sname with a specimen (foliage) on a sheet Contemporaryhorticulturalauthors on conifers,
at BM, which is a part of collections made by such as Gordon (1862) and Carriere(1867), under-
C. J. W. Schiede and F. Deppe. At W, a few pine took to comment on them, but neitherhad a sound
cones have been preserved from the Endlicherher- taxonomic knowledge of the Mexican pines known
barium, among which is a cone with a small label at that time on which to base their judgements.
with the inscriptionby J. Lindley: "P montezumae Gordonsynonymized many of Roezl's names, often
Lamb."andon the reverse:"of this cone [Link] incorrectly,while Carrieretended to accept most of
drew his plate." It does not say who collected it, yet them, even validating several of Roezl's nomina
it perfectlyfits Lambert'sillustration,even as to size, nuda by providingthem with [Link]
and must be consideredtype [Link](1842) (1868), the more critical botanist, made short shrift
referredmaterialdistributedby the HorticulturalSo- of them by reducing the entire list (and more-see
ciety of London (Lindley?) to his P. montezumae below) of names to synonyms of seven already
var. lindleyi, which is a taxonomic synonym. known species (see also Shaw, 1909).
Roezl's catalogue of Mexican pines. Benedict What happened to the material on which these
Roezl (1824-1885) was the son of a Czech gardener, names were based and who saw it? Shaw (1909),
who emigratedto Mexico in 1854, where he main- who checked Gordon's reference (1862: 71, foot-
tained a nurseryand a plantation(Mabberley,1985). note), found that Roezl's collection was lost, which,
In June 1857 he issued a "Cataloguedes graines de for all we know, is the case. He believed that it
coniferes mexicains,qui se trouventchez B. Roezl & might be "impossible today to find a complete au-
Cie. a Napoles, pres Mexico," printed in Mexico thenticset." But threeherbariawere probabledeposi-
City. In it he named and described 82 new species tories, and these were checked in 1994 by A. Farjon:
of Pinus, giving brief characters,remarkson their I) Lindley's herbariumin CGE and/or herbariato
beauty or excellence, and often a statement as to which he distributedmaterialthroughthe Horticul-
where he had found them. Almost all of them are turalSociety of London;2) Parlatore'sherbariumin
from the Valle de M6xico and its surroundingmoun- Fl; and 3) the material Carriere had seen, which
tains, many from the same mountain or locality. could have gone to Paris (P). Little of it remains in
There is no attemptto compare them. Many of the Lindley's herbarium (CGE), but a few mounted
descriptionsare completely overlappingeach other. cones are there, bearing scant information as to
In 1858, a second catalogue was published in Paris, provenance. In Fl and P there are incomplete but
on plant seeds for sale at Roezl & Co., in which substantialsets, the most comprehensive of sheets
several new names appear,in addition to the 1857 and separate cones being the one in Fl. In P, the
names; but these, with one exception, received a material of branches and cones was wrapped up
summarydescriptionin groups, which rendersthem in coarse paper and stored with the carpological
invalidly published. collections. There is nothing in W, where Lindley
The obvious mercantile purpose (especially sug- had sent otherearly pine collections for the attention
gested in that many names are honouringEuropean of Endlicher.

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136 Flora Neotropica

In the register of accessions at FI (Botanical Mu- are two polymorphictaxa rangingfrom CentralMex-
seum) from 1841 to 1938, on page 33 underthe date ico to [Link] are also closely related to
14 October 1862, an entry is made for 435 specimens the high-altitudespecies P hartwegii, but more dis-
of conifers taken from E Parlatorefrom his trip to tinct from it. The morphologicalcharactersused to
London. Parlatore had bought Roezl's specimens delimit P. nmotezumae and P devoniana are largely
from John Standish (1814-1875), gardener of the quantitative and some are even subjective (Carva-
Duchess of Gloucester at Bagshot Park, Surrey,En- jal & McVaugh, 1992). The extremes of both taxa
gland, while visiting the HorticulturalSociety in are quite distinct,but many of the charactersused are
August 1862. Parlatorewrote his determinationsand much weaker than those preferredin a phylogenetic
"[Link]." referringto his treatmentin the Pro- species concept (e.g., Stevens, 1991; Thiele, 1993).
droinms (Parlatore, 1868) on the labels bearing A more or less congruentcombination of character
Roezl's names. There are sheets and separatecones states (e.g., narrowerleaves, shorterfascicle sheaths
in glass jars, but not all of these match up: there are and leaves, and smallercones) is then used to distin-
sheets without cones and vice versa. The sheets bear guish P [Link] not all collections combine
original labels with the names in a large hand, also these [Link] (1909) therefore united both
found on the wrappings in P. This is believed to be under P montezumae, but Martinez (1945, 1948)
Roezl's writing;the rest is probablyby variousother could not accept this and went off in the other
people who added "Mexico" (in Italian or French) directionby describinga rangeof varietiesand forms
and donnedepar M. Roezl" and/orsometimes a loca- underboth species. In the presenttreatment,lumping
tion. In the catalogue of the Museum botanique(P), of the two seems prematurewithout furtherdetailed
under "entrees sorties" 1833-1864, there is on page research,which should aim firstof all at findingtruly
78 an entry under 18 December 1858, No. 30: "M. fixed characters (Davis & Nixon, 1992), possibly
Roezl (de Mexico) Echantillonsde Pins (cones) du molecularones.
Mexique, 67" [= 67 specimens]. The labels have F- Pinus Jmonitezumaevar. gordonianca: Gordon
numbers(accession) and also "Roezlnumbers";these (1847) gave a validating description of P gordoni-
probably refer to one of his catalogues, but not to ani, a name coined by Hartweg for one of his
the one publishedin 1858 in [Link] is no proof collections from Mexico. No specimen had been
that Carriereever saw this material,but it may well found at K or elsewhere that could be interpreted
be that he did not annotate it. The wrappings had as original material; the illustration accompanying
become extremely brittle and have been removed, Gordon'sdescriptionwas designatedas the lectotype
retaining any pieces with text. In G. R. Shaw's (Carvajal& McVaugh, 1992). A single cone at W.
collection of pines at Harvard University (A) are from Lindley to Endlicher,labeled "P gordoniana,
kept an album and a box with photographsof the Hartweg," could be original material. The leaves
seed cones then at P and leaf fascicles, possibly also Gordon described as slender and very long, to 40
from P, which Shaw probably received after the cm; the cone illustrated is ovoid-attenuate, 12 cm
publicationof his "Pines of Mexico" in 1909. long (natural size) and has flat to slightly raised
Investigation of this material shows that Roezl apophyses and obtuse umbos. Shaw's (1909) treat-
issued many more names than the ones published in ment of P montezumae var. lindleyi Loudon, fol-
the 1857 catalogue. Parlatoredid extremely well in lowed by Martinez(1945, 1948), disagrees with what
synonymizing them, save for the distinctionbetween Loudon had in mind (see also Carvajal& McVaugh,
P montezumtaeand P gordoniana, which he did not 1992), which had ratherthick, rigid and short leaves,
make, and a few errors with P hartwegii which we and a cone with raisedapophyses. So P montezumnae
now define in a narrower sense. Only a few of var. lindlevi auct., non Loudon, is indeed in need of
Roezl's describednames are now left withouta trace another name. But this need not be a new one: P
of original material in institutional herbaria. For gordonlianaK. T. Hartwegex G. Gordon is available
some of these, the descriptionis unambiguousas to for it, as it agrees with both Shaw's and Martinez's
where to place it. If cited by Parlatore(1868), he concepts. We, indeed, believe it to be distinctenough
could have had a cone that subsequentlywent astray. to be recognized at the rank of variety, for which
These names have here been synonymized along Silba (1990) provided the correct nomenclature.
with those for which a lectotypehas been designated. Carvajaland McVaugh(1992), in synonymizing P
The rest, only a few names now, is listed with montezumaevar. mnezacibrana with P mnonitezumrae,
inserted cedis. suggested a possible hybrid origin of this taxon
Pi'mlt montezcinae and its close ally P devoniana involving P montezumaeand P. hartvegii. Including,

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Systematic Treatment 137

as they did, MartSnez 3438 (MEXU; Fig. 32G) in PittusmichoacanaMartinez,Anales Inst. Biol. Univ.
that concept would bring P. montezumnae var gor- Nac. Mexico 15: 1. [Link]:
doniana as we understand it into that putative orbit May 1940,M. Martinez3443 (holo-
Parangaricutiro.
type,MEXU).
of hybridization between two closely related, though Pitus michoacana Martinez var. corliuta [Link]-
ecologically distinct, taxa. ales Inst. Biol. Univ. Nac. Mexico 15: 1. 1944. Type.
Mexico. Jalisco: Tequila Iprobablyon Volcin de Te-
16. Pinus devoniana Lindley, Edwards's Bot. Reg. quila, S of Tequila].Oct 1941, M. Martinez3446 (holo-
25: 62. Aug 1839 [& Allg. Gartenzeitung 7: 324. type. MEXU).
Pinus michoacainaMartinezvar. quevedoi Martinez,An-
1839]. Type. Mexico. Hidalgo: Cerro Ocotillo, ales Inst. Biol. Univ. Nac. Mexico 15: 1. 1944. Type.
"found on the Ocotillo between Real del Monte Mexico. San Luis Potosi: El Tablon[= SierraEl
and Regla," 1838, Hartweg s.n. (lectotype, W, here Tablon?]. Oct 1940. M. Martinez 3448 (holotype.
MEXU).
designated). Fig. 33 Pintis michoacana Martinezvar. cornutta Martinezforma
Pinlts macrophyllcLindley, Edwards'sBot. Reg. 25: 63. naivaritana Martinez,Pinos Mexic., ed. 2, 266, fig. 215.
Aug 1839 [& Allg. Gartenzeitung7: 325. 18391;Pinus 1948. Type. Mexico. Nayarit:Compostela, May 1941.
iontezuniae A. B. Lambertvar. ntacrophylla(Lindley) M. Martitez 3447 (holotype, MEXU).
Parlatorein Candolle, Prodr. 16(2): 399. 1868. Type.
Pinls michoacanc Martinez forma procera Martinez.
Mexico. Hidalgo:CerroOcotillo, "foundon the Ocotillo PinosMexic.,ed. 2, 257, fig. 207. [Link].
between Real del Monte and Regla," 1838, Hartweg Jalisco: Tuxpan, Los Mazos. n.d.. M. Martinez 2433
s.n. (lectotype, W, here designated;isolectotype. K).
(lectotype,F. heredesignated).
Pinusfilifolia [Link]'sBot. Reg. 26: 61. 1839. Pionus nichoacana Martinez forma trumi,ida
Martinez.
Type. Guatemala."Murray,HabitatGuatemala"(CGE), PinosMexic.,ed. 2, 258, fig. 208. [Link].
"Santiago, Guatemala" (P), locality not stated (W), Oaxaca:San SebastianEtla ['Ixtlahuaca. Etla"on the
1838?, [H(artwegs.n.] ex herb. Lindlev3179 (lectotype, label,but the localitycited in Los PinosMexicanos,
CGE, here designated;isolectotypes, P, W). ed. 2 (1948), is Telixtlahuaca,
Oaxaca],Oct 1940,M.
Pinlls grenvilleaeG. Gordon,J. [Link]. London2: 77. Martinez3444 ([Link]).
1847. Type. Mexico. Nayarit:Cerro San Juan, Hartiveg
s.n. (holotype, W).
Pinus wincesterianaG. Gordon,J. Hort. Soc. London 2: Tree, medium or sometimes large, height to 20-
158. 1847. Type. Mexico. Nayarit:"Cerrode San Juan 30 m, dbh to 80-100 cm. Trunkmonopodial,usually
(or Saddle Mountain)near Tepic," 1845, Hartweg s.n. erect, slender or massive in old solitary trees. Bark
(holotype, W. "Pinus Winchestericna";isotype, Fl).
Pinuls michoacaensis Roezl, Cat. Grain. Conif. Mexic. thick, scaly, very rough, with elongated plates di-
26. 1857. Type. Mexico. Roezl s.n., original material vided by deep longitudinal black fissures, reddish
not located; photograph of seed cone from Roezl's brown to dark brown, on young trees rough, soon
collection at P in G. R. Shaw's pine collection (lecto- [Link] of first order long, spreadingand
type, A, here designated).
Pinus magnifica Roezl, Cat. Grain. Conif. Mexic. 28. assurging,or ascendingnearthe top, often persistent;
1857. Type. Mexico. Michoacan:In mtns. near Morelia, branchesof higher orders sparse, spreadingand as-
1857, Roezl s.n. (lectotype, FI, here designated). cending, the highest order branchesassurging,form-
Pinus ocanmpiiRoezl, Cat. [Link]. Mexic. 28. 1857. ing an open, broad pyramidal or domed crown.
Type. Mexico. Michoacan:"On hacienda of Melchior Shlootsvery thick (15-20 mm), rigid, curved, very
Ocampo, near Morelia,"1857, Roez s.n. (lectotype, Fl,
here designated;isolectotypes. A, P). rough and scaly, with persistent, decurrentpulvini.
Pijnus zitacuarensisRoezl, Cat. Grain. Conif. Mexic. 29. Cataphylls large, up to 20-25 mm long, subulate,
[Link]. Mexico. Michoacan:Zitacuaro.1857. Roezl scarious,recurvedor reflexed,darkbrownto blackish
s.n. (lectotype, Fl, here designated). grey, with erose-ciliate margins, persisting ca. one
Pinus nec-plus-ultra Roezl, Cat. Grain. PI. Mexic. 6. season longer than the leaf fascicles. Vegetativebuds
1858. Type. Mexico. No locality given, 1858, Roezl s.n.
(lectotype, Fl, here designated). large, terminalbuds 20-40 mm long, lateralssmaller,
Pinus zitccuarensis Roezl var. nitidacRoezl ex Carriere, ovoid-acute, not resinous; the scales imbricate but
Trait6Gen. Conif.. ed. 2. 2: 554. 1867. Type. Mexico. spreadingapically, subulate-linear,orange-brownto
Location unknown, 1857. Roezl s.n., original material reddishbrown, with long-ciliate (3-5 mm) margins,
not located.
Pinus verschaffeltiiRoezl ex Carriere.Trait6Gen. Conif., giving the buds a villose appearance. Fascicle
ed. 2, 2: 555. 1867. Type. Mexico. Location unknown, sheaths very long, up to 30-40 mm, with ca. 10
1857, Roezl s.n. (lectotype, P, here designated;isolecto- imbricate,orange-brownscales with whitish ciliate
type, Fl). margins, persistent, on mature fascicles (20-)25-
Pinus ptawlowskiana Roezl ex Carriere, Trait6 Gen.
35(-40) x 2-3 mm, tightly enclosing the fascicle,
Conif., ed. 2, 2: 556. 1867. Type. Mexico. Roezl s.n.,
original materialnot located; photographof seed cone turningdark brown or blackish, often very resinous.
from Roezl's collection at P in G. R. Shaw's pine Leaves in fascicles of 5, rarely 4 or 6, in dense,
collection (lectotype, A, here designated). wide-spreading tufts toward the ends of upturned

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138 Flora Neotropica

'
i

FIG. 33. Pinus (Ievolnia(li (A-C. Higman. Padilla & Styles 62; D. G. Stead & Styles 28IA; E, F. Hughes 1349). A.
Shoot with leaf fascicles, cataphylls. bud and immatureovuliferous cone. B. Fascicle sheath. C. Cross section of leaf. D,
E. Maturecones. F. Basal seed scales of cone attached to branch.G. Seed scale (three views). H. Seed with articulate
wing. (Magnifications:
win~. (Magnifications:A. B. D-H.
A, B. D-H, xxO.5: C. x
O.5:C. x30.)
30.)

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Systematic Treatment 139

branches, persisting 2-3 years, rigid, straight, or Guerrero,Oaxaca, and Chiapas. Guatemala:In the
flexible and drooping, (17-)25-40(-45) cm X 1.1- southernhighlands. In both countries it is a constit-
1.6 mm, with serrulate margins, acute-pungent, uent of relatively open, often secondary pine-oak
bright,lustrous(dark)green. Stomataon all faces of forests; secondly of secondarymixed pine forest, or
leaves, in (4-?)5-9(-10) lines on the abaxial face with Pinus oocarpa invading burnedmountainsides.
and 3-6 lines on each adaxial face. Leaf anatomy: It is well-adaptedto withstand fires by its juvenile
Cross section triangularor transverse-triangular,with "grassstage" similar to that of P palustris of the SE
flat or convex abaxial side; cuticula and epidermis United States. The pines most commonly associated
thick; hypodermismulti-layered,with (irregular)in- with it vary somewhat from north to south and
trusions into the mesophyll; resin ducts large, 3(-4), include P. oocarpa, P. montezumae, P. pseudostrobus,
medial, near the edges, sometimes 1-2 (smaller) P. maximinoi, and, less frequently, P. cembroides at
ducts internal;stele terete or weakly trigonous;outer lower altitudes and P hartrwegiiat higher altitudes.
walls of endodermisnot thickened;vascularbundles Common are also Quercus and Liquidambar and,
2, distinct but approximate,often partly surrounded in the understorey,Calliandra, Leucaena, Acacia,
by sclerenchym layers. Pollen cones crowded near Dodonaea, Gaultheria, and Mimosa. Its altitudinal
the proximalend of a new shoot in short,often dense rangeis (700-)900-2500(-3000) m. Pinus devoniana
clusters, subtended by long, subulate bracts with grows on a variety of soils, often of volcanic origin.
ciliate margins, cylindrical, 20-40 mm long, pink- The climate is warm-temperateto subtropical,with
purplish,turninglight brown. Microsporophyllspel- annualprecipitation1000-1500 mm and a dry season
tate-cordate,with a radially striate abaxial surface from November to May. Phenology:Time of pollen
and hyaline-erose [Link] cones subterminal, dispersal not recorded.
solitaryor in pairsor whorls of 3-4. Immaturecones
initially often erect, after pollination recurving on [Link]-
Representative
PAS:Comitan. 11 Mar 1993. Higman. Padilla & Styles 10
thick, bracteate peduncles, ovoid-oblong, purplish (EAP,FHO,[Link]);Bochil,14 Mar1993,Higilan,
brown,maturingin two years. Maturecones on short, Padilla & Styles 23 ([Link]. MO, ESNAC): Motozintla.
persistent peduncles, leaving a few scales on the 16 May 1945. Matuda 5534 (MEXU. MO, NY); 7 km
branch when falling, variable in size and shape, from Comitanat FranciscoSarabia, I Apr 1974. Styles 121
(FHO); 4 km E of Amatenango del Valle, along Hwy.
typically ovoid-oblong, with an oblique base, or 190 to Comitan, 27 May 1985, Villaseior & Thomais829
curvate to cornute (often by insect-damage), 15-35 (MEXU, NY). GUERRERO: Omiltemi, Plan de Potrerillos.
x 8-15 cm when open. Seed scales ca. 175-225, 24 Apr 1985, CasteloN. 446 (MEXU);along rd. to Chilpan-
partingwhen the seeds are mature,spreadingwide, cingo, 5 km fromChilapa.24 Mar 1993, Higmatn,Padilla &
Styles 54 (EAP. FHO, MO, ESNAC); SierraAlquitran,10
straight or slightly recurved, thick woody, rigid or Jun 1968, Kruse 1778 (MEXU); 2 km from Los Morros
flexible with some force, margins thin, light brown
along rd. to Filo de Caballos, 5 Mar 1984, Lorea 2986
or purplishblack on the abaxial side, (light) brown (FHO, MEXU); near Taxco. Jun 1940. M. Martitnez3445
with lighter marks of seed wings on the adaxial (MEXU); W of Petlacala.4 Jan 1938, Me.ria9083 (F. K.
side. Apophysis mostly raised, transversely keeled, MO, NY, S, UC. US); 1I km W of Chilpancingo, 13 Nov
rhombic in outline, with irregularsides, up to 25 1973, Quero et al. 2264 (FHO, MEXU, UC); Filo de
Caballo, 20 Apr 1985. Soto N. & Aureoles C. 8281 (FHO).
mm wide, often radially striated, in various shades HIDALGO: 20 km NE of Pachuca along rd. to Huasca at
of brown. Umbo dorsal, raised, flat or depressed,up El Octal, Gardner& Knees 5259A (E). JALISCO:Cerro La
to 7 mm wide, terminatingin a small, usuallydecidu- Bufa, 2 Feb 1927, Mex'.r 1627 (A, BM, F. MICH. NY.
ous prickle, grey-brown. Seeds obliquely broad UC); Talpa de Allende, on rd. to Cuale, 8 Nov 1984.
Rusliforth609 (E); Sierra del Tigre, 10 km from Ciudad
ovoid, flattened,8-10 X 5-7 mm, light brown,often Guzmfin, 10 Mar 1974, Styles 51 (FHO). MICHOACAN:
with dark spots. Seed wings articulatebut effective, CerroAzul, nearMorelia,28 Mar 1910,Arsene5477 (MO);
held to the seed in two oblique claws, obliquely Dos Aguas, 19 Feb 1980, Klaus s.n. (WU); Los Conejos,
ovate to oblong, with a straightside, 25-35 X 10- 10 km SW of Uruapan, 1900, Little 17837 (K. MS, US);
17 km along rd. from Ciudad Hidalgo to Mil Cumbres.9
15 mm, light brown with darker stripes. Seedling:
Feb 1980, Stead & Styles 466 (FHO). MORELOS: Cuerna-
Numberof cotyledons not observed;many seedlings vaca, Nov 1979, Klaus s.n. (WU). NAYARIT: Cerro Los
develop a "grass stage" with delayed apical growth Lobos, NE of Santa Mariadel Oro, 19 Aug 1959, Feddema
of the stem. 664 (MEXU. MICH);8 km W of Chapalilla, I Nov 1984,
Distribution and ecology (Map 14). Mexico: In Marshall 84F-104 (FHO); loc. unknown, Feb 1941. M.
Matrtinez2413 (F, GH, US); El Tortuguero,30 Oct 1979,
Nayarit,Jalisco, Zacatecas,Aguascalientes,San Luis Salazar J. I (MEXU); 6 km NW of Xalisco. 30 Mar 1987.
Potosi, Queretaro,Hidalgo,Michoacan,Mexico, Dis- Tellez V 10121 (MEXU). OAXACA: Dirt rd. to San Lucas.
trito Federal, Morelos, Tlaxcala, Puebla, Veracruz, near Hwy. 190, ca. 30 km from Oaxaca. 23 Apr 1994.

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140 Flora Neotropica

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~~~~~~~~~~~~~~~~~~I'
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:

dc]rlnn ....~~~~~~~~~Mp1. Piu

Farjon & Mejia 314 (E, FHO. HEH, K, MEXU, U); La Sacatepeque. 22 Mar 1979, Stead & Styles 278 (FHO).
Soledad Tectitlan, 26 Apr 1986, Gereau & Martin 2000 HUEHUETENANGO: Along Hwy. CA I between Huehueten-
(FHO, MO); Presa Benito Juarez,near Santiago Lachiguiri, ango and Colotenango,above the Rio Cuilco, 19 Apr 1994,
ca. 55 km NW of Tehuantepec,29 Mar 1989, Hughes 1349 Farjon & Mejia 309 (E, FHO, K, MEXU, U, HEH);
(FHO. K. MEXU); Portillo de Nejapa, 17 km NW of El Malacatacingo,3 Apr 1993, Higman, Padilla & Styles 62
Coyul. 8 Nov 1993, Hughes 1773 (FHO, K, MEXU); (EAP, ESNAC, FHO, MO); near village of Gracias a Dios
Ayoquezco, 29 Oct 1979, Ortega C. s.n. (MEXU); 5 km E W of Finca la Trinidad, ca. 25 km N of Nenton, 2 Apr
of San PedroCoatlan, 18 Mar 1988, TorresC. et al. I 1850 1989, Hughes 1357 (FHO, K); 13 km from Aguacatan. 5
(MEXU). PUEBLA:Near Calchicomula, Jul 1901, Rose & Apr 1979, Stead & Styles 339 (FHO). QUEZALTENANGO:
Hay 5781 (A); ZACATECAS: Nochistlan, n.d., M. Martiinez Sacatepequez, Mar 1892, Donnell-Smith 2632 (K, US);
2429 (A). WITHOUT LOCALITY: Specimen labeled "Pinus along Hwy. CA I. near Chiquibal, 18 Apr 1994, Farjon &
filifolia" presentedto Parlatore,1862, ex herb. Balfour s.n. Mejia 308 (E, FHO, HEH, K, MEXU, U); 2 km SW of
(FI); specimen labeled "Pinus Winchesteriana,Marquisof village Alotenango, on track to Volcin de Fuego, 9 Mar
Winchesterspecimen of seeds 1845," ex herb. Gordon s.n. 1984, McCarter& Styles 184 (FHO); Santa MariaCauque,
(K, det. not certain);specimen labeled "Pinus Devoniana," 7 Feb 1978, Stead & Styles 142 (FHO). SOLOLA:Volcin
1857, Roezl s.n. (Fl). SantaClara.5 Jun 1942, Stevermark47038 (F). TOTONICA-
GUATEMALA. BAJAVERAPAZ: Cubulco, Sep 1870, PAN: Casa Blanca, 19 km along rd. from Totonicapan to
Bernoulli & Cario 1085 (GOET). CHIMALTENANGO: Lo- Santa Cruz del Quiche, 12 May 1979. Stead 310 (FHO).
cality not stated, 3 Jul 1951, Schwerdtfeger44 (F); Zara-
goza, along Pan American Hwy. between Patizicia and
Uses. Pinus devoniana is a common tree through-
Chimaltenango,22 Apr 1979, Stead & Styles 381 (FHO).
GUATEMALA: Along rd. from San Jose Ocafia to San Juan out the mountainous parts of southern Mexico and

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Systematic Treatment 141

Guatemala, often encountered in open, degraded leaves (45 cm). Like the othermaterial,it was distrib-
pine-oak or pine forest. As it is not a very tall, uted throughthe HorticulturalSociety of London,of
straight-boledtree and has brancheslow on the trunk, which Lindley was secretary, to estate owners in
it is not often selected as a timber tree. On the GreatBritain,and also to Endlicherin Vienna,where
other hand, it is locally used and, like other pines, good cone specimens (W) have "miraculously"sur-
increasinglyfor firewoodby the growing ruralpopu- vived the ravages of the Second WorldWar.
lation. Gordon's (1847) names P grenvilleae and P.
This species-in its more extreme forms easily wincesterianawere probablybased on a single gath-
distinguishable from its relatives (in particularP. ering made by Hartwegon Cerro de San Juan near
,nontezuinae)by its thick, shaggy branches,its very Tepic in the state of Nayarit. Because no original
long and stiff, lustrous green leaves with very long, material has been found in Gordon's herbariumat
dark, resinous fascicle sheaths, and the very large, K, Carvajaland McVaugh(1992: 51) designatedthe
thick woody cones-occurs in a bewildering array relevant published illustrationsof P. grenvillae and
of polymorphismsespecially of cone forms. Also, P. wircesteriana as the lectotypes of these species.
intermediatesoccur between this taxon and P. mon- However, as with Lindley's taxa, there is original
tezumae, with charactersof either species shared in material of these in W and Fl (the seeds of "P.
populations of one or the other. Carvajal and Winchesteriana"in K cannotbe identifiedunambigu-
McVaugh (1992) compared this situation with the ously) which must be considered the holotypes (W)
species complex which Stead and Styles (1984) and isotype (FI), of Gordon's taxa. They are to
called the "pseudostrobus"group, and spoke of the supersedethe lectotypes (illustrations)designatedby
"montezumae"group, in which P hartwegii is to be Carvajaland McVaugh(Art. 9.13 in Greuteret al.,
included. Shaw (1909) considered them as a single 1994).
species, P. montezumae, with four varieties: var. It has, in addition, been possible to retracemany
montezumae,var. lindlevi, var. rudis, and var. hart- of Roezl's "new species" to original materialin A,
wegii, but in his monograph(Shaw, 1914) he dropped CGE, and especially Fl and P (see for discussion
even these. He considered the wide range in size under P montezumae),and they have accordingly
of both leaves and cones of no taxonomic value and been lectotypifiedhere. None of them deserves rec-
believed that they probably increased with warm, ognition as distinct taxa, as Parlatore (1868) had
subtropicalclimatic conditions furthersouth. Num- alreadyconcluded.
bers of leaves per fascicle increase in a similar Martinez (1945, 1948), reacting to Shaw's very
fashion, according to Shaw, and he believed all inclusive treatmentof P. montezumae,set up a "grupo
these characters to be "accurately correlated with Michoacana"with a new species: P [Link]
altitude and exposure." His lumping (Shaw, 1914), considered this species likely to correspond with
however, did not prevent him from recommending Lindley's species P devoniana, P russelliana, and
monographicstudy of the "montezumae"group, an P. macrophylla,but probablyfelt he could not use
undertakingwe still await today (Styles, 1993: 409). any of these names as his only source of information
The confusion originates at least in part also for them seems to have been Shaw (1909, 1914). He
from a profusion of names published by Lindley, proceeded to describe a number of varieties and
Gordon, and Roezl, and then again after Shaw's formsbased on sizes and shapesof cones, apophyses,
monographic work by Martinez, who included a and umbos of the scales. However,to his six varietal
number of varieties and forms under a new name: shapes of apophyses (Martinez, 1948: 259) several
Pinus michoacana. more, as well as intermediates,could be added from
Lindley (1839) described three "quite new" spe- the materialhere examined.A furtherproblemis that
cies from C. T. Hartweg'scollections of pines from they do not correlate with cone sizes and shapes,
Mexico, all gathered from the vicinity of Real del such as Martinezsuggested.
Monte in Hidalgo. Hartwegbroughtcones (for seeds) Geographically,there is in our view no clear sepa-
and foliage (judging fromthe illustrationsmade from ration into subspecific taxa and in Guatemala,from
it in Loudon, 1842), but the latter has rarely been where Martinez did not study any collections, as
preserved. We regard P devoniana and P macro- many taxa could be assembled. Perry (1991), who
phylla, as a single taxon for which the first name followed Martinez'snomenclature,especially main-
should be retained. A year later, Lindley (1840) tained var. cornuta but, apart from his findings in
published P filifolia from Hartweg's collection on turpentine compounds, gave no new evidence.
the Volcan de Fuego in Guatemala,with very long Curvedcones do not always coincide with pyramidal

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142 Flora Neotropica

apophyses and/orraisedumbos (e.g., Farjon& Mejia light yellowish green to [Link] on

309). This "variety"seems to be sympatricwith the all faces of leaves, in 3-4 lines on the abaxial face
species sensu stricto. That none of these authors and 3-4 lines on each of the adaxial faces. Leaf
had a clear notion of Lindley's concept of Pinus anatomlv:Cross section triangular;hypodermismulti-
devoniana is aptly illustratedby the lectotype in W. layered, with numerous intrusions into the meso-
It consists of three cones, two of them (bound with phyll, some connecting with the endodermis(septal);
rope to preventthem from opening) ca. 25 cm long, resin ducts (2-)3, medial, large; stele terete; outer
only 5-6 cm wide and very "cornute,"and all three cell walls of endodermisthickened;vascularbundles
labeled in Lindley's handwriting "P Devoniana 2, connate but distinct. Pollen cones numerous,
Lindl."Very recently,materialfrom very near to the crowded near the proximal end of a new shoot,
type locality (or at it) was collected (Gardner & subtendedby subulatebracts,forminglong, spiculate
Knees 5259A, E) with cones matchingthe type col- clusters, cylindrical, 20-25 x 4-5 mm, pinkish
lection at W. We agree with Carvajaland McVaugh brown at [Link] peltate,
(1992) that there are, from the existing evidence, no ca. I mm diam., with an erose-denticulate, light-
convincing reasons to maintain these varieties nor coloured [Link] cones subterminal,solitary,in
to make the necessary new combinations under P pairsor whorls of 3-4 on stout, recurved, 10-15 mm
devoniana. However, the polymorphyof the "mton- long peduncles which fall with the cone. Immature
tezumale"group would appear to ask for more de- cones on bracteatepeduncles, ovoid, with pungent
tailed research using different and perhaps more umbos, ca. 15 x 10 mm, purplish or glaucous.
useful approaches, of which molecular methods Maturecones ovoid to ovoid-attenuatewhen closed,
should particularlybe mentioned. often slightly curved, more broadly ovoid with a
flattenedbase when opened, then 7-10 x 5-7 cm.
17. Pinus douglasiana Martinez, Madrono 7: 4. Seed scales ca. 110-130, partingto release the seeds
1943. Type. Mexico. Nayarit:Cerro Juanacata,Jul except at the base, (thin) woody, oblong, straightor
1940, M. Martfinez3429 (holotype, MEXU). slightly curved, light or dark brown abaxially, light
Fig. 34 brown, with often indistinct lighter marks of seed
wings [Link] nearly flat or raised and
Tree, medium to tall, height to 20-45 m, dbh to transverselykeeled, radiallystriateor grooved, rhom-
80-100 cm. Trunkmonopodial, straight, columnar, bic or pentagonal in outline, nearly symmetrical
terete and slender, usually clear of branches2/ of its aroundthe cone, 11-17 mm wide, light brown, red-
length. Barkrough,scaly, divided into large, irregular dish brown or dark brown. Umbo dorsal, raised,
plates and deep fissures,dark reddishbrown, weath- transverselyrhombic,5-7 mm wide, darkerthan the
ering grey-brown; on young trees and branches apophysis, prickle absent. Seeds obliquely obovoid,
smooth or scaly, reddish brown. Branches of first slightly flattened, 4-5 x 3-3.5 mm, light grey or
order long, slender, spreading, or ascending in the brown, often with dark spots. Seed wings articulate
upperpartof the crown; of second and higherorders but effective, held to the seed by two oblique claws,
spreading or ascending, flexible. Crown in young oblong, with a straightand a curved side, 18-24 X
trees pyramidal, in older trees generally rounded, 7-9 mm, light yellowish brown, translucent,some-
dense or open. Shoots uninodal, smooth or rough times with a darkerbrown tinge. Number of cotyle-
with prominent, decurrent, mostly exfoliating pul- dons not observed.
vini, dark brown, not glaucous. Cataphyllssubulate- Distribution and ecology (Map 9). Mexico:
caudate, spreading or recurved, with erose-ciliate Mainly in Jalisco, Michoacan, Mexico, and N Mo-
margins, brown. Vegetativebuds ovoid-conical, the relos, but extending northwardinto Nayarit and the
terminal buds 15-25 x 10-15 mm, the laterals crest of the Sierra Madre Occidental on the border
smaller, not resinous, brown;the scales free, spread- between Sinaloa and Durango; southward it occurs
ing or recurved,subulate-linear,with ciliate margins. locally in Guerreroand Oaxaca. The latter was col-
Fascicle sheaths persistent,remaininglong, (15-)20- lected as Pinuspseudostrobus;in recentyears several
35 mm, with 7-10 imbricate,bracteate,light lustrous populationsformerlyso identifiedhave turnedout to
brownscales with ciliate margins,weatheringgreyish be P douglasiana (cf. Critchfield & Little, 1966:
[Link] in fascicles of 5, rarely4 or 6, spread- map 52). It is a constituentof mostly mixed pine or
ing and drooping in dense tufts, persisting 2-2.5 pine-oak forests at (1100-)1400-2500(-2700) m in
years, slender, lax or sometimes more rigid, 22-35 warmto temperateclimatic zones. The annualprecip-
cm X 0.7-1.2 mm, margins serrulate,apex acute, itation differs with altitude but is roughly 1000 mm

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Systematic Treatment 143

'd,

FIG. 34. Pinus douglasiana (A, C, D, Styles 24: B, Stiles 76; E, Higman, Padilla & Stiles 51; F-H, Farjon & D.
Mejial344). A. Branch with leaf fascicles, pollen cones, and immatureovuliferous cone. B. Shoot with flushing leaf
fascicles and young ovuliferous cones. C. Leaf fascicle. D. Cross section of leaf. E, E Ovuliferouscones. G. Seed scale
(two views). H. Seed (two views). (Magnifications:A-C, E-H, x0.5; D, x40.)

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144 Flora Neotropica

in most areas. Common associated pines are Pilns dermis were found to be similar in P maximinoi
pseudostrobus,P herrerae,Pi leiophylla, P lawsonii, where the two species are sympatric (i.e., in the
P ayacahuite in the southern part of its range, and southernpart of the range of P douglasiana). Shaw
sometimes P oocarpa and P devoniana at lower (1909) had observed the same structureof the hypo-
elevation and drier sites, [Link] the highest dermis in P mnaximinoi, including Hartweg 620, the
and wettest sites it can occur with Abies, Picea (in type collection from Guatemala.
Durango). or Cupressus lusitanica. In many areas Distinction between P doulglasiana and P pseu-
Quercusspp. are co-dominant;a shift in forest com- dostrobus can be similarly vague, as was also ob-
position toward broad-leavedtrees may occur also served by Stead and Styles (1984). Few individual
by selective cutting of pines. Phenology: Pollen dis- characters appear to have a diagnostic value by
persal was observed to occur in Michoacanat 1500 themselves and only in combinationcan they be used
m near the end of February(Styles 24) and may to distinguish between these taxa. The most useful
occur up to a month later at high altitudes. appearto be the hypodermaldevelopment, which is
"normal"in P pseudostrobus,and the cone peduncle,
[Link]-
RANGO: ChavarriaNuevo. 3 Apr 1991, Styles, Favela et al. which persists on the branchin the latterspecies but
7 (FHO, MEXU). GUERRERO:Carrizalde Bravo, 20 Aug falls with the cone in the other two species. Carvajal
1984, Lorea3124 (FHO, MEXU). JALISCO: Along rd. from and McVaugh(1992) mentionthe decurrent,exfoliat-
Tonayato CiudadGuzman,nearEl Jazmin,4 May 1994, ing bractbases (pulvini) on the shoots of P douglasi-
Farjon& Mejia 344 (E, FHO, HEH, K, MEXU, U); ana and P nmaximinoi versus the persistent pulvini
between Bolafos and Guadalajara,20 Sep 1897, Rose 3025
(US); 10 km fromCiudadGuzmin,at G6mezFarias,21 of P pseudostrobus to distinguish them "quickly
and without much question."However, the frequent
Feb 1980, Stead & Styles 524 (FHO): Sierra el Tigre. 10
km from Ciudad Guzmin, 10 Mar 1974, Styles 50 (FHO, misidentificationof these pines would indicate that
K);Tecatitlan.14 Mar1974,Styles76 (FHO.K);Cerrode such "ease" of identificationis not
Alotica, II Mar 1974, Styles & Kemp60 (FHO). MeXIco: experienced by
Valle Bravo,NW of Lagunade Avandaro,2 Oct 1965, most collectors of pines to date.
Sousa 2602 (MEXU).MICHOACAN: Ca. 3.5 km S of
Paracho de Verduzco, at Km 35 along Hwy. 37, 6 May 18. Pinus maximinoi H. E. Moore, Baileya 14: 8.
1994, Farjon & Mejia 350 (E, FHO, K. HEH. MEXU, U); 1966.
Las Playitas Forest Sta.W of Tumbiscatio,27 Feb 1974, Fig. 35
Styles 24 (FHO. K); Dos Aguas, W of Aguililla, 2 Mar Pinus tenuifolia Bentham, PI. Hartweg. 92. 1842, non
1974, Styles 42 (FHO, K). OAXACA:Benito JudirezNational R. A. Salisbury, 1796; Pinus pseudostrobusLindley var.
Park, 20 Mar 1993, Higman. Padilla & Styles 43 (EAP, (Bentham)
tenuifoilia G. R. Shaw,[PinesMexico]Publ.
ESNAC. FHO, MO); Miahuatlan.22 Mar 1993, Higman, Arnold Arbor. 1: 20. t. 13. 1909;Pinus ldouglasiana
Padilla & Styles 51 (EAP, ESNAC, FHO, MO). SINALOA: Martinez var. mariminoi (H. E. Moore) Silba, Phyto-
Guanera.16 Feb 1980, Stead & Styles 476 (FHO, MEXU). logia68: 50. [Link].
NearGuatemala
City, "CanalesMts.," 1841, Hartweg620 (holotype, K,
Uses. Pinus douglasiana is, along with other spe- ex herb.G. Bentham1854;isotypes,B. BM,[Link],
cies with which it often occurs, an importanttimber K (see below), MO, NY, P, W).
tree in most of its range. It is, however,not especially Piniusescalndoniana Roezl,[Link].24.
1857. Type. Mexico. Mexico: Mt. Tzompoli. 1857.
sought out for tree-breedingpurposes, perhaps due Roe:lIs.n. (lectotype,FI,designatedhere).
to the fact that it is relatively unknownto foresters. PiinushoserianaRoezl, Cat. [Link]. Mexic. 24.
Martinez (1943) distinguished his new species 1857. Type. Mexico. Mexico: Mt. Tzompoli. 1857,
from Pinus pseudostrobus var. tenuifolia (Benth.) RoeZs.n. (lectotype,FI,designatedhere).
Shaw (- P maxininoi H. E. Moore) by the morphol- Piinustiomlpoliana Roezl,[Link].24.
1857. Type. Mexico. Mexico: Mt. Tzompoli, 1857,
ogy of the leaves, which in P douglasiana are less Roezls.n. (lectotype,Fl, designatedhere).
slender, flexible, and drooping. The hypodermis of
this species is very strongly developed; the scleren- Tree,medium to tall, height to 20-40(-50) m, dbh
chyma cells form several intrusions into the meso- to 70-90(-I(X)+) cm. Trunk monopodial, straight,
phyll, often connecting with the endodermis. The slender,terete,the lower half of the bole usually free
cones are rather similar but have a thicker, more of branches. Bark thick on the lower part of the
raised apophysis of the seed scales and are smaller trunk,with relativelysmooth, longitudinalplates and
than those of P pseudostrobus. These differences deep longitudinal fissures, grey-brown, on young
were confirmedin a morphometricstudy using Prin- trees and branchesmore or less smooth, grey-brown.
cipal ComponentsAnalysis (PCA) by Stead (1983), Branches of first order slender, longest in the upper
and the taxon was recognized at species rank by partof the crown, often distinctly whorled,spreading
Stead and Styles (1984). The intrusionsof the hypo- or ascending; branches of higher orders sparse,

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Systematic Treatment 145

' "F

/
scale
(three .Seed.
views). (MagA, B X.5;C,x4

FIG. 35. Pinus maximinioi(A-C, F. Hughes 1264; D. Stead & Sn/les 135; E. Hignian, Padilla & Sn/?aes13; G-1.
McCarrter/46). A. Branchwith leaf fascicles and pollen cones. B. Leaf fascicle. C. Cross section of leaf. D. Shoot with
flushing leaves and pollen cones. E. Shoot with flushing leaves and ovuliferous cones. F, G Ovuliferous cones. H. Seed
scale (three views). I. Seed. (Magnifications:A. B. D-IA XO.5 C, X40.)

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146 Flora Neotropica

spreading,slender and flexible, forming a pyramidal darker than the apophysis, weathering grey. Seeds
crown in young trees and an open or dense, rounded obliquely ovoid, slightly flattened,4-6 x 3-4 mm,
crown in maturetrees. Shoots uninodal,with promi- ochraceous or dark brown, with or without dark
nent, decurrentpulvini, green or light brown, rarely spots. Seed wings articulate, effective, held to the
glaucous. Cataphylls subulate-caudate, soon reseed by two oblique claws, oblong, with a straight
flexed, 10-15 mm long, brown with ciliate, hyaline side, widest near the middle, 13-22 x 4-8 mm,
margins, persisting longer than the leaf fascicles. yellowish brown, [Link] cotyledons
Vegetativebuds ovoid-conical, the terminalbuds 15- not observed.
20 mm long, the laterals smaller, not resinous; the Distribution and ecology (Map 15). Mexico:
scales free and spreading,subulate-linear,with ciliate Mainly in the southern half, in Sinaloa, Jalisco,
margins, light brown. Fascicle sheaths 15-25(-30) Michoacan, M6xico, Hidalgo, Tlaxcala, Puebla, Ve-
cm long, persistent,consisting of 8-10 initially dis- racruz, Guerrero,Oaxaca, and Chiapas; and in the
tinct, imbricate, orange-brown scales with ciliate- mountainousparts of Guatemala,Honduras,El Sal-
white margins, later (lustrous) grey-brown to grey. vador,and NW [Link] occurrencein Sinaloa
Leaves in fascicles of 5, rarely 4 or 6, in dense appears to be a clear disjunction (see also Perry,
tufts, persisting 2-2.5 years, slender, lax, drooping, 1991). Pinus maximinoi is a species with a wide
sometimes pendulous,20-35 cm X 0.6-1(-1.1) mm, ecological amplitude,occurringfrom wet subtropical
with serrulatemargins,acute, (yellowish) light green forest, where it is a gap pioneer, well up into the
to [Link] on all faces of leaves, in cooler cloud forestson high mountainsin Mesoamer-
2-4 lines on the abaxial face and in 2-3 lines on ica. In Mexico it also occupies drier sites as a
each of the two adaxial faces. Leaf anatomy: Cross constituent of pine or oak-pine forest or woodland.
section triangular;hypodermismulti-layered,thicker Its altitudinal range is great: (450-)600-2800 m,
in the marginal areas, often with some intrusions with an optimum at 900-1800 m. In the northwest
into the mesophyll, especially on the abaxial side of of its range the species occurs at 1500-2800 m. It
the leaf where they may contactwith the endodermis; occurs on a variety of soils under various climatic
resin ducts (2-)3(-4), medial;stele terete;outerwalls conditions; in Mesoamerica annual precipitation is
of endodermal cells thickened; vascular bundles 2, in the range of ca. 900-2500 mm, with the wettest
with the two xylem strands often connate. Pollen conditions on the Atlantic and Pacific slopes of the
cones crowded nearthe proximalend of new leading mountains. Under these conditions it occurs fre-
shoots, sometimes also on lateral shoots, densely quently with Pinus tecunumanii and Liquidambar
clustered, subtended by subulate-lanceolate,brown styraciflua, at lower altitudes with P oocarpa. In
bracts, cylindrical, 30-40 x 5-8 mm when full- secondary broad-leavedforest other pines may join
grown, light pinkish brown, turning darker. Seed in: P devoniana, P pseudostrobus, and pine wood-
cones subterminal, solitary or in pairs on distinct, land may prevail undera regime of grazing or burn-
stout, curved peduncles which fall with the cones. ing, with the undergrowthdominated by grasses or
Immaturecones ovoid or elliptic, up to 15 x 10 mm, the fern Pteridium aquilinur. In central Mexico, it
with small, spreadingspines, lustrouslight brown or grows at the higher sites with Abies religiosa, P
glaucous. maturing in two seasons. Mature cones ayacahuite, P patula, P pseudostrobus,P douglasi-
narrowlyovoid to ovoid-attenuatewhen closed, more ana, and often Quercus in mixed pine or oak-pine
or less ovoid, slightly curved, with an obliquely forest, where precipitationis more moderatebut the
flattenedbase when opened, (4-)5-10(-12) X (3-) seasonal temperaturerange greater,with some frosts
4-8 cm when open. Seed scales (100-)120-160, occurring in winter. Phenology: The time of pollen
readily parting to release the seeds, thin woody, dispersal of this species is found to be February-
flexible, oblong, straight, spreading 90? or often Marchin Mesoamerica;data for centralMexico were
(strongly) reflexed, mostly symmetrical around the not available.
cone, dark (purplish) brown, with a distinct, light
Representative specimens examined. MEXICO. CHIA-
brown band below the apophysis abaxially and two PAS:At Toliman, 38 km W of Huixtla, 1972, Breedlove &
light brown marks of seed wings on the adaxial Thorne30968 (MEXU, MO); above Colonia Chiapasalong
side. Apophysis flattenedor slightly raised and then rd. to Leyra,II Mar 1993,Higman,Padilla& Styles 13
transversely keeled, 8-15 mm wide, irregularly (EAP,FHO,HEH,MO);Soyalo, 13 Mar 1993,Higman.
Padilla & Styles 22 (EAP, FHO, HEH, MO); San Luis, near
rhombic to pentagonal in outline, variably light
Siltepec, 23 Jan 1945, Matuda5317 (NY); La Grandeza,19
brown. Umbo dorsal, raised, curved, transverse- May 1945, Matuda5583 (NY). GUERRERO: Rinc6n Viejo,
rhombic in outline, 4-8 mm wide, up to 4 mm long, CerroCabezade Toro,10 km W of Aguade Obispo,26

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Systematic Treatment 147

.-i.
LP L CY ? I 1 _ ? 1 ? 1 L ~ ?~ l I~a -. r .
A r~~~~~~~~~~~~~~~~~~~~~~~~~~
. ?
?
L

Solid

- ~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~~~
* ? h
15 Pinus mawninn
Map
IM P n

**0t4~~~~~~~~~4

Feb 1969, Kruse 2406 (MEXU); PuertoSoleares, 6 km SE as determined by A. Farjon). EL PROGRESO: W end of
of El Carrizal. 3 Mar 1984, Lorea 2973 (FHO, MEXU). Sierra de las Minas, above Morazan, 16 May 1979. Stead
JALISCO: Sierra Manantlan,16 Apr 1987, Hughes & Styles 332 (FHO). GUATEMALA:Chinautla."in montibuspraerup-
140 (FHO, K); Mascota, 14 Aug 1976. Lamas & Torres60 tisque 'Canales' dictis. necnon ad pagum Chinanta [Chi-
(UC): along trail from San Sebastianto Arroyo Seco, 8 Jan nautla] prope Guatemala. et summis jugo Choacas prope
1927, Mexia 1421 (BM, F, MICH, MO, UC). MBxico: Salama," 1842, Harhtweg 92 (NY); Chinautla.3 Jun 1977,
Temascaltepec.4 May 1934, Hinton 5775 (NY); Valle de Mittak 8488 (FHO, M). SACATEPEQUEZ: Santa Maria
Mexico, at Tenango, 10 Dec 1950. Matuda 20873 (NY); Cauque, along rd. from Guatemala to Chimaltenango, 7
Nevado de Toluca, 15 May 1954, Matuda et al. 31020 Feb 1978, Stead & Stiles 141 (FHO. MEXU). SOLOLA:
(MEXU). MICHOACAN: Campanario,near Morelia. 4 Jul Between San Pedro and Santiago Atitlan. 4 Apr 1993.
1911,Arsene 6041 (P, US); CerroSan Miguel, 21 Apr 1979, Higman, Padilla & Styles 65 (EAP. FHO. HEH. MO).
MadrigalS. 3283 (ENCB, INIF,MEXU, MS); Uruapan,25 SUCHITEPEQUEZ:Finca Moca, 30 Oct 1934, Skutch 1566
Feb 1980, Stead & Styles 533 (FHO, K); Las Playitas, (F, NY, US).
forest sta. W of Tumbiscatio, 27 Feb 1974, Styles 25 HONDURAS. COMAYAGUA: Plan Cerro Maneadero.
(FHO); 8 km from Uruapan,6 Mar 1974, Styles 49 (FHO). 10.5 km E of Lago Yojoa, 14 Mar 1993, Hawkins 631
OAXACA: San Juan Lachao. 22 Mar 1993, Higman. Pad- (EAP, FHO, HEH, MO, TEFH); El Achote. 18 Feb 1928,
illa & Stiles 52 (EAP, FHO, HEH, MO); Las Galeras, ca. Standlev 56144 (A, F, US). COPAN:Cerro Azul National
100 km NE of Oaxaca, 14 Oct 1960, Little 17953 (IFGP, Park, 12 km NW of Florida, 9 Feb 1992, D. Mejia 40
US); San Gabriel Mixtepec. I Feb 1980, Stead & Styles (EAP, FHO, HEH, MO, TEFH). CORTES: Cusuco National
421 (FHO, K). SINALOA: San Ignacio, Jan 1940, M. Marti- Park, near visitors ctr., 18 km W of San Pedro Sula, 21
nez 2392 (US); PuertoBlanco, Jun 1940, M. Martinez3428 Mar 1993, D. Mejia 359 (EAP, FHO. MO. TEFH); Cerro
(MEXU). VERACRUZ: Las Visas, 2 km E of rd. Xalara- San Ildefonso, between Buenos Aires and Banaderos, 27
Visas, 30 Aug 1989, Zamora C. 1002 (MEXU). Mar 1963, Molina R. 11600 (EAP, F, NY. US); 14 km from
GUATEMALA. ALTA VERAPAZ: Finca Chajsei, S of Cofradiaalong rd. to Buenos Aires, 14 Mar 1978. Stecad&
Coban, 3 Feb 1978, Stead & Stiles 134 (FHO); between Styles 254 (FHO). FRANCISCO MORAZAN: Between El
Segoum and the mines of Caqupej, 3 Feb 1978, Stead & Hatillo and Lo de Ponce, rd. into Montana La Tigra. 24
Styles 135 (FHO, K). BAJAVERAPAZ: El Rancho, 6 Jan Nov 1958, Molina R. 8685 (A, EAP, F, US). INTIBUCA: El
1906, Kellerman5026 (UC). CHIMALTENANGO: Between Achote, above the plains near Siguatepeque, 28 Jul 1936.
Antigua and Lake Atitlan. 18 Feb 1938, Schrenk 2 (MO, Yunckeret al. 6205 (A, F, K, MICH, MO. NY, S). LA PAZ:

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148 Flora Neotropica

[Link] Marcala,13Apr1994,Farjon& Mejia297 (E, material could be traced and is here designated as
FHO, HEH, K, MEXU, U). LEMPIRA:
La Planta, between lectotypes of these obsolete names. (For a discussion
electricpowerplantand Don Tomascampsite,19 Nov of Roezl's catalogues, names, and collections, see
1991,Hawkins98 (EAP,FHO,MO,TEFH);10 km along
SW to Candelaria.
rd. fromErandique 2 km NE of Cerro under P. montezumae.)
Congolon,9 Mar1991,Hughes1428(FHO);[Link] As has been discussed under P. douglasiana, the
NationalPark,8 km SW of Gracias,19 May 1992, D. distinctionbetween P maximinoiand P douglasiana
Mejia 113 (EAP, FHO, MO, TEFH). OLANCHO: El File, has proved to be difficult, leading to misidentifica-
13kmNNEof Catacamas, 2 Jun1992,HawIkins461 (EAP,
tions in the field, in herbaria,and in the literature.
FHO,MO,TEFH);MontafaCoyolito.15 km NE of San
Esteban.14 Sep 1983,McCarter146(FHO). Stead and Styles (1984) concluded that P maximinoi
EL SALVADOR. SANTAANA:Cerro Montecristo, 16 is essentially a Mesoamerican species that extends
km N of Metapan,23 Feb 1989,Hughes1264 (FHO,K, into centralMexico as far as Michoacanand Jalisco,
LAGU). but is there restricted to higher altitudes and the
San Rafael del Norte, 2 Mar
NICARAGUA. JINOTEGA:
1977,Chaplinc312 (FHO).19 Jan 1970,Kemp12 (FHO). Pacific side of the mountains. Gross morphology
shows differences in leaf width, with very thin leaves
Uses. Pinus ilaximinoi is an importanttimbertree in P mtaximinoi(not exceeding I mm), with (as
in most of its range, where it is exploited with other a result) fewer stomatal lines. There is, however,
tall-growing pines. It has also been the subject of considerable overlap in the length of the fascicle
experimentalforestryin varioussubtropicaland trop- sheaths, which contradictsthe figures given by Stead
ical countries underprogramssuch as those initiated and Styles (1984) and Carvajaland McVaugh(1992),
by the Central America & Mexico Coniferous Re- the latterauthorsonly citing specimens from western
sourcesCooperative(CAMCORE).The wood is rela- Mexico. The ovuliferous cones are usually smaller
tively soft and light and easily worked. in this species and have thin woody scales with
Bentham (1842), when describing various plants flattened apophyses. Such differences are quantita-
collected by C. T. Hartweg, named and described tive and, to some extent, even continuous; often a
Hartieg 620 from Guatemala as Pinus tenuifolia. certaindeterminationcan be made only by consider-
which is a later homonym of P tenuifolia R. A. ing a combinationof characters.
Salisbury(1796), a synonym of P strobusLinnaeus.
Moore (1966) correctedthis nomenclaturaloversight, 19. Pinus lumholtzii B. L. Robinson & Fernald,
but in the meantime P. tenuifolia Benthamhad been Proc. [Link]. Arts 30: 122. 1894. Type. Mex-
reduced to a variety of P pseutdostrobusby Shaw ico. Chihuahua:Near Coloradas, May 1893, Hart-
(1909). Later,Shaw (1914) synonymized it with the man 541 (holotype, GH; isotypes, A, K, US).
latter species, but Martinez(1945, 1948) restoredit
Fig. 36
as a species, using the illegitimate name.
Hartwegcollected this pine nearthe city of Guate- Pinutspatltul B. C. Seemann,[Link].Herald336. 1856;
mala in the "CanalesMountains."The type collection non Schlechtendal
& [Link].
"BetweenMesquitaland SantaTe-
Durango/Nayarit:
(K) consists of two sheets, one labeled "620 Guate- resa,"1850,Seetmaunn
1961(holotype,K;isotype,BM).
mala Hartweg,"the other "620 Pinus sp." and with
two locations given. It is thus not certain that both Tree, medium size, height to 20 m, dbh to 50-70
sheets representthe same gathering,despite the same cm. Trunkmonopodial,straight,terete, usually erect.
[Link] firstsheet, from W. D. Hooker's herbar- Bark thick, scaly, very rough,divided into elongated,
ium, bears the inscription"Pinus tenuifolia Benth."; irregularplates and deep, wide, longitudinalfissures,
the second, from G. Bentham'sherbarium,lacks this greyish brown to dark grey, on young trees and
name. Both are sterile shoots; separate cones have branches scaly, flaking very soon, reddish brown,
not been found. The specimen from the Bentham turning grey. Branches of first order long, slender,
herbariumis considered the holotype; the one from spreading or ascending, persistent; branches of
Hooker's herbariumis possibly an isotype. On the higherordersslender,flexible, drooping,the ultimate
reverse of the BM sheet, three localities are men- branches ascending. The crown is broad, rounded
tioned, with the "CanalesMts." as the [Link]- and usually open. Shoots ridged between decurrent
tions like this seem to serve as much to describe pulvini, initially glaucous, turning reddish brown,
where the taxon occurred as to indicate the locality then grey. Cataplhlls subulate, up to 10 mm long,
where it was collected. scarious, with ciliate-hyaline margins, red-brown,
Roezl (1857) describeda few "new species" from early deciduous. Vegetative buds ovoid-conical,
Mt. Tzompoli near Mexico City, of which original acute, the terminalbuds ca. 15 x 8 mm, the laterals

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Systematic Treatment 149

o,
AA
G

~,vty ~ ~ ... F

FIG. 36. Pinus lumholtzii(A, C. Wilkinson& St$les 3; B. D, E, G. H. Stead & Styles 474; F, Hughes 973). A. Shoot
with bud and leaf fascicles. B. Shoot with leaf fascicles and disintegratingfascicle sheaths. C. Fascicle sheaths. D. Cross
section of leaf. E, F. Ovuliferous cones. G. Seed scale (two views). H. Seeds with articulatewings. (Magnifications:A.
B. E, F. x0.5; C. G, H, x I; D, x 30.)

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150 Flora Neotropica

smaller, resinous; the scales imbricate,more or less Distribution and ecology (Map 16). Mexico: In
free at base, narrowlytriangular,acute, with ciliate the Sierra MadreOccidental,in Chihuahua,Sinaloa,
margins, reddish brown. Fascicle sheaths (20-)25- Durango,Nayarit,Jalisco, Zacatecas,Aguascalientes,
35 mm long on juvenile fascicles, consisting of ca. and [Link] altitudinalrangeof this species
10 initially imbricate,ligulate-subulate,thin, reddish is (1500-)1700-2600(-2900) m, which corresponds
brown scales, soon coming apart, forming a tuft of to the lower and middle slopes of the Sierra Madre.
curling scales at the base of the fascicles before they It grows usually mixed with several species of Quer-
fall at maturity,leaving the fascicles bare. Leaves in cus and other pines in pine-oak forest, or on the
fascicles of 3 (exceptionally 2 or 4), remote, usually wetter western slopes of the SierraMadre, in mixed
extremely pendulous, persisting two years, lax but pine forest. Associated pines are, e.g., P leiophylla,
not thin, (15-)20-30(-40+) cm X (1-)1.2-1.5 mm, P arizonica, P douglasiana, P teocote, and P oo-
with serrulatemargins,acute, light green. Stomataon carpa: on more mesic sites one can expect Pinus
all faces of leaves, in 8-14 lines on the convex avacalhuite and Pseudotsuga menziesii, while in dry
abaxial face, in 4-5 lines on each adaxial face, habitatsP. cembroidescan occur with it. The annual
conspicuous. Leaf anatomy: Cross section trans- precipitation,mostly as summer rains, is a moderate
verse-triangular,with convex abaxial side; hypoder- 500-600 mm except on the driest and wettest sites.
mis multi-layered,especially in the marginalregions, Phenology:Time of pollen dispersal not observed.
with minor intrusionsinto the mesophyll;resin ducts
[Link]-
4-10, medial, sometimes 1-4 internal; stele wide, HUAHUA:SierraCanelo,Rio Mayo,24 Sep 1936,Gentry
oval in cross section; outer cells of endodermis not 7 Jul 1936.
2873 (A. F, K, MO,S, UC, US);Basaseachic,
thickened; vascular bundles 2, distinct, the xylem LeSueur557 (ENCB,F,MO,UC,US);PuebloNuevo,Mar
strandsoften connate. Pollen cones concentratednear 1895, Lumholtzs.n. (A); Mt. Mohinora, 1 Sep 1898. Nelson
4914 (F). DURANGO:
LasVeredas,"km20.8 LasVeredas,"
the proximal end of a new shoot but sparsely clus- 2 Oct 1960, Courbasson & Mahieux s.n. (P); Miradorlos
tered, subtended by imbricate, thin, reddish brown Balcones,S of Durangoalongmainrd.,2 Apr1991,Styles,
bracts,cylindrical, 20-30 X 5 mm when mature,at Favela et al. 4 (FHO). JALISCO:Chiquilistlan. 28 May
first pinkish, turningyellow. Seed cones subterminal 1892,M. E. Jones492 (US);Sierrade la [Link]
or lateral,solitary (sometimes in whorls of 2, rarely rd. to Mascota,ca. 12 km NW of Los Volcanes.23 Oct
1952,McVaugh 13733(BM);25 km E of Arandas,14/15
3) on 10-15 mm long, curved peduncles, breaking Nov 1970,McVaugh 24330(MICH);CerroEl Picacho,E
off easily and remaining with the soon deciduous of Tenamaxtlan, 25 Dec 1985, Perez dle la Rosa 1024
cones. Immaturecones broadlyovoid to subglobose, (MEXU);12 km S of Etzatlan,on [Link] Ameca,23
with antrorselycurved spines, maturingin two sea- Oct 1904, Pringle 10014 (A, E, [Link],K. M.
sons. Mature cones ovoid to ovoid-attenuatewhen MEXU,MICH,);9 km fromTapalpatowardChiquilistlan,
22 Sep 1986, Wilkinson & Styles 3 (FHO). NAYARIT:
closed, ovoid to ovoid-acute when opened, then (3-) off [Link] Tepic,14Feb 1980,
MesetaJuanacata
3.5-5.5(-7) X (2.5-)3-4.5 cm. Seed scales ca. 70- Stead & Styles 474 (ENCB, FHO, MO). SINALOA: Rosario,
90, parting gradually,spreading wide or the lower just W of El Palmita, 31 Jan 1962, Breedlove 1693 (DS);
scales remainingclosed, thick woody, oblong, widest 5 km E of Los [Link] rd. from Mazatlinto
Durango.30 Apr 1987, Hughes973 (FHO,MEXU);El
near the apex up to 10-12 mm, the smaller basal Palmito, 28 Dec 1983, Sanders et al. 4449 (UC).
scales remaining connate, dark red-brown on the
abaxial side, dark brown with light brown marks Uses. Pinus lumholtziiis generally known as "pino
of seed wings adaxially. Apophysis slightly raised, triste"for its striking pendulous foliage. It has been
thickened along the distal margin, obscurely trans- overexploited with other pines in some areas for
versely keeled, those on the basal scales gibbous, timber,but due to its scatteredoccurrence,especially
rhombicto pentagonalin outline, ochraceousto red- in pine-oak forest and in drier sites, it is not a
dish brown. Umbo dorsal, often tilted toward the commercially importanttree.
distal marginof the apophysis, flat or slightly raised, Shaw (1909) cited Pinus patula sensu Seemann
up to 5 mm wide, with a minute, soon deciduous (1856), non Schlechtendal& Chamisso (1831) as a
prickle,grey-brown,often darkerthanthe apophysis. synonym of P lumholtzii. Seemann 1961 (BM, K)
Seeds obliquely obovoid, slightly flattened,3-5 mm indeed belongs to this species, and Seemann's inter-
long, dark brown, often with black dots. Seed twings pretationof P patula is incorrect.
articulate,effective, held to the seed by two oblique Robinson and Fernald (1894), in an account of
claws, oblique, widest below the apex, (8-)10-14 X new plantscollected on an expedition led by archae-
4-6 mm, light yellowish or greyish brown. Number ologist Carl Lumholtz,cite Hartman541 as the only
of cotyledons not observed. collection underthis new species and apparentlydid

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Systematic Treatment 151

- 1., a I

'''' ~~~~~~~~~~~
L
a

i~~~~~a
..

I- ~~ ~ ~ ": ? ~ ?I '~

?CI.~~~~~~~~~~~I

r?'
Ma6 iuiIn/ot, (crls at/ arpn/ tinls

not consult other herbariato find if this strikingly 60 or 100-125 cm. Trunkmonopodial,erect, straight
different pine had not been collected earlier. or tortuous,terete, often clear of branches for ?Vof
Carvajal and McVaugh (1992) cite a collection tree height. Bark thick, rough, scaly, breaking into
(McVaugh24330, MICH) with non-pendulous leaf small or large, longitudinal plates and shallow fis-
fascicles; a similarspecimen is representedby Styles, sures, darkbrown to grey-brown,on young trees and
Favela et al. 4 (FHO). Both have ratherrigid, short, branches thin, scaly, reddish brown. Branches long,
spreading leaves 14-15 cm long but otherwise are often tortuous,spreading,forminga roundedor irreg-
entirelysimilar to typical P [Link] collec- ular,open crown. Shoots roughand scaly, with prom-
tions were made in Jalisco and Durango,respectively, inent, decurrentpulvini, reddish brown. Cataphylls
and, being apparently very rare, represent single 10-15 mm long, subulate, recurved, scarious, with
"aberrant"individuals within populations of "nor- erose-ciliate margins, brown, weathering blackish
mal" P lumholtzii. grey. Vegetativebuds ovoid-oblong to fusiform, the
terminal buds 15-25 mm long, the laterals ovoid-
20. Pinus oocarpa Schiede ex Schlechtendal, Lin- acute, smaller, not resinous; the scales imbricate,
naea 12:491. 1838. with free, recurved apices, subulate, ciliate at mar-
Tree, medium to tall, height to 10-15 or 30-35 m gins, brown. Fascicle sheaths up to 25 mm long,
(dependenton variety and site condition), dbh to 40- persistent,not reducedin length, with imbricate,red-

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152 Flora Neotropica

brown scales, weatheringto nearly black. Leaves in xaca, S Veracruz,and Chiapas; widespread in the
fascicles of 3-5, in rigid tufts on thick branchlets, highlands of Guatemala,in Honduras,El Salvador,
persisting2-3 years, straight,rigid or less commonly and NW Nicaragua. Reports (Critchfield & Little,
lax, (I l-)14-25(-30) cm x 0.8-1.6 mm, with serru- 1966; Perry, 1991) of P oocarpa from Belize are
late margins, acute-pungent, lustrous (yellowish) here considered to refer to P tecunumanii.
green. Stomata on all faces of leaves, in 6-8(-10)
lines on the abaxial face and in 3-5 lines on each
of the adaxial faces. Leaf anatomy: Cross section Key to the varieties of Pinus oocarpa
(transverse-)triangular, with a convex abaxial side; i. Leavesin fasciclesof 5 (sometimesalso 3-4 on
treeswithmostly5), (17-)20-25(-30)cm X 0.8-
hypodermis irregularly developed, with intrusions 1.4 mm;seedcones5-8(-10) x 5-9(-12) cm.....
into the mesophyll often connecting resin ducts to ........................................................ 20a var. oocarpa
the endodermis; resin ducts 4-8, usually septal, 1. Leaves in fascicles of 3 (rarely 4), (11-)14-17
sometimes medial or external;stele wide, triangular (-20) cm x 1.2-1.6 mm; seed cones 3-5.5 x 3-
in cross section; endodermis with uniform, thin- 5(-6) cm...............................................20b var. trifolittal
walled cells; vascular bundles 2, approximate, or
with the xylem strands connate. Pollen cones 20a. Pinus oocarpa Schiede ex Schlechtendal var.
crowded near the proximal end of a new shoot oocarpa. Type. Mexico. Michoacan: "Zwischen
but loosely arranged,subtendedby scarious bracts, Ario und dem Feuerberge Jorullo." Schiede s.n.
(no original material has been found in HAL or
oblong-cylindrical, 15-20 x 5-6 mm, pink or red-
dish, turningyellowish brown. Seed cones subtermi- elsewhere); neotype: Michoacan:Las Playitas For-
est Sta., along track to Cahas, Styles 36 (neotype,
nal, at the base of a subsequentshoot, solitary or in
whorls of 2-4 on stout, up to 35 mm long, recurved, FHO, here designated;isoneotype, K). Fig. 37
bracteate peduncles which eventually fall with the Pinus oocarpa Schiede ex [Link]
cones which are persistent for several years after Endlicher,Syn. Conif. 152. 1847. Type. Mexico. [Hart-
weg s.n.] ex herb.LindleY3150 (CGE, labeled as "Pinus
seed dispersal. Immaturecones ovoid to subglobose,
oocarpol)ites.H. L.," is possibly original material).
often wider than long, purplish, then light brown, Pilnusoocarpa Schiede ex Schlechtendal var. mlcflantzoi
maturingin two seasons. Maturecones broadlyovoid Martinez, Anales Inst. Biol. Univ. Nac. Mexico 11:
to subglobose, semi-serotinous, when opened often 70. 1940. Type. Mexico. Hidalgo: San Jose Ocotillas
wider than long, with a flattenedbase, 3-8(-10) x ("habitatin Ocotillos, prope Huasca"), Mar 1939, M.
Martfinez3460 (holotype, MEXU-2 sheets).
3-9(-12) cm when open. Seed scales ca. 70-130,
parting slowly except the proximal scales (semi- Tree, medium to tall, height to 30-35 m, dbh to
serotinous), thick woody, symmetrical, oblong, 100-125 cm. Leaves in fascicles of 5 (sometimes
straight or slightly recurved, dark purplish brown, also 3-4 on trees with mostly 5), straight, rigid or
with light brown marksof seed wings on the adaxial less commonly lax, (17-)20-25(-30) cm x 0.8-1.4
face. Apoplhysisnearlyflat or slightly raised, in some mm. Seed cones 5-8(-10) x 5-9(-12) cm when
cones pyramidal, (weakly) transversely keeled, open. Seed scales ca. 100-130.
rhombic to pentagonalin outline, the apical margin Distribution and ecology. Distribution as given
entire or undulating, lustrous, ochraceous to light for the species. It has the largest range of all of the
brown, weatheringgrey. Umbo dorsal, flat or raised, pines in our region and is especially abundant in
sometimes curved, obtuse or rarely with a minute [Link] extends over a NW-SE distance of
prickle, dark grey-brown. Seeds obliquely ovoid, ca. 3000 km and consequently is found under very
slightly flattened,4-8 X 3-4.5 mm, blackish grey, different ecological conditions. This is expressed in
often with black spots. Seed wings articulate,effec- its altitudinalrange, (200-)500-2300(-2700) m, and
tive, held to the seed by two thin claws, 8-18 x 4- in the variation of annual precipitation. 700-3000
8 mm, greyish brown. Seedlings: Numberof cotyle- mm. These variations are greatest in Mesoamerica
dons not observed. Juvenile leaves persist 3-4 years and are less pronouncedparticularlyin the far NW
and are ca. 30 x 0.5 mm; seedlings 2-4 years of part of the range. Especially precipitationlevels in-
age may sprout from the root after fire. crease toward the southeast. Yet its optimum is at
Distribution (Maps 17, 18). Mexico: From the middle elevation (1000-1500 m) with abundantrain-
Sierra Madre Occidental SE to Mesoamerica, in S fall. Seasonality is mainly expressed in a (long) dry
Sonora, Sinaloa, SW Durango,Nayarit,S Zacatecas, season from October to June in much of its range.
Jalisco, Michoacan, Mexico, Distrito Federal, Hi- Nearly everywhere, fire is an integral part of the
dalgo, N Puebla, Morelos, Tlaxcala, Guerrero,Oa- ecosystem, but man-madefires, often deliberate,add

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Systematic Treatment 153

" ii.
~~~ ~~~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
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l. MapP i usocra a.[i/laa

substantially to the frequency with which they occur. MO); Soyalo. 13 Mar 1993. Hignman,Padlilla& StYles21
Pin'us oocarpa has semi-serotinous cones and is (EAP, FHO, HEH. MO). GUERRERO: Chilapa. 24 Mar
1993, Higman. Padilla & Styles 53 (EAP. FHO, HEH.
adapted to tire, at least at natural frequencies. It Near rd. to Zimapanmines, ca. 7 km from
MO). JALISCO:
occurs in usually open woodland or forest, often in
Hwy. 200 NE of Tuito. 3 May 1994. Farjon & Mejia 342
pure stands or as a constituent of pine-oak woodland. (E, FHO. HEH, K, MEXU. U): Sierra de JuanacatlAn.
Other pines commonly associated with it are P. at La Laguna, n.d.. Nelson 4119 (A, US); "hills near
engelmannii, P leiopphylla, and P douglasiana in the Guadalajara."n.d., Pringle 2455 (BM. F. GOET, K. LE.
M, MEXU. MO, NY): near CiudadGuzm;in. 14 Mar 1974.
northwest and P. maximinoi, P. devoniana, and P
Styles 75 (FHO, K). MICHOACAN: Las Playitas. 1974.
teclunluanii to the southeast. If fires are less fre- Styles 37 (FHO). OAXACA: 38.6 km SW of San Jeronimo
quent, there can be an understoley of shrubs with, Coatlin, 30 Nov 1990. Campos V 3478 (MEXU); 16 km
e.g., Calliandra, Acacia, Leucacena, Hybosena, Byr- NW of El Camar6n along Hwy. 190, 13 Aug 1975, D(-
viclse& Davidse 9602A (MO); ca. 30 km N of Presa Benito
sonimna,and Leucothoe, but often the burning favours
Juarez, near Santiago Lachiguiri. 29 Mar 1989. Hughes
Pteridium aquilinumn or grasses. Phenology: Pollen 1348 (FHO. K, MEXU); 2 km from Gueva de Humboldt.
is released in February-March. 6 Mar 1985. McCarter & Styles 348 (FHO). SINALOA: El
Caj6n, 25 Nov 1975, Nash et al. L19563 (NY); at Hotel
Representative specimens examined. MEXICO. CHIA- Villa Blanca. 3 km SW of Santa Lucia. 8 Jan 1983.
PAS:8 km N of Ixtapa. 12 Feb 1972, Breedlove 24140 Worthingtonet al. 9368 (NY). SONORA: Sierrade Alamos.
(MO, NY); between La Pimienta and El Eden, II Mar near Alamos, 1910, Rose et al. 13087 (F. GH. MO. NY.
1993. [Link] & Styles 12 (EAP, FHO, HEH, US).

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154 Flora Neotropica

ISW~~~~~~~~~~~~~~~~~~"

'1:
go?~~ S. i~~~~~~~~~~~~~~~~~~~~~~
~ 2
?~ ~~~~[Link] waravr vcra

GUATEMALA. BAJA VERAPAZ:El Bucaral. 3 Mar Dec 1838 [capt. Beechey's voyage], Barclay 2698 (BM).
1970. Kemp 70 (FHO); San Jeronimo, Sierra de Chuacus. MORAZAN: CacaguatiqueMtns., ca. 3.5 km E of General
2 Feb 1978, Stead & Styles 126 (FHO). EL QUICHE: Calder6n, 1942, Tucker 766 (F, K, LL, MICH, US). SAN
Chichicastenango, II Aug 1976, Mittak 8258 (FHO); Cu- MIGUEL:Volcan San Miguel, on NW slope, 24 Feb 1942,
nun, 12 Aug 1976, Mittak 8261 (FHO); 12 km N of San Tucker982 (UC). SANTAANA:Cerro Montecristo, 16 km
PedroJocopilasalong rd. to Sacapulas,8 Feb 1978, Stead & N of Metapan, 23 Feb 1989, Hughes 1261 (FHO, K,
Styles 158 (FHO); Chicalte. N of Pozuelo (Cerro la Cam- LAGU).
paia), 9 Feb 1978. Stead & Styles 168 (FHO). GUATE- NICARAGUA. ESTELI:CerroQuiabu,8 km from Esteli,
MALA: Along rd. between San Jose Pinulaand Mataquescu- 19 Oct 1979, Grijalva & Araquistain668 (FHO, MO, NY):
intla, 17 Apr 1994, Farjon & Mejia 301 (E. FHO, HEH, 10 km SSE of Esteli along Hwy. I, I Sep 1983, Nee 27719
K, MEXU, U). TOTONICAPAN:3 km beyond Pologna along (FHO, MO, NY); 14 km S of Esteli. along rd. to San
rd. to Huehuetenango, McCarter & Hughes 53 (FHO). Francisco "TierraBlanca," I Sep 1983, Nee et al. 27730
ZACAPA:Sierra de las Minas, along rd. from Guanacaste (FHO, MO, NY). LE6N: Las Mangas (Bonete), 25 Apr
to Mal Paso, near El Lobo, 8 Mar 1978, Stead & Styles 1974, Chaplin c28(509) (FHO); on volcanic hill above Las
233 (FHO). Mangas, 21 Dec 1969, Kemp 1/69 (FHO); Cerro Bonete,
HONDURAS. CHOLUTECA: Near Choluteca, 3 Jun on hills W of Rio Grande de Matagalpa, 10 Mar 1974,
1965. Lent611 (F, MO, TEX). COMAYAGUA:Los Limones, Styles 54 (FHO, K). MATAGALPA: Locality given as
19 Aug 1988. Cornelius 34 (HEH). COPAN:"Near the 12?55'N,85?47'W, 1971, Kemp87 (FHO).
border"(with Guatemala), 16 Jul 1973, Hazlett546 (HEH);
Ostumdn, 22 Nov 1969, Molima 24741 (F). CORTES:15
km W of San Pedro Sula, along rd. between Cofradiaand 20b. Pinus oocarpa Schiede ex Schlechtendal var.
Buenos Aires, 23 Mar 1993, D. Mejia 376 (EAP, FHO, trifoliata Martinez, Pinos Mexic., ed. 2, 308, figs.
MO, TEFH). EL PARAfSO:10 km S of San Lucas, 17 Mar 252-253. 1948. Fig. 37C-D, G
1982, Hughes & Styles 108 (FHO). FRANCISCO MORAZAN:
El Hatillo, 20 Mar 1985, C. Garcia 102 (NY); 27 km S of Pinus oocarpa Schiede ex Schlechtendalforma trifoliata
Tegucigalpa, 31 Jul 1979, Trott et al. 183 (MO, NY). Martinez, Anales Inst. Biol. Univ. Nac. Mexico 16:
INTIBUCA: Cerro San Crist6bal, W of Esperanza, I Mar 297. 1945 (nom. inval., sine descr. lat.). Type. Mexico.
1985, Rainos 138 (NY). Durango:Sierra de Chavarria,May 1941, M. Martinez
EL SALVADOR. LA UNION:Sierra de Conchagua, 3 3458 (holotype, MEXU).

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Systematic Treatment 155

\ i,.:If \!D F ' '-'

I a *. .R

FIG. 37. A, B, E, F, H-J. Pinus tncairpa var. oocarpa (A, B. Stead & Styles 233; E, Kemp87; F. McCarter& Hughes
55; H, Kemp70; I, J, Stead & Styles 158). C, D, G. P oocarpa var. trifoliata (C, D, Styles. Favela et al. 8; G. Frrjoll &
D. Mejia 313). A. Branch with leaf fascicles and immaturecone. B, C. Leaf fascicles. D. Cross section of leaf. E.
Immatureovuliferous cones. F, G. Ovuliferous cones. H, I. Seed scales. J. Seeds with articulatewings. (Magnifications:
A-C, E-G, J, x0.5; D, x35; H. , x 1.)

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156 Flora Neotropica

[Link] size, height to 10-15 m, dbh to 40- terial could be found (Dr. U. Braun, pers. comm.,
60 cm. Leaves in fascicles of 3, rarelyof 4, straight, 1993) and nothing has been found elsewhere. For
rigid, (11-)14-17(-20) cm x 1.2-1.6 mm. Seed this reason, a specimen collected by Brian T. Styles
cones 3-5.5 x 3-5(-6) cm when open. Seed scales in Michoacan, with characters as described by
ca. 70-90. Schlechtendal,has been chosen as the neotype.
Distribution and ecology. Mexico and Mesoamer- Throughout its vast range, this species remains
ica. Found scattered throughoutthe entire range of remarkablyconstant morphologically. A relatively
the species (not restrictedto W Mexico, as mapped small number of infraspecific taxa have been de-
by Perry, 1991). Its range must yet be considered scribed, mostly by [Link] are var. micro-
imperfectly known. It is probably sympatric with phvlla Shaw (1909), var. ochoterenae Martinez
[Link] in several places, but we have observed (1940), var. manzzanoi Martinez (1940), and var.
small isolated stands of trees having leaves consis- trifoliata Martinez (1948). The first two are now
tently 3 in a fascicle. The altitudinal range of the usually recognized as belonging to distinct species
collections here cited is 800-2500 m. It seems to be (Styles, 1976; Styles & McVaugh, 1990), as they are
a smaller tree, but ecological (edaphic?) factors may in this work. Varietymanzanoi is not well defined.
be largely responsible for this, and perhapseven for Martinez(1940) gives a numberof characterstates
some of its other features. which appearto fall within the variationobserved in
numerouscollections studied here of what we con-
[Link]- sider to be P oocarpa var. oocarpa, both from
RANGO: Chavarria Nuevo, 3 Apr 1991, Styles, Favela et
(l. 8 (FHO, MEXU). MICHOACAN: 5 km from Uruapan Mexico and from Central American countries. The
experimental forest, 6 Mar 1974, Styles 45 (FHO, K). six illustrationsgiven by Martinezof transversesec-
OAXACA: Along rd. from Oaxaca to Tuxtepec (Hwy. 175), tions of leaves, showing medial resin ducts, are
[Link] SanLucas,ca. 30 kmfromOaxaca,23 Apr
slightly misleading: they are free in the mesophyll
1994,Farjon& Mejia313 (E, FHO,HEH,K, MEXU,U); due to the lack of intrusions of the
10 km N of Solade Vega,in the SierraMadredel Sur,12 hypodermis
Nov 1983,Hughes384 (FHO,MEXU);6 kmS of Portillo connecting the endodermis, which is the case in
de Nejapa,along Pan AmericanHwy. from Oaxacato some leaves. The ovoid cone of the type specimen
Tehuantepec,25 Jan 1980, Stead & Styles 383 (FHO); near (Martinez 3460, MEXU) is somewhat atypical but
Miahuatlan [Link] PuertoAngel,26 Jan well within the normalvariationfound in P oocarpa.
1980, Stead & Styles385 (FHO);15 km from Oaxaca
towardTuxtepec,25 Jan 1980,Stead& Styles393 (FHO); Martinez(1945) at first recognized var. trifoliata
13 km fromMiahuatlinalongrd. to PuertoAngel,3 Feb as only a form of P oocarpa, but as he failed to
1980, Stead & Styles450 (FHO);10 km from Oaxaca provide a diagnosis in Latin, his combination was
towardTuxtepec, 26 Mar 1974, Styles 100 (FHO, K). not validly published. Later (Martinez, 1948), he
HONDURAS. INTIBUCA:Arrayanes, valley of Rio
raised it to a variety in the second edition of his
Otoro, along rd. from Toro to La Esperanza,19 Feb 1981.
Stead & Styles 727 (FHO). book on the pines of Mexico. This was mainly
because the 3-leaved characterstate was found to be
Uses. Throughoutits range, Pinus oocarpa is an more constant and [Link] leaves are also
importanttimber tree, especially for local and re- (on average) shorter and thicker and consequently
gional markets. It is also widely tapped for resin. more rigid than those of var. oocarpa, while the
The greatly increasedfrequencyof fires, often delib- cones appear to be smaller. It is also often a more
erately ignited, poses a serious threatto the develop- "depauperate"tree, and one could ask the question
ment of mature stands of this pine. The problem is as to how much of this difference has environmental
especially poignantin Mesoamerica,where both fires ratherthangenetic causes. As the presentdistribution
and infestationsby the pine bark beetle (Dendrocto- map shows, it is likely that its range is imperfectly
nus mexicanus) are frequentlyfollowed by "lumber known, althoughwith the massive collecting that has
salvationoperations"and subsequentconversion into been undertakenin Mesoamerica in recent decades,
pasture(Perry, 1991). one would expect more localities there if it is at all
Schlechtendal (1838) gave a rather concise and common.
vague description of Pinus oocarpa, but character-
ized the typical, broad cones [Link] 21. Pinus praetermissa Styles & McVaugh,Contr.
C. J. W. Schiede is credited with its discovery Univ. Michigan Herb. 17: 310, fig. 2. 1990. Type.
"zwischen Ario und dem Feuerberge Jorullo" (in Mexico. Nayarit: Cerro Juanacata, 14 Feb 1980,
Michoacan) as well as with its name, no collection Stead & Styles 475 (holotype, FHO; isotypes,
was specifically [Link] HAL no original ma- ENCB, MEXU). Fig. 38

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Systematic Treatment 157

N ''

? .
| .:D :;.'

: '. ..

FIG. 38. Pinus proetermissa (A-C, Steadl& Stleus 475; D. Wilkinson& Styles 2; E-G, Phre de la Rosa 1499). A.
Shoot with leaf fascicles. B. Leaf fascicle. C. Cross section of leaf. D. Closed ovuliferous cone. E. Opened ovuliferous
cone: (a) lateral view, (b) proximal view, the basal scales missing. F. Seed scale (three views). G. Seed. (with) articulate
wings. (Magnifications:A, B. D-G. 0.5; C. X40.)

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158 Flora Neotropica

Piulls ooc'arpaSchiede ex [Link]/k base when opened and missing the proximal scales,
G. R. Shaw, [Pines Mexico] Publ. Arnold Arbor. I: 27, (4-)5-6.5(-7) x (5-)6-8 cm when open, falling 1-
t. 20, figs. 2, 5, 8-11. 1909. Type. Mexico. Sinaloa: 3 years after maturity. Seed scales ca. 100-120,
Colomas, 1897, J. N. Rose 1755 (lectotype, US 300624-
5, designatedby Styles & McVaugh,1990; isolectotype, parting wide at maturity,the proximal scales some-
A). times abscising before the cone falls, thin woody but
rigid, oblong, straight or recurved, more or less
Tree, small to medium, sometimes bushy, height symmetrical, chestnut-brown, with light brown
to 10-15(-20) m, dbh to 30 cm. Trunkmonopodial, marksof seed wings on the adaxial side. Apophysis
tortuousor curved, sometimes branchinglow. Bark flat to slightly raised, radially striate or ridged to
relativelythin, rough and scaly, exfoliating with thin, transverselykeeled, rhombicto pentagonalin outline,
grey-brownplates, irregularlyfissuredlongitudinally, with a rounded or angular upper margin, lustrous
on young trees and branches thin, rough, reddish light brown or yellowish brown. Umbo dorsal, flat
brown. Branches long, spreading,tortuous,not pen- or slightly raised, 2-4 mm wide, more or less rhom-
dulous, forming a broad, irregular, open crown. bic in outline, obtuse, grey-brown. Seeds obliquely
Shoots slender,with weakly developed, early eroding ovoid, slightly flattened, 5-8 X 3-4 mm, blackish
decurrent pulvini, soon scaly and flaky, reddish grey or with black dots. Seed wings articulate,effec-
brown, turninggreyish. Cataphyllssmall, 7-10 mm tive, held to the seed by two thin claws, thickened
long, subulate, not recurved, scarious, with erose- above the base, obliquely obovate-oblong, 12-18 x
ciliate or hyaline margins, light brown, deciduous 5-8 mm, greyish brown. Number of cotyledons not
with leaf fascicles or slightly later. Vegetativebuds observed.
oval-oblong to cylindrical, the terminal bud 10- Distribution and ecology (Map 19). Mexico: S
15 mm long, the laterals ovoid-acute, smaller, not Sinaloa, Nayarit, and Jalisco. Its distribution must
resinous;the scales imbricate,straight,subulate,with still be relatively unknown,due to likely misidentifi-
ciliate or hyaline-erose margins. Fascicle sheaths cation with P oocarpa; however, it is probably re-
short, 10-14 mm long, with chartaceous,light brown stricted to the region covered by McVaugh's(1992)
scales with white or hyaline margins, persistent, Flora Novo-Galiciana. Perry (1991) reports this
weathering dark brown to grey. Leaves in fascicles taxon from nearConcordiain Sinaloa (as P oocarpa
of (4-)5, in dense, lax tufts on ultimate branches, [Link]),but his map gives a wider distribu-
persisting up to 3 years, straight, delicate but not tion in thatstate which is not substantiatedby herbar-
pendulous, (8-)10-16 cm X 0.5-0.8 mm, serrulate ium [Link] praetermissaoccurs in open, dry
at margins,acute, bright green. Stomlataon all faces pine-oak woodlandsor tropicalbroad-leavedforests,
of the leaves, in (2-)3-5(-6?) lines on the convex often on rocky slopes. Its altitudinalrange is 900-
abaxial face and 2-4 lines on each of the adaxial 1900 m. The dry season is from November to May,
faces. Leaf anatomy: Triangular in cross section; annualprecipitationvaries between 1000-1500 mm.
epidermis thin; hypodermis 1-2-layered, uniform, Associated pines are P devoniana, P luimholtzii,
sometimes locally thicker,especially in the marginal and possibly P oocarpa, but furtherinvestigation is
areas; resin ducts 1-2(-4 accordingto Loock, 1950), necessary to evaluate its sympatry with these and
internal,occasionally I septal; stele terete or nearly other pines. Phenology:Time of pollen dispersal not
so in cross section; cell walls of endodermis small, observed.
thin-walled; vascular bundles 2, the xylem strands
connate. Pollen conies crowded near the proximal
[Link].
end of a new shoot, subtendedby light brown bracts, ISCO:Mun. La Manzanilla,Las Canoas, 1900, Diaz L. 1724
ovoid-oblong to cylindrical, 10-15 x 5 mm, pink along rd. from Magdalenato
(ENCB);Hostotipiquillio,
to reddish, turning yellowish brown. Seed cones Ixtlandel Rio, 21 Jan 1988, Perezde ia Rosa 1499 (FHO);
Volcan Tequila, 25 Oct 1970, Webster& Breckon 15970
subterminal,commonly solitary, occasionally oppo-
(MEXU, MICH); Hostotipiquillio, 21 Sep 1986, Wilkin-
site, on up to 35 mm long, recurved,slender pedun- son & Styles 2 (FHO). NAYARIT: Between La Laguna and
cles, deciduous, non-serotinous,falling without the Santa Maria del Oro, 4 Dec 1977, Chazaro 736 (ENCB):
basal scales and peduncle in most cases. Immature Mun. de Jesus Maria,Santa Teresadel Nayar,5 Apr 1978.
cones broadlyovoid to subglobose (wider thanlong), Diaz L. 9582 (ENCB, GUADA); Ixtlin del Rio. Jul 1940.
M. Martinez 3558 (F); Jala, Jan 1940, M. Martilez 4085
with acutish apex, purplish,soon light brown, matur-
(US);Juanacatic,1900,M. Martinez4096 (IFGP);Ixtlan
ing in two seasons. Maturecones broadly ovoid to del Rio,Jul 1940,M. Martinez4097 (A, GH);La Laguna,
subglobose (wider than long), often with a flattened 3.5 km E of SantaMariadel Oro, 15 Sep 1960,McVaugh

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Systematic Treatment 159

MapU19.
praetermisscl. Pinus

am. ~ ~ . - -

. . -.

'at' . . .. i I
Map 19. Pi,ws raeierii1?(

19030 (MICH);Tepic, 5 Jan 1892. Palmer 1998 (A, F, NY. species. The ovuliferous cones, although strikingly
US): "3.5 miles W of Jalisco." 8 Aug 1956, Phillips s.,. similarto those of P oocarpa. show some anatomical
(DS, MICH, UC); Sierra Madre,near Colomas, 1897. Rose traits resulting in a distinct characterwith biological
1997 (A, NY, US). SINALOA: "San Ignacio,"Jan 1940. M.
Martinez4095 (NY). significance. Unlike in P. oocarpa. the seed scales
partwidely upon maturity,the proximalscales persist
Uses. This scattered species is rarely recognized on the peduncle or break off, and the cone falls
as distinct from P. oocarpa by foresters, and conse- after the seeds are released. In contrast, cones of P
quently no specific uses are known. It is likely to be oocarpa are semi-serotinousand persistent,a charac-
exploited for timber together with other species. ter common in pines of fire-subclimaxcommunities.
Shaw (1909) considered the widespreadtaxon Pi- Seed dispersal in P. praetermissa is therefore inde-
nus oocarpa to be a species easily recognized by its pendent of recurrentfires and this in turn suggests
characteristichard,broadlyovoid, lustrouscones and that the seedlings of this species can compete with
the prevalence of septal resin ducts in the leaves. other plants more successfully. The leaves are quite
Because of this typology, he described a pine from different in both species, with a more delicate mor-
Colomas, Sinaloa (Rose 1755), and one from Tepic, phology in P. praetermissa resulting in very slender,
Nayarit(e.g. Palner 1998. Rose 1997), with similar lax leaves with fewer lines of stomata per unit leaf
cones but with relativelyshortand slender,lax leaves width; the hypodermisis thinnerand there are fewer,
in fascicles of 5 as P. wocarpa var. microphylla. internalresin ducts in this species. There are further
Subsequentauthors(Martinez, 1945, 1948; Loock, differences in the shoots, and P praeternmissahas
1950; Farjon, 1984; Perry, 1991) accepted this treat- smaller, less persistent cataphylls. These characters
ment, while Silba (1986) even sank it into synonymy and the likelihood that the two species are allopatric
with P. oocarpa. Recently, revision (Styles & indicate that they are ecologically different as well.
McVaugh, 1990) of the material studied by Shaw No specimens with these charactershave been found
and Martinez,in additionto collections more recently elsewhere in the very extensive range of P oocarpa;
made, has led to recognition of this taxon as a good it seems likely that they are restrictedto a limited

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160 Flora Neotropica

area of the Pacific slope (Carvajal & McVaugh, (narrowly) ovoid or ovoid-attenuate when closed,
1992). usually slightly curved, more or less asymmetrical
at base, 5-10(-12) X (3-)4-6.5 cm when open. Seed
22. Pinus patula Schiede ex Schlechtendal& Cham- scales ca. 100-150, serotinousor partingsome time
isso, Linnaea6: 354. 1831. after maturing, tenacious or deciduous with the pe-
duncle, (thin) woody, oblong, usually curved when
Tree, usually tall, height to 35-40 m, dbh to 90- spreading, the proximal scales connate, purplish
100 cm. Triunkmonopodial, erect, straight, slender, brown to dark brown, with light brown marks of
terete, commonly free of branchesfor 2/ or more of seed wings [Link] to slightly
its length. Bark thick, rough and scaly, with large raised, transverselykeeled, more or less gibbous on
elongated plates and deep, longitudinalfissuresespe- the proximalscales, more so on one side of the cone,
cially on the lower partof the bole, darkgrey-brown rhombic to obtrullatein outline, upper marginacute
below, lighter above, on young trees and branches or obtuse-rounded,abaxial surface striateor smooth,
thin, scaly, reddish brown. Branches of first order (lustrous) ochraceous or yellowish brown. Umbo
long, slender, spreading or slightly ascending; of dorsal, flator raised,often sunken into the apophysis,
higher orders slender, drooping or subpendulous, 3-7 mm wide, grey, with a minute,deciduousprickle.
the ultimate branches pendent, forming a conical to Seeds obliquely ovoid, flattened, 4-6 x 2-4 mm,
domed, open crown. Shoots often multinodal,rough blackishgrey. Seed wings articulate,held to the seed
and scaly when the leaf fascicles have fallen; foliage by two oblique claws which partly cover the seed
shoots with prominent,decurrentpulvini, yellowish on one side, obliquely ovate-oblong, 12-18 x 5-
brown to reddish brown. Cataphy!llssubulate, re- 8 mm, light brown with dark stripes. Number of
curved at apex, scarious, with erose-ciliate margins, cotyledons not observed.
brown, early deciduous. Vegetativebuds oval-oblong Distribution (Maps 16, 20). Mexico: In Tamauli-
to cylindrical, the terminal bud 15-20 mm long, pas, Queretaro, Hidalgo, Mexico, Distrito Federal,
the laterals shorter, brown, not resinous; the scales Morelos, Tlaxcala, Puebla, Veracruz, Oaxaca, and
spreading, subulate, ciliate at margins. Fascicle Chiapas.
sheaths initially long, 20-30 mm, with 6-8 imbricate,
chartaceous,whitish yellow to orange-brownscales,
persistentbut reducedto 12-15 mm in maturefasci- Key to the varieties of Pinus patula
cles, tight, slowly weatheringto [Link] I. Seed cones on (very)shortpeduncles,seemingly
in fascicles of 3-4(-5), in drooping tufts, typically sessile, tenacious, persistent, serotinous. (5-)7-
10(-12) cm long, (narrowly) ovoid when closed.
in two rows on either side of the upturnedshoot,
broadly ovoid when open.......................22a. var. patult
persisting 2-3 years, thin, lax, drooping to pendent, 1. Seed cones on distinct, curved peduncles, decidu-
( I1-)15-25(-30) cm x 0.7-0.9(-1) mm, serrulateat ousaftera fewyears,openingreadily,5-8 cm long,
the margins,acute, pale green to darkgreen. Stomata narrowly ovoid to ovoid-attenuate when closed,
on all faces of the leaves, in 4-6(-7) lines on the (narrowly)ovoid when open...................................
......................................... 22b. var. IongipledluncultaI
convex abaxial face and 2-3(-4) lines on each adax-
ial face. Leaf anatorny: Transverse-triangularto trian-
22a. Pinus patula Schiede ex Schlechtendal &
gular in cross section; the hypodermis usually with
Chamisso var. patula. Type. Mexico. Veracruz:
a double layer of cells; resin ducts (1-)2-3(-4),
Cruz Blanca, "Inter Cruz blanca & Jalacingo"
medial, occasionally I internal, stele oval in cross
(sheet I of 4 at HAL), Nov 1828, Schiede & Deppe
section; outer walls of endodermalcells not thick-
1108 (lectotype, HAL, here designated).
ened; vascularbundles 2, distinct, the xylem strands
often connate. Pollen cones crowded near the proxi- Fig. 39A-F
mal end of new shoots, spreading, subtended by Pinus patuki Schiede ex Schlechtendal& Chamisso var.
scarious bracts. ovoid-oblong to cylindrical, 15-20 macroca(rpa M. T. Masters,[Link]..ser. 3, 9: 438.
x 5-6 mm, pinkishyellow, turningyellowish brown. fig. 92. [Link].,ser. 3, 9: 435. fig.
92. 1891. (lectotype, here designated).
Seed cones subterminalor lateral, in whorls of 2 to
many, rarely solitary, persistent or deciduous, on Tree, as in the species. Seed cones (very) short
short or moderately long (to 20 mm) peduncles. pedunculate, seemingly sessile at maturity, in whorls
Immaturecones ovoid, on short, or long, recurved of 2 to many, tenacious, persisting many years, serot-
peduncles with persistent cataphylls, purplish,soon inous, ovoid when closed, broadly ovoid and (5-)7-
light brown, maturingin two seasons. Maturecones 10(-12) x (3-)4-6.5 cm when open. Seed scales ca.

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Systematic Treatment 6
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6

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,~~~~~~~1 _...41'''?' -t~1:.. ' ?. .~ ?

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~~~~~~~~~
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?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~1 '' "~ ?''".

? ? ? ~. i.? r
.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
I~L? I~ , J~k ' " L. i ?.??; ,?.-. .'.;? .
..~... .i': . ? .. . .
i
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,
........ . , . -~~.:
..~?,. ,..
Map 2 ....:
.Pinus va.,Ioni
. . ,....
euclt ..
patula .:

120-150, (thick) woody, rigid, many of the proximal Be,u,Oi 4526 (TEX. UC. US). HIDA(LG;O: Along Hwy. 85
scales remainingconnate. NE of [Link] hamlet of Maguey Verde. 28 Apr
1994. IF(rjorl& Meji' 331 (E. FHO. HEH. K. MEXU. U):
Distribution and ecology. Mexico: In a few local- Mineraldel Monte. Rancho del Guajolote. 1839. Hclrtwneg
ities in Tamaulipas,in Quer6taro,Hidalgo, Mexico, 443 (BM. Fl. K). 1839. Hartweg s.n. (P): Tenango. 9
Distrito Federal, Morelos, Tlaxcala, Puebla, Vera- Nov 1985. Lorewce & Hernilddez 4910 (MEXU); Vega
cruz, Oaxaca, and Chiapas. Its main distributionis Quemada. 16 Jul 1977. Mitutik8500 (BANS. FHO. M): 14
on the Sierra MadreOrientaland the high mountains km SW of Huachinango.29 Apr 1983, Nee & Tv,lor 26971
(NY): Honey railway sta.. 25 Nov 1903. Pringle 8785 (A.
of east-centralMexico. Its altitudinalrangeis (1400-) BM. E. GOET. K. M. MEXU. MICH. MO. NY); Sierrade
1800-2800(-3300) m. Pinus patula var. patula is Pachuca. 10 Dec 1907. Pritgle s.n. (MICH): Huachinango.
generally restrictedto humid, subtropicalto warm- 18 Mar 1980. Stead & Styles 663 (FHO): 29 km NE of
temperate sites, with annual precipitation 1000- Valle Metquititlin. 4 Apr 1987. Zepeda 34 (CIIDIR). MEXIco:
de Mexico. "toret de la Deserta Vieja." 7 Sep 1865.
2200 mm. On the eastwardslopes of the Sierra Ma-
Bottrgeal 904 (Fl. K. P. S. US): Valle de Mexico. 7 Jan
dre Occidental these amounts are enhanced by fre- 1951. MatIudi 20879 (NY). [Link]:Ca. 4 km W of
quent fog outside the rainy season. It occurs in a Huitzilac, S slope of Scrjana de Ajusco. 14 Jul 1960.
variety of mesic forest types, associated with other liti. & Koeppen 221 (UC); Sierra Encantada. along rd.
from Mexico City to Cuernavaca. 10 Apr 1965. R. C.
pines such as Pinus pseudostrobuxs P. greggii, P
Pala'cioss.N. (ENCB). OAXACA:Benito Juirez National
maxriminoi, P. hartwegii, P. cacalcchite; on drier sites [Link] Yuvila. 20 Mar 1993. [Link] & Styles 42
it is replaced by P. teocote, P. [Link], and P (EAP, FHO, HEH. MO): on the summit of Mt. Talea. 1841.
leiophylla. In some locations Abies religiosa occurs Liebhma,t 14805 (MO. S. UC); along rd. from Oaxaca to
with it. Mixed pine-oak forests are most common in Tuxtepec at Brecha 1010. 28 Mar 1974. Styles 113 (FHO.
1841. Liebhminas.n. (NY):
the interior, while on the Atlantic slope numerous K). PUEBLA:Chinantla. May
Chinantla. 1841. Lielbimtai14803 (MO). QUERITARO: Pi-
are
angiosperms,especially Liquidambar. (co-)domi- nal des Amoles. 1900. Nelso,n 3936 (A. US). SAN LUiS
nant. Phenology: Pollen dispersal is recorded in POrosi: SierraConeja. near Llano de Coneja. 13 Jun 1948.
Februaryand March (Critchfield, 1966b). NewuIrtn44 (US). TANIAULIPAS:E of Dulces Nombres. 18
Jul 1948. Meyer & Rogers 2802 (E. MO): G6mez Farias.
[Link]- n.d.. Sulllivra 518 (TEX). TI,AXCALA: Ci6nega Larga. be-
TRITO FEDERAL: Desiertode Las Leones. 24 Sep 1961. tween Zacatlln and Tlaxco. 26 Nov 1960. Madrigal S.

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162 Flora Neotropica

I; i

FIG. 39. A-F. Pinus patula var. paltul(i(A-C, Higiman,Pdilli & Styles 42: D-F, Firjon & D. Mejtia331). G-l. P
p1tmiu1var. longipedluncultua (b)ock 113A). A. Shoot with (flushing) leaf fascicles. B, H. Leaf fasciles. C. Cross section
of leaf. D. Ovuliferouscone. E. Seed scale (three views). F. Seed, articulatewings. G. Branchwith foliage and ovulifer-
ous cones. I. Immatureovuliferous cone. (Magnifications:A. B. D-l, x 0.5; C. x 50.)

s.n. (INIF); near Tlaxco, Dec 1962. Savior 22 (IFGP). Nee & Diggs 24931 (F, FHO, MEXU. NY, XAL): Los
VERACRUZ: Pico de Orizaba, 1855, Muller517 (NY): Mun. Altos, 11 Mar 1983, Nee & Tvalor25891 (CAS, F, K. MO,
Acajete. 5 km SE of Las Vigas along Hwy. 140. 1900, NY. TEX,); Nueva Vaqueria, 12 km from Excola, 1900,

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Systematic Treatment 163

Nee & TaYlor27061 (F, K, NY, TEX, XAL); on Cerro 6 km N of Huayacocotla.

20 Nov 1982, Acosta S. &
VolcanilloN of Cofre de Perote, near Las Vigas, Aug 1894, Barrios 299 (MEXU).
Stahl s.n. (JE); Las Lajas, 21 Mar 1980, Stead & Styles
695 (FHO); near Perote, 1I Jun 1974, Duncan 2451 (UC).
UNKNOWN LOCALITY:Locality not stated, 1857, Roezl s.n. Uses. This species is one of the most important
(FI);"Malpaysde la roya,reg. frig.."Nov 1828,[Deppe pines for timber in Mexico, as it grows quickly and
s.n.] ex herb. Lindley s.n. (CGE), June 1829, Schiede s.n. producesa long, straightbole, for the most partfree
(HAL). of branches. It is also widely introduced in other
tropical countries in plantation forestry, where in
some cases it has become problematicas an invasive
22b. Pinus patula Schiede ex Schlechtendal &
Chamisso var. longipedunculata Loock ex Martf- species.
Schlechtendal and Chamisso (1831) gave under
nez, Pinos Mexic., ed. 2, 333, figs. 276-280. 1948
C. J. [Link]& F Deppe 1108 a concise descrip-
("longepedunculata"). Type. Mexico. Oaxaca:Ran- tion of a 3-leaved
cho Benito Juarez, (Rancho Tablas), Jul 1947, pine, of which they had seen no
cones ("strobilo . . . nobis deficiente") in the mate-
Loock 113a (holotype, PRF-n.v.; isotype, MEXU).
rial presentedto them. At HAL there are 4 sheets (3
Fig. 39G-1 with this collection number),all with small, sterile
branchlets. Sheet I of 4, marked by A. Farjon, is
Tree, as in the species. Seed cones distinctly pe- designated as the lectotype. Another
specimen at
dunculate, peduncles up to 20 mm long, curved, HAL collected in June 1829 by Schiede (&
Deppe?)
falling with the cones after a few years. Cones has 4 immatureovuliferouscones. The sheet at CGE
opening readily, narrowly ovoid to ovoid-attenuate (collected by Deppe, Nov. 1828) only has a sterile
when closed, (narrowly)ovoid when open, then 5-8 branchlet.A box with cones
(Lindley's herbarium
X 3.5-4.5(-5) cm. Seed scales ca. 100-120, (thin) cat. no. 3194),
despite E. L. Little's annotation,bears
woody, somewhat flexible, partingreadily to release no proof that it belongs with the sheet and seems to
the seeds. be of two separate gatherings of this species. The
Distribution and ecology. Mexico: Known from serotinous cones are thereforedescribed from other
only two localities in Hidalgo and Veracruzbut more materialnot associated with the protologue.
widespreadin Oaxaca and centralChiapas. Probably Gordon (1875) described Pimlls )lpatula(var.) mac-
incompletely known. Ecology and phenology are rocarp,a based on collections by C. J. W. Schiede
similar to [Link], but its ecological amplitudeis with a cone size "from6-7 inches long and 2 inches
narrower;it apparentlydoes not occur on the interior broad",which, if correct, would be quite outside the
mountainsof Central Mexico, where conditions are
rangeof this species as here defined. Masters(1891)
drier and seasonal temperaturefluctuationsmore ex-
formallypublishedthis combinationand providedan
treme. illustrationof the cone. No specimen has been traced
in the herbariato match the descriptionsand/or the
Representative [Link]- illustration. The illustration in Masters (1891) is
PAS:4 km E of San Crist6balde las Casas,alongrd. to therefore
Las Ollas, 20 1994. & Mejhi 310 (E, FHO.
designated as the lectotype.
Apr Farjon
HEH. K, MEXU. U). HIDALGO: I km W of Llanos Gran- Martinez (1948) ascribed his new variety P piatula
des. 23 Oct 1976, Medina 1673 (ENCB. MEXU). OAXACA: var. longip/edunculatato Loock. who coined the
[Link],SantaMartaLatuviin the name on the label of the type specimen. He described
SierraNorte,28 Feb 1986,AcostaS. & Neris.n. (MEXU); it as different from "typical"P patuilain having 2
[Link],26.7 km SW of SanJer6nimo.2
Dec 1990,Camnpos V 3516 (MEXU);SanPabloMacuilti- resin ducts in the leaves instead of 3 and in having
anguis, I May 1977. Echienique53 (MEXU);San Juan pedunculate(5-15 mm) cones versus sessile cones.
Atepec, 23 Apr 1994, Farjon & Mejia 316 (E, FHO. Perry(1991) allows it occasionally 3 resin ducts.
HEH,K, MEXU,U); Lachatao.6 km fromSan Antonio Barrett (1972), in comparing many specimens
along rd. to San PabloYaganiza,10 May
Cuajimoloyas from most of the range of P. patula, found clinal
1986, Gereau & Mover 2145 (FHO, MEXU, MO): Benito
Juarez National Park. 20 Mar 1993. Higman, Padilla & variationfrom northto south in which the predomi-
Styles 40 (EAP,FHO, HEH, MO); along Hwy. 175 between nant numberof leaves per fascicle increases from 3
Oaxaca and Tuxtepec,between Ixtlin de Juarezand Llanos to 4(-5). A similar cline was reportedfor the length
de las Flores, 15 Mar 1984, McCarter & Styles 266 (M, of the cone peduncles,but accordingto Styles (1976)
MEXU), 15 Mar 1984, McCarter & Styles 271 (FHO,
the distributionof peduncle length is more random.
MEXU). McCarter & Styles 273 (MEXU); Mun. Santa
CatarinaLachatao,Santa MartaLatuvi in the Sierra Norte, Styles (1976) concluded that "it agrees phenotypi-
8 Feb 1986, Stinche:A. & Neri s.n. (MEXU). VERACRUZ: cally in all botanical characters"with those men-

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164 Flora Neotropica

tioned by Martinez (1940) under P oocarlpa var. long, light brown,persistentbut reducedto 8-10 mm
oclhoterenae and synonymized both under P patula. on maturefascicles, weatheringlight grey. Leaves in
He later expressed a different view (Styles, 1984) fascicles of (4-)5, rarely 3, in open or more dense
but stopped short of formally synonymizing these tufts at the ends of branches, persisting 2-3 years,
two taxa under his newly created combination P slender, lax, erect or slightly drooping, 12-18(-22)
/patula subsp. tecunumanii. From his determinations cm X (0.5-)0.6-0.8 mm, with serrulate margins,
of specimens it appearsthat P oo(arpa [Link]- acute, light green to yellowish green. Stomata on all
nae Martinezwas treatedas a synonym of P. patula faces of leaves, in (3-)4(-5) lines on the convex
subsp. teclunumanii(see under P tectnumanii), but abaxial face and usually 3 lines on each adaxial
that at least more-recentlycollected material (Hig- face, sometimes [Link] anatomy:Cross
mnan,Pradillt& Styles 40) was again accepted as P. section triangular,with convex abaxialside; hypoder-
patiula(subsp.) [Link] the mis uniform,with 1-2 layersof cells; resinducts (1-)
field and study of type materialby A. Farjonconfirm 2-3(-5), septal, sometimes 1-2 small ducts internal;
recognition of this taxon as a variety if peduncle stele terete to obtuse-triangularin cross section;
length is coupled with persistence and serotiny, outer cell walls of endodermis thickened; vascular
which may point to differentadaptivetraitsbetween bundles 2. with connate xylem [Link] cones
the two taxa. crowded near the proximal end of a new shoot,
ovoid-oblong to cylindrical, 12-18 X 5-6 mm, sub-
tended by scarious bracts, purplish yellow, turning
23. Pinus jaliscana Perez de la Rosa, Phytologia54:
290. 1983. Fig. 40 light brown. Microsporophyllspeltate, the distal part
ca. 1.5 mm wide. Seed cones subterminal,solitary
Piiius pulHllt Schiede ex Schlechtendal& Chamisso var. or in whorls of 2-3 on stout, curved, 7-15 mm long
j(lisc(tniI (Perez de la Rosa) Silba, Phytologia 58: 368. peduncles which remainattachedto the fallen cones.
1985. Type. [Link]:Along rd. 25 km from Immaturecones ovoid, 10-15 mm long, purplish
Zimap;in mine,takingoff Hwy.2(X)just N of El Tuito.
19 Jun 1983,[Link] la Rosa370 (holotype,IBUG; red, maturingin two seasons. Maturecones ovoid-
[Link],F, GH. K. [Link].P, US). oblong to ovoid-attenuate when closed, often
Carvajal,Phytologia59: 139. 1986:
Pinlls mac(iauglhii oblique at base, (4.5-)6-8.5(-9.8) cm long, (3-)4-
Piius oocarpa Schiede ex Schlechtendalvar. mac- 5(-6) cm wide when open. Seed scales ca. 90-115,
viaughii(Carvajal) 68: 57. [Link].
Silba,Phytologia
[Link]:[Link] de Puriticaci6n, El Salto partinggraduallyexcept for the proximalscales, ob-
del Rinc6n, 24 Oct 1983. Ctrvnajtl 4588 (holotype. long, straightor slightly reflexed, thick woody, pur-
CREG-n.v.; isotype. MICH). plish brown on the abaxial side, dark brown, with
light brown marks of seed wings, on the adaxial
Tree, medium to tall, height to 25-30(-35) m, side. Apophysisslightly raised to gibbous, more pro-
dbh to 60-80(-100) cm. Trunkmonopodial, erect, nounced on one side of the cone toward the base,
straight,terete, taller trees with clear boles for 2/3or transversely keeled, rhombic to pentagonal in out-
more of total height. Bark moderatelythick, rough, line, upper marginssometimes crenate, radially stri-
scaly with irregular,longitudinalplates and shallow ate, lustrous ochraceous or light brown. Umbo dor-
fissures, reddish brown to greyish brown, on young sal, flat to slightly raised, 3-4 mm wide, with a
trees and branches thin, scaly, exfoliating in thin minute, deciduous prickle, concolorous or greyish
flakes. Branchesof firstorderrelativelylong, slender, brown. Seeds obovoid, slightly flattened, 3.5-6 x
spreading or ascending: branches of higher orders 2-3.5 mm, dark grey-brown. Seed wings articulate
slender, flexible, slightly pendulous, forming a but effective, held to the seed by two oblique claws,
rounded, rather open crown. Shoots smooth, with obliquely ovate-oblong, 13-17 X 6-8 mm, yellow-
faint ridges between weakly developed pulvini which ish, translucent,with grey or black tinge. Numberof
disintegrateearly in the second or thirdyear, orange- cotyledons not observed.
brown, later [Link], subulate, Distribution and ecology (Map 21). Mexico: Jal-
recurving at apex, scarious, erose at margins, light isco, in the NW partof the SierraMadredel Sur, on
brown with hyaline margins,early abscised. Vegeta- the Pacific slope, mainly in the Sierra de Cuale
tive buds ovoid-oblong to conical, the terminalbuds (Sierrael Tuito). It occurs at 800-1200(-1650) m in
10-15 mm long, the laterals ovoid-acute, smaller, mountainous terrain on deep, acidic soils derived
not resinous; the scales imbricate,free and reflexed from granitic rocks. The climate is subtropical,with
at apex, subulate, light brown with ciliate margins. a 5-6-month dry season from December to May and
Fascicle sheaths of juvenile leaves up to 15 mm annual precipitation of 1000-1500 mm depending

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Systematic Treatment 165

F
D

D A

:~~~~

FIG. 40. Pintusjaliscana (A. B, D, Wilkinson& StYles 1; C. E-G, Perez de la Rosa 348; H, Perez de la Rosa 354).
A. Branchwith leaf fascicles and immatureovuliferous cone. B. Leaf fascicle. C. Cross section of leaf. D, E. Ovuliferous
cones. F. Seed scale (three views). G. Seed. H. Pollen cones. (Magnifications:A, B, D-H, x0.5; C. x40.)

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166 Flora Neotropica

on altitudeand exposition. Pinusjaliscana is a local kind and seem to serve no other purpose than to fix
constituentof pine or pine-oak forests with a limited anticipated [Link] comparison of the
extent in most localities known so [Link] above cited specimens with material studied of P
and widespread species that grow with it are P herrerae,we presentthe following summaryof char-
maximinoi and P oocarpa, and at slightly higher acter states:
altitudes also P douglasiana. Several species of
Pinus herrer(e Pinus jaliscana
Quercus are often co-dominant; Clusia salvinii is
also common. Phenology: The time of pollen dis- leavesin fasciclesof 3 leavesin fasciclesof
persal is not recorded;the occurrenceof (old) pollen (4-)5, rarely3
cones on a specimen collected in November (Perez leaves0.7-0.9 mmwide leaves(0.5-)0.6-0.8 mm
de la Rosa 354) cannot be taken as evidence for a wide
late (second) period of dispersal. resinductsinternal, resinductsseptal,some-
rarelyseptal times 1-2 internal
Representative specimens examined. MEXICO. JAL- matureseedcones(2-) matureseedcones(4.5-)
ISCO:[Link] ca. 4 kmN of ElTuitoto Zimapan 3-3.5(-4) cm long 6-8.5(-9.8) cm long
mine, 7 km from Hwy. 200, acrosscreekon hillside,3
May1994,Farjon& Mejia340 (E, [Link],K, MEXU, In additionto size, which from these figuresseems
U); El Tuito,nearZimapanmine, 16 Nov 1983,Perezde to be distinct, the cones of P jaliscana often remain
la Rosa348 (FHO);El Tuito,17 [Link] Zimapan
closed in the lower '/ or so, resembling those of P
mine, 16 Nov 1983,Perezde la Rosa354 (FHO,IBUG);
3 km N of RanchoEl Saucillo,[Link] SanSebastian, patula.
II Sep 1983,Perezde la Rosa386 (IBUG);La Estancia,
25 km alongrd. to San Sebastian,betweenBalconesand
24. Pinus tecunumanii Eguiluz & J. P. Perry,Revista
La Bulera,20 Apr [Link] la Rosa 1089 (IBUG);
along rd. to Los Lobos,5 km N of Cuale,21 Apr 1987, Ci. Forest. 8: 4. 1983. Fig. 41
Perez de la Rosa 1352 (IBUG); Milpillas, 17 May 1986,
RevnaB. 225 (IBUG);ElTuito,[Link] Zimapan Pinus patula Schiede ex Schlechtendal& Chamisso
mine.20 Sep 1986,Wilkinson& StylesI (FHO). subsp. tecunumanii (Eguiluz & J. P. Perry) Styles,
[Link].13:50. 1984.
Uses. No particularpreferenceto single this spe- [Link]:San Jeronimo,"col-
lectedat finca INAFOR,60 m (NE) down fromthe
cies out as a source of timber is known to occur; it towerof GUATEL," Eguiluz2 (holotype,GH).
is locally exploited with other pines that grow with Pinus oocarpa Schiede ex [Link]
it and are more abundant. Martinez,[Link]. [Link]: 65.
1940 ("ochoterenai"). Type. Mexico. Chiapas:"Las
On a field trip to the Sierra de Cuale in western Casas"[San Crist6balde las Casasl, Jun 1939. M.
Jalisco, P6rez de la Rosa (1983) initially identified Martineo-3461 (lectotype,[Link]).
an uncommon pine there as P herrerae; it was,
however, later described as a new species. Pinus Tree, usually tall, height to 50-55 m, dbh to 120-
herrerae does occur in Jalisco, with leaves in fasci- 140 cm. Trunk monopodial, erect, straight, terete,
cles of 3(-5?) and somewhat largercones than else- bole of large trees usually clear of branchesfor 20-
where. The septal resin ducts in the leaves of the 30 m. Bark on lower part of the trunk thick (ca. 5
new taxon appeared to be a more fixed character cm), grey-brown, above 3-4 m thinning, reddish
distinguishing it. brown,on young trees and branchesthin, exfoliating
As Carvajal and McVaugh (1992) pointed out, in thin flakes, light reddish brown. Branches long,
none of the differences between the species, which slender, spreadingor slightly ascending, the higher-
neither P6rez de la Rosa (1983) nor Carvajal(1986) order branches flexible but not pendulous. Shoots
compareddirectly with each other in their tables or rough, with persistent bases of decurrent pulvini,
otherwise, are absolute. However, there is probably reddishbrown,often glaucous. Cataphyllsspreading
additional material collected from Jalisco and filed or recurved after the leaves have fallen, subulate,
in herbaria under P herrerae which belongs to P erose-ciliate at margins,scarious, brown, weathering
jaliscana. A more detailed study of this materialand to blackish grey. Vegetativebuds oval-oblong to cy-
visits to possible localities may reveal not only a lindrical,the terminalbud 15-20 mm long, the later-
greater range of the latter than presently known, als smaller,not resinous;the scales spreading,subu-
but also provide a better understandingof species late, ciliate at [Link] sheaths initially long,
delimitation and relationshipsbetween this and re- up to 25 mm, (light) orange-brown,the imbricate
lated taxa. The new combinationsproposedby Silba scales with ciliate, whitish margins,in maturefasci-
(1985b, 1990) are scarcely based on researchof any cles reducedto 12-18 mm, persistent,tight, weather-

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Systematic Treatment 167

~~J
.

! ~~
I*W? ~ ~ ~ ~ . "?

Dm4
I~~~~~~~~~~~~~I
. ~ ..~ . ~ . ~ ~ ~.

am4~~~~~~~~~

i$s I '.$
Map21~Piw aicn tinls) itwii(ice)

ing slowly, grey-brown, sometimes blackish brown. cones ovoid to broadly ovoid, with acutish apex
Leaves in fascicles of 4(3-5), in lax, spreadingtufts, when closed, nearly symmetrical or asymmetrical,
persisting2-3 years, droopingbut not pendent,(14-) with a roundedbase in open cones, (3.5-)4-7(-7.5)
16-18(-25) cm X 0.7-1(-1.3) mm, serrulateat mar- X (3-)3.5-6 cm when open. Seed scales ca. I(X)-
gins, acute, brightgreen. Stomata on all faces of the 140, partingusually within 1-2 years after maturity,
leaves, in (3-)5-8 lines on the convex abaxial face oblong, straight or slightly curved, the proximal
and in 2-5 lines on each adaxial face. Leaf anatomy: scales remainingconnate, purplishbrown to reddish
Triangularto transverse-triangularin cross section; brown, with light brown marksof seed wings adaxi-
hypodermis with a double layer of cells except in ally. Apoph,sis raised, transverselykeeled, on proxi-
marginalareas where it is thicker;resin ducts (2-)3- mal scales more or less gibbous, rhombic to obtrul-
4(-5), medial, occasionally I internal, or septal by late in outline, or with a rounded,but crenate, upper
connection with additional hypodermal cells; stele margin,striate, dull light brown or slightly lustrous.
transverse-obovatein cross section; cells of the endo- Umbo dorsal, flat or slightly raised, 3-5 mm wide,
dermis thin-walled or rarely slightly thickened;vas- with a minute, deciduous prickle, grey. Seeds
cular bundles 2, the xylem strands connate. Pollen obliquely ovoid, slightly flattened,4-7 x 2-4 mm,
cones crowdednearthe proximalend of a new shoot, dark grey-brown, with blackish dots, or blackish
spreading, subtended by scarious bracts, ovoid-ob- grey. Seed wings articulate, effective, held to the
long to cylindrical, 15-20 X 5-6 mm when shedding seed by two thin claws, obliquely ovate, 10-13 x
pollen, yellowish, turning yellowish brown. Seed 4-8 mm, grey-brown. Cotyledons (4-)5-6(-7), of
cones subterminal,in whorls of 2-4, rarely solitary, equal length.
on (15-)20-25 mm long peduncles, semi-serotinous, Distribution and ecology (Map 22). Essentially
persisting 1-3 years after shedding seeds, falling Mesoamerican,distributedthroughoutlow- to high-
with the peduncles. Immaturecones broadly ovoid, montane regions of Chiapas (Mexico), Guatemala,
with acutish apex, on long, recurvedpeduncles with Belize, Honduras,El Salvador,and [Link]
persistentcataphylls,purplish,soon light brown, ma- oocarpa [Link](here treatedas a synonym)
turing in two seasons (or equivalent time). Mature was reported from Nicaragua into Oaxaca; it was

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168 Flora Neotropica

!B I G

FIG. 41. Pinus teclnumnanii(A, C. E, F, H. Grijalva & Sediles 5330; D, Styles 115; G, I. J. M. Martinez 3564). A.
Branch with leaf fascicles. B. Leaf fascicle. C. Cross section of leaf. D. Flushing shoot with two immatureovuliferous
cones. E-G. Ovuliferous cones. H, I. Seed scales (various views). J. Seed with wing. (Magnifications:A, B, D-J. x 0.5;
C., x30.)

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SystematicTreatment 169

i i.i. .

I~~~~~~~? Srr . ? ??? 1?.

L~~~~~~~~~~~~~
" ~ ~ ~ ~ ~~a 2 Pnslcnmni

also said to occur in SE Guerrero(Perry, 1991). The pas, and may be earlier at lower altitudes in tropical
altitudinalrangeof this species is considerable:(300-) regions.
550-2500(-2900) m. In Belize it is found at 300-
760 m. It is a major constituent of more or less Representative specimens examined. MEXICO. CHIA-
PAS:El Chilil, 13 Mar 1993, [Link] & Styles 20
open to closed-canopy pine and pine-oak forests in (EAP, FHO, HEH, MO); Jitotol, 14 Mar 1993, Higinan.
climatic zones that receive annualrainfallof at least Padilla & Styles 25 (EAP. FHO, HEH. MO): "Las Casas"
I(X)Omm, and up to 2500-3000 mm in some places. [San Crist6bal de las Casasj. Nov 1939. M. Martinez s.n.
The dry season is usually long, lasting from Novem- (MEXU). Apr 1939, Martinez 3564 (MEXU); near San
Crist6balde las Casas, 3 km along rd. to Ocosingo. 6 Oct
ber to May, so that at lower to middle altitudes fires 1983, McCarter & Hughes 58 (FHO): near San Crist6bal
are an integral phenomenon in the ecosystem (the de las Casas. 22 Jul 1977, Mittak 8519 (FHO. M): 2 km
frequency,however, having been greatly accelerated along rd. from Ocosingo to Comitin. 31 Mar 1974. Styles
119 (FHO). OAXACA: Yavezia. Dec 1842. Liebnmann 14792
by humans). Here, open pine stands with grasses,
Pteridium aquilinum, Rubus, Calliandra, and Leu- (F, GH, UC. US): San Bartolo. Dec 1842. Liebmamn14794
(F. US); 15 km from Juquila along rd. to Hwy. 113. Sierra
caena are predominantas long as the disturbances de Miahuatlan,25 Feb 1985. McCarter549 (FHO).
do not lead to [Link] less disturbed GUATEMALA. BAJAVERAPAZ: Cubulco. Sep 1870.
areas, mostly at higher altitudes, P. tecunumanii is Bernoulli& Cario 1079 (GOET);Sierrade Chuacus."Finca
often associatedwith other pines, such as P. oocarpa, Nacional San Jer6nimo."19 Feb 1983, Hughes 302 (FHO);
Rabinol.28 Jun 1951. Schwerdtfeger42 (M). CHIQUIMULA:
P. maximinoi, P. pseudostrobus, and, at the more San Jose La Arada, 30 Sep 1976, Mittak 8356 (M). EL
mesic sites, P ayacahuite and P strobus var. chia- PROGRESSO: Bucaral,W slopes of Sierrade Las Minas, 28
pensis. Abies guateinalensisand Cupressuslusitanica Mar 1972, Styles 149 (FHO). EL QUICHE:Chichi-
are other conifers on these high mountainridges. On castenango. 4 Jul 1951, Schwerdtfeger48 (M). GUATE-
MALA:E of San Jose Pinula. mtns. above Finca Agua Tibia.
the Atlantic slopes in Chiapas, a mixed angiosperm
7 May 1979, Stead 272. 273 (FHO), 21 Mar 1979. Stead &
forest with Liquidambar,Magnolia, Clethra, Carpi- Styles 270 (FHO). HUEHUETENANGO: Along rd. 4K from
nus, Symplocos, Quercus, and many other species Huehuetenango.22 Feb 1938, Schrenk 7 (A, MO): Sierra
predominates;and Pinus tecunumanaii and other pines Los Cuchumatanes, 6 Jul 1951, Schwerdtfeger52 (M).
occur either on poorer sites or at an earlier stage in TOTONICAPAN: Momostenango, 11 Mar 1993, Higinan.
Padilla & Styles 8 (EAP, FHO. HEH, MO); Pachoc, 12
a sere leading back to dominance of broad-leaved Nov 1976. Mittak & Perry 8370 (FHO). ZACAPA: San
trees. Phenology: Pollen dispersal occurs in Febru- Lorenzo, 27 Feb 1980. Mittak 9134 (BANS. M).
ary-March in the highlandsof Guatemalaand Chia- BELIZE. CAYO:Mountain Pine Ridge. Angel's Drive.

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170 Flora Neotropica

13 Apr 1983, McCarter 130 (FHO); Millionario, in centre (1984), studied the pines of Guatemalain connection
of Chiquibul Forest Reserve, 14 Apr 1983, McCarter 135 with a pathological outbreak of a bark beetle and
(FHO); Mountain Pine Ridge, Little Granite Basin, 1960, came to the conclusion that a species new to science
Little 17983 (IFGP,K, US).
HONDURAS. COMAYAGUA: Plan Cerro Maneadero, occurred there. He named it "Pinus tecunmumtaamii,"
10.5 km E of Lago Yojoa, 14 Mar 1993, Hawkins 630 misspelling the name while commemorating the
(EAP, FHO, MO, TEFH); 6 km SSE of Siguatepeque, 16th-centuryIndian leader Tecun Umin, but his de-
"bosqueESNACIFOR,"7 Oct 1983, Hernlidez. & Cornejo scription is invalid under the Code (Art. 36.1, see
s.n. (HEH); Cerro Granadilla,10 km E of Comayagua,29
Feb 1988, McCarter 556 (EAP, FHO). CORTES:Cerro
Greuteret al., 1994). Eguiluz and Perry(1983) later
Cantiles, 20 km W of San Pedro Sula, in Cusuco National described the species as new (no type specimen had
Park, 18 Mar 1995, D. Mejia 321 (MO); near visitor ctr., been indicated by Schwerdtfeger).using the correct
18 km W of San Pedro Sula, in Cusuco National Park, 21 orthographyand designatinga type specimen.
Mar 1993, D. Mejifa358 (EAP, FHO. HEH, MO, TEFH). Martinez(1940) described a new variety of Pinus
LA PAZ: 15 km W of La Paz, 7 Feb 1983. McCarter &
Styles 72 (FHO). LEMPIRA:El Moj6n. 14 km SW of oocarpa, var. ochoterenae, from Chiapas, Mexico.
Gracias, in Celaque National Park, 17 May 1992. D. Mejiat In the protologue, he did not designate a type speci-
108 (EAP, FHO. MO, TEFH); between electric plant and men but cited collections he had seen from three
Don Tomas campsite, in Celaque National Park, 19 Nov locations: Ocosingo, Coapilla, and Las Casas [San
1991, Hawkins 99 (EAP, FHO, MO, TEFH): Cerro del
Crist6bal de las Casas]. On some of the labels he
Cleto, S of El Cutal, 5 Mar 1988, McCarter 574 (EAP,
FHO, HEH). OLANCHO: La Chorrera,15 km NW of Cata- wrote "var. nov." and "tipo."Martinez3461 is here
camas, in Agalta National Park,4 Jun 1992, D. Mejia'160 designated as the lectotype. Martinez enumerated
(EAP, FHO, MO, TEFH); Montafa de Coyolito, 15 km NE five characters that differentiate var. ochoterenae
of San Esteban, 7 Feb 1984, McCarter226 (FHO). from P oocarpa. The first characterstate-leaves in
EL SALVADOR. CHALATENANGO: Los Esesmiles, E of
La Palma, 6 May 1959, Alien & van Severen 7325 (EAP, fascicles of (3-)4-5 (mostly 4 in Martinez 3461)-
F, GH, NY, US). SANTAANA:Cerro Monte Cristo, 16 km is intermediatebetween P oocarpa and P patula.
N of Metapan, in Cerro Monte Cristo National Park, 23 The medial resin ducts described by Martinezare a
Feb 1989, Hughes 1263 (FHO, K, LAGU), 27 Feb 1979, state rarely encountered in P oocarpa but are the
Perry SALI (BANS).
rule in P patula. The cones of Martinez'staxon have
NICARAGUA. ESTELI:Llana Almaciguers, 4 km S of
Rio Estanzuela, 29 Sep 1980, Gumtidnet al. 1261 (FHO, charactersof both P oocarpa and P patula but are,
HNMN, MO, NY); 10.6 km W of bridge at La Trinidad, on the whole, less persistent and serotinous than
along rd. from Hwy. I to San Nicolas, 31 Aug 1978, either of these. Field studies carried out by Styles
Stevens 10270 (FHO, MO); Cerro Tomabu, 16 Oct 1979,
(1976) in the highlandsof Chiapaspointed to further
Steventset al. 14917 (FHO, MO, NY). JINOTEGA: 4.7 km
S of Jinotegaon old rd. to Hwy. 3 near Matagalpa,25 Aug differences with "typical"P oocarpa, which is sym-
1978, Stevens 10177 (FHO, MEXU, MO, NY). MATA- patric in this region.
(ALPA:Yucul, 20 May 1987, Grijalva 5097, 5098 (FHO); In their paper describing P tectunutanaii,Eguiluz
La Riconada, ca. 12 km SW of Dario, 24 Jul 1987, Gri- and Perry (1983) presentan evaluation of compara-
jalva & Sediles 5297. 5303 (FHO), 21 Aug 1987, Gri- tive charactersof this species and P oocarpa var.
jalva & Sediles 5326, 5330 (FHO); Cerro de la Olla, El
Ocotal, 10 km N of Waswali, 20 Jun 1982, Hughes 203 ochoterenae, using morphologicalas well as chemi-
(FHO). NUEVASEGOVIA: El Rodeito, 3.5 km W of Dipilto, cal (terpenoids) characters or traits. These show
Jul 1977, Tomlin189 (FHO, MO, NY). slightly greater mean size for most characterstates
in P oocarpa var. ochoterenae. However, several of
Uses. While occasionally still abundant and of them indicated as highly significant-the sequence
very tall stature, this pine is now usually scattered cone size, seed scale size, seed wing size-are,
in small, disjunct populations and has been depleted in fact, fully [Link]
by overexploitation and forest clearing, especially at leaves only the size of the cones to be considered,
lower elevations, to such an extent that many of which, singularly, being much more variable than
these populations are now vulnerable to extinction their limited samples suggest, taxonomicallyis not a
(Dvorak & Donahue, 1992). This taxon has received very informativecharacter.
considerable interest from foresters as a species for Meanwhile, Barnes and Styles (1983), Styles and
potential plantation forestry to be introduced in tropi- Hughes (1983), and Styles (1984) had concluded that
cal countries. A comprehensive collection of seed and P teciunumaniishould be classified as a subspecies
specimens was carried out by the Oxford Forestry of P patula. In the opinion of the authors, the
Institute (OFI), as well as by other organizations, applicationof statistical methods supportedthe clas-
throughout its entire range. sification of this taxon as distinct from P oocarpa.
Schwerdtfeger (1953), as explained by Styles However,a complex sequenceof calculationsinvolv-

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Systematic Treatment 171

ing continuouscharacterstates is necessary to arrive for this taxon. They also maintained P oocarpa
(by their method) at identification,whereby a possi- var. ochoterenae, which in Perry's (1991: 188-189)
bility of error remains (McCarter & Birks, 1985; morphologicalcomparisonappearedto score 7 times
Birks & Barnes, 1985). They found such a high level towardP patula and only 3 times towardP oocarpa.
of "continuitiesin the variationpatterns"within the Finally,a molecularapproachto the problemusing
two taxa that "orthodox taxonomic methods . . . RAPD markers on bulked DNA samples of these
are largely unsatisfactory."The second study looked pines (Grattapigliaet al., 1992) pointed at an "unex-
primarilyat growth performancesof provenances(in pected distinctivenessbetween P. patula and P tecu-
part sampled from those used in the first study) in nunanii." Until more, and perhapsmore informative,
plantation trials, which indicated genetically based DNA charactershave been found, the debate is likely
differences between them. It is difficult, however, to to circle aroundthe interpretationof old data. Intro-
determine the significance of these outcomes for gression, assumed to play a role especially at lower
species delimitation, given the great geographical altitudes,with P oocarpa (Dvorak& Donahue, 1992)
and ecological ranges of the taxa involved. may complicate even this more direct genetic ap-
Eguiluz (1986) undertooka comparativephenetic [Link] these reasons, it seems prudentto recog-
and biochemical study to determine the taxonomic nize this taxon in its original rank of species until
relationshipsof [Link] two closest its relationshipwith P. oocarpa and P. patula and
related taxa, P oocarpa var. ochoterenae and P their respective varieties has been clarified, if it ever
patula [Link],as well as P maximinoi will be.
(but not P. oocarpa var. oocarpa or P patula var.
patula) were chosen for this evaluation of relation- 25. Pinus durangensis Martinez,Anales Inst. Biol.
ships. Apart from the apparent undersampling(15 Univ. Nac. Mexico 13: 23, figs. 1-4. 1942. Type.
trees from one locality for each of the two varieties Mexico. Durango:El Salto, Sep 1940, M. Martline
against more than 100 trees from five localities in 3477 (holotype, MEXU; isotype, ENCB). Fig. 42
Guatemalafor P tecunumanii),there is the familiar Pinus durcangensisMartinezforma quinquefoliataMarti-
misunderstandingamong forest geneticists turningto nez. Pinos Mexic..ed. 2, 232. 1948. [Link].
Chihuahua:Bocoyna,Feb 1942, M. Martinez3454
phenetics that a t-test performed on the means of
(holotype,MEXU;[Link]).
morphological measurementswith a "high signifi- Pinus martineziiE. Larsen,Madroio 17: 217. 1964;
cance level" is proof of taxonomic distinction at Pinus douglasiana Martinezvar. martinezii(E. Larsen)
species level. In reality,the characterstates found in 68: 49. [Link]-
Silba,Phytologia
the analysis are not only quite similar (e.g., a mean can: NearParacho,10 Jan 1960,Hinds & Ltrsen65
leaf length of 17.5 cm and 19.4 cm found for P ([Link];isotypes.A, [Link]).
tecunumaniiresp. P oocarpa var. ochoterenae) but Tree, usually tall, height to 35-40 m, dbh to 80-
also of a continuous nature,with overlappingstates 100 cm. Trunkmonopodial,erect, straight,in forest
in many trees. Furthermore,of the morphological stands with clear boles for '/ to /4 of total height.
charactersused, several are interdependent,such as Bark rough, scaly, breaking into large, irregular,
length of seed scale and seed wing, and cannot be elongated plates divided by longitudinal, shallow
"addedup" to argue differences or [Link] fissures, dark brown, weathering grey, on young
outcome of the cluster analysis, which placed P trees and branches rough and scaly, reddish brown.
tecunumnaiiat a greater"taxonomicdistance"from Branches of first order long, slender, the lower
the two varieties of P. oocarpa and P patula than branches usually curved downward; branches of
these are from each other, is for these reasons not higher orders spreading or ascending, ultimately
conclusive. more or less pendent. Shoots uninodal, rough with
In a detailed study of this taxon, Styles and decurrent,persistent pulvini, orange-brownor red-
McCarter (1988) clearly point out the difficulties dish brown, usually glaucous. Cataphylls large, ca.
with the identification,especially in distinguishingit 15 mm long, 3-4 mm wide at base, subulate, soon
from P oocarpa "at middle elevations (900-1600 m) reflexed, scarious, with erose-ciliate margins and
on sites where the two species occur together."Yet caudateapex, darkbrown. Vegetativebuds (broadly)
they maintainStyles's classification of the taxon as ovoid, the terminalbuds 15-20(-25) X 10-15 mm,
P patula subsp. tecunumanii,but agree that its status the laterals smaller, not resinous; the scales free,
is likely to remain controversial. Indeed, Squillace spreadingto recurvedat the base of the bud, subu-
and Perry (1992), in a paper using mainly terpene late-linear,orange-brownwith dark, ciliate margins
compounds as characters,maintainedspecific status and a caudate apex. Fascicle sheaths long, 20-30

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172 Flora Neotropica

"./N :

6lir i.

1f1
G

FIG. 42. Pinus durangensis (A, Spellenberg et ul. 10063; B, C, Hinds & Larsen s.n. ("paratype");D, Styles. Fave/a
et al 10;E.
E Pirez de la Rosa 528). A. Branch with leaf fascicles. B. Leaf fascicles. C. Cross section of leaf. D, E.
Ovuliferous cones. F. Seed scale (three views). G. Seeds. (Magnifications:A, B, D-G, x 0.5; C, x40.)

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SystematicTreatment 173

mm, persistent but reduced to 10-15 mm with a in Jalisco). Critchfieldand Little (1966b) mappedP
thickened base in older fascicles, with ca. 10 imbri- durangensis in N Michoacan but did not mention
cate, appressed, scarious, orange-brownscales with Larsen'staxon as a synonym. In the Sierra Madreit
ciliate margins, weathering grey-brown. Leaves in is an important constituent of the "yellow pine"
fascicles of (4-)5-6(-7, rarely8), in dense, spreading forest, where it occurs in pure stands or mixed with
or drooping tufts, persisting 2-2.5(-3?) years, several other species of pine-e.g., P arizonica, P
straightor slightly curved, 14-24 cm x 0.7-1.1 mm, leioph!lla, and P engelmanii--or in pine-oak for-
marginsserrulate,acute to pungent,yellowish green ests. Its altitudinalrange is extensive: (1400-)1600-
to glaucous-green. Stomata on all faces of leaves, 2800(-3000?) m, but most common at 2000-2800
with 3-4 lines on the convex abaxial face and 3-4 m. Climatically,the SierraMadreis warm-temperate,
on each adaxial face. Leaf anatomy: Cross section but with cold spells during the short winter at the
triangular;hypodermismulti-layered,slightly intrud- [Link] in the range
ing into the mesophyll;resin ducts (2-)3(-5) medial, of 700-1200 mm, most of which occurs in the
the three largerones near the margins;stele more or [Link] pine is adapted to grow on shallow,
less terete;outer cell walls of endodermisthickened; rocky soils, but its better stands are found on deeper
vascularbundles2, connatebut [Link] cones soils, where it can successfully compete with most
crowded near the proximal end of new shoots, sub- other pines. The soils are derived mostly from volca-
tendedby subulatebractswith ciliate margins,ovoid- nic rock. At the highest elevation it occurs with
oblong to cylindrical,yellowish brown when mature. Abies and/or Cupressus lusitanica: at the lowest with
Seed cones subterminal,solitary,in pairsor in whorls Juniperus deppeana and Pilnus oocarpa. Other pines
of 3-4 on short, stout, usually persistentpeduncles, are P m,ontezumae, P teocote, and, in the southern
eventually deciduous. Immaturecones broadlyovoid partof its range, P [Link]: Pollen is
to subglobose, ca. 15 X 10-12 mm, with pungent dispersedduringApril-May.
umbos, purplishor glaucous, maturingin 2 seasons.
Maturecones ovoid or broadlyovoid when opened, [Link]-
Representative
HUAHUA: Cascada Basaseachic. II Nov 1986. Spellenberg
slightly flattened or obtuse-conical at base, often et al. 10063 (FHO, MEXU. NMC). Spe/llenberget. al
slightly curved, 5-9(- 11)x 4-6(-7) cm when open. 10065 (NY). DURANGO: SE of Guachichilas,80 km N of
Seed scales ca. 90-120, partingreadilyto release the Estaci6nCoyotes. 5 Nov 1970. Breedlove 18826 (MO): La
seeds except those at the base of the cone, thick Ciudad, 30 Sep 1960. Courhasson & Mahieux s.n. (P). 30
Apr 1987, Hughes 976 (FHO. MEXU); 5 km W of El
woody, broadlyoblong, straightor recurvednear the Salto,nearsawmillLa Lajita,30 Apr 1987.H,ughes980
base of the cone, purplishbrown to blackish brown ([Link]);MesaLagunitas. [Link] 1940.
abaxially, light brown with lighter marks of seed [Link].3583(F);[Link] [Link]-
wings [Link], sometimes flat on n;ez 3661 (NY); Mesa de Sandia. 15 Oct 1905, Pringle
the basal scales, prominently transversely keeled, 10140(A, GOET.K. M, [Link],NY.W); Durango,
5 Apr 1991, Styles. Favela et al. 10 (FHO). JAIISCO: Las
rhombicto pentagonalin outline, more or less sym-
Parejitas,7 km N of Ayotitlin.17 Apr [Link]
et
metrical around the cone, ochraceous to light (red- al. 6559 ([Link]);SierraManatlan.
CerroLas Ca-'
dish) brown. Umbo dorsal, raised, slightly recurved, pillas. on summit,16 Apr 1987. Hughes& Styles 138
transversely keeled, 5-6 mm wide, with a small, (FHO);23 kmS of Tenzompa at LosAmoles.21 Feb 1983.
[Link] la Rosa528 (FHO);CerroPrieto.15 Sep 1897.
persistent (or deciduous) prickle, usually grey in Rose2379 (A. MICH,US);"SierraMadreW of Bolanos."
mature cones. Seeds obliquely ovoid, slightly flat- 7-9 Sep 1897,Rose3004(NY).MICHOACAN: LlanoLargo.
tened, 5-6 X 4-4.5 mm, light brown to grey, with 1.5 km NE of Los Azufres,19 Mar [Link] al.
small, darkspots. Seed wings articulatebut effective, 919 (FHO);ca. 6 kmS of Paracho,10Jan [Link]&
Larsen s.n. (paratypeof P. nartinezii, FHO); Paracho. 9
held to the seed by two flimsy claws, oblique, 14-
May 1984. Perez de la Rosa 654 (FHO). SONORA: Sierra
20 X 6-9 mm, light greyish brown,translucent,with 17Jun 1953,Marshall121(UC).
Huachinera,
a blackishtinge. Numberof cotyledons not observed.
Distribution and ecology (Map 23). Mexico: Uses. Pinus durangensis,as a pine tree with a tall,
Mainly in the S Sierra Madre Occidental, rare in E straightbole, has been much sought after as a timber
Sonora and Chihuahua,common in Durango,Zaca- tree and exploitation has been severe, especially in
tecas, and N Jalisco, more scatteredfurthersouth in Durango (Perry, 1991), where extensive, more or
Jalisco and N Michoacin. Populationsin these latter less pure stands are now rare. In mixed forest and
two states have been classified as a distinct species, remote areas, matureto aged trees still occur abun-
Pinus nmartinezii (Larsen, 1964; see also Perry, 1991, dantly,but in the near futureprotectionmay become
who reportedthis taxon from the Sierrade Manatlin necessary to avoid over exploitation.

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174 Flora Neotropica

S? *.-,. . .. -

-4. .Ji. .

i.~ ~~~~~~~~~~a 23 sus svclgns

As Martinez(1942) has pointed out, Shaw (1909) form with 5 leaves (rarely 6) per fascicle was de-
cited under P. montezumae a collection (Pringle scribed: P durarcgensisf. quinquefoliata, based on
10140, A, K, MEXU, NY, W) from the Mesa de la Martinez3454 (MEXU). No other distinctive charac-
Sandia, Durango, which he seems to have referred ters were mentioned or observed on the type speci-
to later(Shaw, 1914) as P ponderosa var. nacrophy- men. The variation in leaf numbers appears to be
lla (= P engelmtannii) and not var. arizonica as greater than Martinez believed, and we do not con-
Martinez understood it. These specimens have leaf sider this taxonomically significant.
fascicles of 5-6 (rarely 7) leaves. Other collections Larsen (1964) described P martinezii from Mi-
from Durango confirm its affinity with P arizonica choacan, which he distinguishedfrom P durangensis
and P engelmannii and its high number of leaves by its glaucous leaves 20-28 cm long (23 cm on
per fascicle, although Martinez(1942) thought it to average)and its deciduous, very small umbo prickles.
be more similar to P. montezumnae. Nevertheless, he The type cited-Hinds & Latrsen65, 10 Jan 1960-
recognized it as distinct and proceededto describe it matches specimen AFS 13084/A at FHO, which is
as a new species. In the protologue, no specimen indicatedas "s.n." on the sheet and "holotype"on a
was indicatedas the type. In MEXU, Martinez3477 field label. It is here consideredto be the holotype. In
bears his annotation "Pinus durangensis sp. nov. the protologue,CANB is indicatedas the herbarium
Tipo" and this specimen must be recognized as the where the type was to be kept, but it is not there;
holotype. the material was sent on "for safekeeping" to FHO
In his revised monographof the pines of Mexico and elsewhere. Due to some confusing remarks
(Martinez, 1948, which has a title changed from Las (Larsen, 1964) as to the relationshipof this species,
pinaceas [Link] Los Pinos mexicanos), a new the identity has been viewed in that light by Perry

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Systematic Treatment 175

(1991), who failed to refer to the type to solve the (6-)9-12 lines on the convex abaxial face and 4-5
problem. The morphological differences are minor lines on each adaxialface. Leafanatomy:Transverse-
and very inconsistent,e.g., glaucousness is not corre- triangularto triangularin cross section; hypodermis
lated with leaf length. Larsen'sspecies is here treated double- to multi-layered,often thickest in marginal
as a synonym of P durangensis, although some areas; resin ducts (1-)2-6, internal and medial,
reservationsaboutthis treatmentwere held by Carva- mostly internal;stele oval in cross section; cells of
jal and McVaugh(1992), in the probabilitythat more endodermis small, not or slightly thickened exter-
material would later become available than they nally; vascularbundles2, [Link]
had at their disposal. Stead and Styles (1984) have cones crowded near the proximal end of a new
searchedthe type locality without [Link] local- shoot, subtendedby scarious bracts,ovoid-oblong to
ity was revisited by A. Farjon in 1994, and it ap- cylindrical, 10-20 x 5-6 mm, yellowish green,
peared almost lumbered of mature pines. No trees turninglight brown. Seed cones subterminal,some-
attributableto this species could be found. times solitary, commonly opposite on short, to ca.
10 mm long, stout, curved peduncles which usually
26. Pinus lawsonii Roezl ex G. Gordon & Glendin- fall with the cone. Immaturecones ovoid, 10-15 mm
ning, Pinetum, Suppl. 64. 1862. Type. Mexico. B. long, purplish red, soon brown, maturing in two
Roezi s.n. (original materialnot located). Michoa- seasons. Mature cones narrowly ovoid to ovoid-
can: "Hillsides near Uruapdn,"9 Oct 1904, Pringle attenuatewhen closed, more or less asymmetrical,
10141 (neotype, NY, here designated;isoneotypes, ovoid when open, with a more or less flattenedbase,
A, BM, E, F, GOET, K, M, MO, S, US). Fig. 43 then 5-8(-9) x 4-6(-7) cm. Seed scales ca. 70-100,
Pinus altamiranoi G. R. Shaw in Sargent,Trees & Shrubs parting to release the seeds, thick woody, oblong,
1: 209, t. 99. 1905 ("Altrairani"). Type. Mexico. Mi- recurvedwhen fully spread,purplishblack abaxially,
9 Oct 1905,Pringle
choacan:"HillsidesnearUruapan." dark brown, with two light brown marks of seed
10017(lectotype.A, heredesignated; A,
isolectotypes.
BM,GOET,K, M, MICH,MO,P. US, W). wings on the adaxial side. Apophysisslightly raised,
transversely keeled, rhombic in outline, the upper
Tree, medium to tall, height to 25-30 m, dbh marginundulate-crenate,radiallystriate,light brown
to 75 cm. Trunk monopodial, straight, sometimes to grey-brown. Umbo dorsal, pyramidal, curved,
tortuous, terete. Bark thick, rough and scaly, with transverse-rhombicin outline, 3-4 x 4-5 mm, lack-
broadand deep longitudinalfissures,outer barkdark ing a distinct prickle, grey-brown. Seeds obovoid,
blackish brown, innerbarkdarkpurplishred (slash), slightly flattened, 4-5 mm long, dark brown. Seed
on young trees and branchesthin, exfoliating in thin wings articulate,functional,held to the seed by two
plates, reddish brown. Branches of first order long, oblique claws, 12-16 x 5-6 mm, translucent,light
spreading horizontally,assurgent in the higher part brown with a dark tinge. Numberof cotyledons not
of the crown;branchesof higherordersstout, flexible observed.
but not pendulous, forming a broadly domed but Distribution and ecology (Map 21). Mexico:
open and irregular crown. Shoots smooth, ridged Ratherscatteredin S Mexico, from the Eje Volcanico
between dwarf shoots (fascicles), with weakly devel- Transversal south into the highlands of Oaxaca;
oped pulvini, orange-brown,often glaucous. Cata- found in Michoacan,Mexico, Morelos, DistritoFed-
phylls subulate-lanceolate,ca. 10 mm long, scarious, eral, (one locality in) Veracruz,Guerrero,and Oa-
erose-ciliate at margins,dull brown, soon deciduous. xaca. Its altitudinalrange is 1300-2600 m. It is a
Vegetativebuds ovoid-oblong to cylindrical, the ter- constituentof warm-temperateto temperatemontane
minal bud 10-15 mm long, the lateralsshorter,acute, forest or woodland, with annual precipitation600-
not resinous; the scales imbricate,straight,subulate, 1500 mm and a dry season from November to May.
ciliate at margins. Fascicle sheaths of young leaves It is most commonly found in pine-oak forest, also
long, up to 25 mm, the imbricate scales lustrous in pine forest with, e.g., Pinus pringlei, P patula, P
orange-brown,persistentbut reduced to 10-15 mm nontezumrae, P. oocarpa, P leiophllla, P herrerae,
in older fascicles, darkred-brown,weatheringblack- P. teocote, and P [Link] sites with sandy,
ish or grey-brown. Leaves in fascicles of 3-4(-5), shallow soil, Juniperus can be co-dominant. Phe-
very seldom 2, in dense or open tufts, persisting 2- nology: Pollen dispersaloccurs November-February
3 years, straightor nearly straight,directed forward (observed on specimens with pollen cones).
to spreading wide, rigid, 12-20(-25) cm x 1-1.2
[Link].
(-1.5) mm, serrulateat margins,acute-pungent,glau- TRIrO FEDERAL:Eslava, May 1939, M. Martinez 3426
cous-green. Stomata on all faces of the leaves, in (MEXU,US). Guerrero:Alongrd. fromFilo de Caballo

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176 Flora Neotropica

I~ ~ ' ~ ~~~
%..
I;' ''
I~~~~, : -.! / ii--.~:i

,.. i,-,,,.

'~
r? ' I"
\,
,?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
\?~~~%~

;"~~~~~~~~~~~~~~r

0 " ' '

4~~~~~~~~~~~~~~~~~~~~:
'~~~~~~~~~~~
,

t
OR~~~~
C~~~~~

~,-,~~y~:,,~..

FI. [Link], i A-. itan& r enintoa 953 n-F Sylsh 5) A w l ef acce n

[Link] wns (Mgiiain:^.BD-.x.5C.x5.

FIG. 43. Pinu~slavvs'.onii (A<. Littmaunn & Pennwington 9053; D-F. Styles IS). A. Branch with leaf fascicles and
immatureovuliferous cone. B. Leaf fascicles. C. Cross section of leaf. D. Ovuliferous cones. E. Seed scale (two views).
F. Seeds, with articulatewings. (Magnifications:A, B, OF, x0.5; C. x35.)

to La Laguna. 8 Feb 1981. Stead et al. 735 (FHO). Ml- (MICH):Agua Blanca, 23 Feb 1974, Styles 15 (FHO); 5 km
CHOACAN:Along rd. from Zitacuaro to Morelia, 69 km from Uruapan,6 Mar 1974, Styles 46 (FHO). MORELOS:
from Zitacuaro,21 Jul 1967, Littmnant& Pennington9053 Cuernavaca,Oct 1904, Shaw s.n. (A). OAXACA: Las Sedas,
(FHO. NY): 28 km E of Morelia. between Rio del Salto n.d., Rose & Hough 4639 (A, NY, US); Nejapa, 17 km E
and La Polvilla. 9 Nov 1961, King & Soderstrom5116 of bridge across Rio Tehuantepecalong Hwy. 190, 27 Dec
(MEXU. MICH, US); Huingo, 24 Nov 1907, Pringle 10399 1962, Tavlor & Tavlor 1183 (MICH); 148 km SE of
(ENCB. K, US); Coru. 29 Oct 1905. Pringle 13711 Oaxaca, 20 Jul 1977, Mittak 8514 (BANS, FHO); 18 km

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SystematicTreatment 177

SE of Miahuatlan,16 Apr 1965, Rzedowski19659 (ENCB). barkorange-red,outer barkgreyish brown,on young

VERACRUZ: Cofrede Perote,10 kmSE of Perote,28 Mar trees and branchesthin, scaly, exfoliating abundantly,
1963,Little18972(INIF,K, US). reddishbrown. Branches long, slender,spreadingor
Uses. Along with other pines, which are often arching, assurgent near the top, the highest-order
more abundantin the mixed pine or pine-oakforests, branches stout, assurgentor tortuous. Crown open,
this species is cut commerciallyfor lumber,although broadly domed or irregular, with sparse main
most trees are of medium size and the boles some- [Link] thick, smooth, prominentlyridged
times [Link] several places with abundanttrees between dwarf shoot bases, with decurrent, soon
of this species, resin may be tappedas well. erodingpulvini, reddishbrown,glaucous. Cataph!lls
Gordon and Glendinning (1862) ascribed Pinus lanceolate to subulate, 10-12 mm long, scarious,
lawsonii to B. Roezl, but the name does not appear becoming reflexed at apex, erose-ciliate at margins,
in either of the two catalogues of Mexican pines reddish brown, deciduous. Vegetativebuds ovoid-
(Roezl, 1857, 1858). Carriere(1867), in evaluating oblong to cylindrical, the terminal bud 10-15 mm
Roezl's species, refers to Gordon's description, in long, the lateralsovoid-acute, smaller, not resinous;
which there is no referenceto either a publicationor the scales imbricate,subulate,straight,ciliate at mar-
a specimen. Roezl coined several additional names gins, reddish brown. Fascicle sheaths of young leaf
on labels that went with the seeds and cones/shoots fascicles up to 20 mm long, the imbricate scales
sent to Europe which were not published by him; it chartaceous,with ciliate margins,red-brown,persis-
is probable that Gordon and Glendinning received tent but reducedto 10-15 mm length and with dark
such material,but that it has not been [Link] red-brown to blackish colour on mature fascicles.
Roezl's collections at Fl and P, nothing has been Leaves in fascicles of 3(-4), in dense, ascending,
found with this name, incidentally one of the very rigid tuftson upturnedbranches,persisting2-3 years,
few Roezl names that representa good species. For rigid, straight, (15-)18-25(-30) cm x 1-1.5(-1.7)
this reason, a neotype has been selected; it seems mm, with serrulatemargins,acute-pungent,lustrous
appropriateto choose the collection C. G. Pringle (yellowish) green or [Link] all
10141 under this name, in view of Shaw's (1909) faces of the leaves, in 9-13 lines on the convex
treatmentand the wide distributionof duplicates. abaxialface and 4-5 lines on each adaxial face. Leaf
Shaw (1905c) described Pilnusaltamiranoi from anatomy: Cross section transverse-triangular,with
near Uruapanin Michoacan,based on C. G. Pringle a convex abaxial side; hypodermis multi-layered,
10017, F Altanirano s.n., and G. R. Shaw s.n. of especially in the marginalareas, with intrusionsinto
October 1904. The collection by Pringle has been the mesophyll; resin ducts 3-7, internal,sometimes
1-2 medial or septal if connected by hypodermal
designatedas the lectotype as it is also cited in Shaw
(1909) underP. Iawsonii and duplicateswere widely cells; stele large, triangularin cross section; cells of
distributed. Shaw synonymized his P altamiranoi the endodermis with uniform thin walls; vascular
there without further discussion-a conclusion ac- bundles 2, separate or the xylem strands connate.
Pollen cones crowded near the proximal end of a
cepted by Martinez(1948) and all subsequentauthors
on Mexican pines. The maximum leaf length of 30 new shoot, subtended by broad, scarious. brown
cm mentioned in the protologue of P altamiranoi bracts,ovoid-oblong to cylindrical, 15-25 X 8 mm
has not been found in the original material, nor in when shedding pollen, yellowish, turning brown.
Seed cones subterminal,solitary or more commonly
any other collections of P Irawsoniiknown to us.
Shaw (1909) later correctedthis himself by giving a in whorlsof 2-4 on short,stout,curvedand tenacious
maximumlength of 24 cm for the species including peduncles,maturecones seemingly sessile. Immature
its synonym. cones ovoid, 10-15 mm long, purplish,maturingin
two years. Maturecones (narrowly)ovoid, slightly
27. Pinus pringlei G. R. Shaw in Sargent,Trees & curvedor asymmetrical,cuneate at base when closed
Shrubs 1: 211, t. 100. 1905. Type. Mexico. Michoa- (some cones are more oblique), 5-8(-10) x 3.5-6
can: Distr. Paracho, "hilltops near Uruapdn,"14 (-7) cm when open, but semi-serotinous(only par-
Oct 1904, Pringle 10019 (lectotype, GH, here des- tially opening), leaving some basal scales on the
branchwhen falling. Seed scales ca. 70-100, parting
ignated;isolectotypes, BM, CAS, E, GOET, K, M,
after a considerabletime except the proximalscales,
MEXU, MO, NY, P, S, TEX). Fig. 44
thick woody, oblong, almost straight, symmetrical.
Tree,small to medium, height to 20-25 m, dbh to purplishblack on the abaxial side, darkbrown, with
90-100 cm. Trunkmonopodial,straight,terete. Bark light brown marksof seed wings [Link]-
thick, rough, with small, scaly plates, fissured,inner sis on one side of the cone nearly flat to slightly

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178 Flora Neotropica

F B 11

~-e
E ~~5]9~~~8'

FIG. 44. Pinus pringlei (A-C, Styles 43; D, F, G, E. Matuda et al. 31041; E. Mexia 9015). A. Branch with leaf
fascicles and immatureovuliverous cones. B. Leaf fascicles. C. Cross section of leaf. D. Branch with ovuliferous cones.
E. Ovuliferouscone. F. Seed scale (three views). G. Seed. (Magnifications:A, B, D-G, X0.5; C. x30.)

raised, on the other (outer) side slightly more pro- brown. Umbo dorsal, flat or depressed, up to 5-6
nounced, these transverselykeeled, rhombic to pen- mm wide but often smaller,with a minute,deciduous
tagonal in outline, lustrous ochraceous or light prickle, blackish to grey. Seeds obovoid, slightly

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Systematic Treatment 179

-~~~~~~~~~~~~~~~~~n,
?? ...:
',?? ~~~~~~~~~~~~~~~~~~~~~.
,ow~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
",~?? ..Fr ?e . 'e
-.r. .......r

?, ~~~~~
pringlei.. Map 24. Pinus
'~~~~'

M 2.
a p~~
Piu prngei

flattened,4-6 mm long, dark brown to grey-black. Chilpancingo, 24 Mar 1993. Higmnan,Padilla & StYles55.
Seed wintgsarticulatebut functional,held to the seed 56 (EAP, FHO, HEH. MO); Sierra Madre del Sur. Distr.
Mina, Petlacala, 1937, Mex.r 9015 (B, DS, F, GH, K, MO.
by two oblique claws nearly enveloping the seed on NY, S, UC); near Chilpancingo, on NE slopes of Sierra
one side, thickened toward the base, 14-18 x 6-8 Madredel Sur, n.d., Nelson 2182 (A. US). Mexico: Valle
mm, translucent,light brown. Numberof cotyledons Bravo, 23 Nov 1952, Matuda 27543 (INIF); near Sultepec,
not observed. Las Mesas, 21 May 1954, Matuda et tl. 31038 (FHO,
Distribution and ecology (Map 24). Mexico: In MEXU). MICHOACAN: Nanchitilla, 26 May 1954, Matuda
et al. 31041 (FHO); near Uruapan, 1903, Shaw s.n. (E, K.
Michoacan,Mexico, Morelos, Guerrero,and Oaxaca,
MICH); 17 km from Ciudad Hidalgo. along rd. to Mil
perhaps in W Puebla. It is a pine of the montane, Cumbres, 9 Feb 1980, Stead & Styles 465 (FHO): along
mesic forests of the Sierra Madre del Sur and parts rd. from Uruapanto Carapan, I Feb 1980. Stead & StYles
of the Eje VolcanicoTransversal,where it is growing 470 (FHO); near Uruapan,Cerro de la Cruz. 25 Feb 1980.
at 1500-2600(-2800) m. This region has a subtropi- Stead & Styles 530 (FHO. MEXU); Las Playitas Forest
Station. W of Tumbiscatio, 28 Feb 1974, Stiles 27, 28
cal to warm-temperateclimate; variable annual pre-
(FHO); 5 km from Uruapan,6 Mar 1974, Stiles 43 (FHO.
cipitation of 1000-2000 mm occurs in summer K). MORELOS:Mountainside above Cuernavaca. 30 Jul
storms from June [Link] is a constit- 1906, Pringle 10340 (ENCB. LL, MICH. SD. TEX. US).
uent of pine and pine-oak forests; associated pines OAXACA:Llano Verde, 72 km from Oaxaca along rd. to
include Pinus douglasiana, P. maxininoi, P. pseu- Huajuapan.4 Mar 1985, McCarter & Styles 340, 341, 342
(FHO); Llano Verde, 65 km from Oaxaca along rd. to
dostrobus,P oocarpa, and P patula. On drier sites, Huajuapan,4 Mar 1985, McCarter & Styles 345. 346, 347
often in degraded or secondary forest, P lawsonii (FHO); ca. 100 km N of Oaxaca. 19 Jul 1977, Mittak8508
and P devoniana are more commonly growing with (FHO. M).
it; in both forest types Quercus spp. are often (co-)
[Link] Donahue(1992) reporta grass Uses. This species is reported to have dense wood
stage of the seedlings of P pringlei. Phenology: and is used for lumber; the resin is also tapped
Pollen dispersal has been observed in February-
commercially. The species is under local pressure by
March. lumbermen, which poses a threat to smaller popula-
Representative specimens examined. MEXICO. tions (Dvorak & Donahue, 1992).
GUERRERO: Near Mezones, along rd. from Chilapa to Shaw (1905b) described this new species from

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180 Flora Neotropica

near Uruapanin Michoacan,and cited C. G. Pringle reddish brown, outer bark dark greyish brown; on
10019, and collections by F. Altamiranoand himself, young trees and branches thin, scaly, exfoliating in
collected there in 1903 and in October 1904. In his thin flakes, reddish brown. Branches of first order
description he notes absence of "bloom"(glaucous- slender, spreading horizontally or curved down;
ness) on the shoots as a possible diagnostic feature, branches of higher orders slender, flexible, slightly
but several later collections are definitely glaucous pendulous, forming an open or dense, pyramidal
(e.g., McCarter & St'les 342 and Sntles 27). The to roundedcrown. Shoots smooth, with prominent,
rigid, thick leaves in fascicles of 3(-4) on very stout decurrent pulvini, orange-brown. Cataphylls subu-
branches and the semi-serotinous cones with only late, curved at apex, with ciliate margins, brown,
slightly raisedapophysesare a good charactercombi- early deciduous. Vegetativebuds ovoid-oblong, the
nation to distinguish this species. Pizns lawsonii, terminal bud 10-15 mm long, the laterals ovoid-
which occurs commonly with it, has leaves in fasci- acute, <10 mm long, not resinous; the scales imbri-
cles of 3-4(-5) which are more slender; the cones cate, subulate,with reflexed apex, ciliate at margins.
are smaller with more protuberantapophyses and Fascicle sheaths initially up to 20 mm long, silvery-
umbos. The two species appear to be genetically yellow, papery, being torn by spreading leaves, in
separatedon the basis of field observations;a molec- mature fascicles reduced to ca. 10 mm, persistent,
ular approachcould perhapselucidate their relation- darkred-brownto [Link] in fascicles of
ship with more accuracy. 3(-2-5), in dense tufts, spreadingobliquely forward,
persisting2-3 years, straightor slightly curved, rigid,
cm x 1-1.4 mm, with serrulate
28. Pinus teocote Schiede ex Schlechtendal & (7-)10-15(-18)
Chamisso, Linnaea 5: 76. 1830. Type. Mexico. margins, acute-pungent,light green. Stomata on all
faces of leaves, in 7-10 lines on the convex abaxial
Puebla/Veracruz:Mt. Orizaba,Sep 1828, Schiede &
face and 4-5 lines on each adaxial face. Leaf anat-
Deppe s.n. (holotype, HAL; isotype BM). Fig. 45
omy: Cross section triangularto transverse-triangu-
Pinus paItuli Schiedeex Schlechtendal & Chamissovar. lar; hypodermismulti-layeredespecially in marginal
strictaBenthamex [Link]. Conif. 157. 1847. areas; resin ducts 2-5, often 3, medial; stele elliptic
Type. Mexico. Michoacin: Near Angangueo, 1840. to transverse-obovatein cross section, outer cell
Hartnieg442 (holotype, K; isotypes. NY. W). walls of endodermis thickened; vascular bundles 2,
Pinus besseriana Roezl, Cat. Grain. Conif. Mexic. 20.
1857; Pinus vilmorinicna Roezl var. besseriana (Roezl) distinctly separate. Pollen cones crowded near the
Carriere,TraiteGen. Conif.. ed. 2. 2: 580. 1867. Type. proximalend of a new shoot, subtendedby scarious
Mexico. RoezI s.n.; original materialnot located. bracts, ovoid-oblong to cylindrical, 10-18 mm x
PinutsvilmorinianaRoezl, Cat. [Link]. Mexic. 20. ca. 5 mm,
1857. Type. Mexico. Roezl s.n.: original material not
yellowish green, turninglight brown after
located. shedding pollen. Seed cones subterminal,commonly
Pinus microcairpaRoezl, non A. B. Lambert (1803), opposite, sometimes 1-3, on short, stout, curved
Cat. Grain. Conif. Mexic. 21. 1857. Type. Mexico. peduncles which fall with the cones. Immaturecones
Michoacan: Morelia. 1857. Roezl s.n. (lectotype, Fl, (broadly)ovoid, 8-15 mm long, purplishred, matur-
heredesignated).
Pinus muilleria,tiaRoezl, Cat. Grain. Conif. Mexic. 21. ing in two seasons. Mature cones ovoid to ovoid-
[Link]:Mineraldel Monte,1857, oblong when closed, slightly asymmetrical, with a
Roezls.n. (lectotype,Fl. heredesignated). broad, flattenedbut oblique base when opened, (3-)
Pinus hlugelii Roezl ex [Link] Gen. Conif., ed. 4-6(-7) x 2.5-5 cm when open, often persistent.
2, 2: 583. 1867. Type. Mexico. Locality not stated, Seed scales ca. 60-100,
parting except those at the
1857, Roez s.n. (lectotype, Fl. here designated).
Pinus teocote Schiede ex Schlechtendal& Chamisso var.
base of the cone (semi-serotinous), thick woody,
imacrocarpa G. R. Shaw, [Pines Mexico] Publ. Arnold oblong, straight or recurved, purplish brown on the
Arbor. 1: 17, t. 10, figs. 5-7. 1909; Pinuls teocote abaxialside, darkbrownto purplishbrown with light
Schiede ex Schlechtendal& Chamissoformaimac- brown marks of seed wings on the adaxial side.
rocarpal (G. R. Shaw) Martinez. Pinos Mexic., ed. 2, in some cones more or
165. [Link]: SanCrist6bal de las Apophysis slightly raised,
Casas. 1895, E. W. Nelson 3218 (lectotype, A, here less flat, slightly more pronouncedon the proximal
designated: isolectotype, US). scales on one side of the cone, transverselykeeled,
apical margin angular or crenate, light brown, in
Tree, medium size, height to 20-25 m, dbh to 75 some cones with radial marks. Umbo dorsal, flat to
cm. Trunkmonopodial, sometimes forked, straight. blunt-pyramidal,transverse-rhombicin outline, up to
Bark thick, rough and scaly, forming longitudinal 6 mm wide, with a minute, deciduous prickle. Seeds
plates divided by deep, wide fissures, inner bark obovoid, slightly flattened,3-5 mm long, dark grey-

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Systematic Treatment 181

N'\II~~ ~I

.I 'I f!

ons
cone.
[Link].
[Link]
secion
Ovlifero? oflaf.
[Link]
hoot
witimmatue
ovu)ifrous
coe.
E.
coe.E
edscl twiws.C
Se ndsewns(antiain'",!.DG
x.;C.x3.

FIG. 45. Pinus teocote (A-C. Stidless.n.: D-G, Mittak8505). A. Branchwith leaf fascicles, pollen cones. and ovulifer-
ovuliferouscone. E. Ovuliferous
ous cones.B. Leaffascicles.C. Crosssectionof leaf. D. Flushingshootwithimmature
cones. F. Seed scale (two views). G. Seed and seed wings. (Magnifications:A, B. D-G. x0.5; C. x 30.)

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182 Flora Neotropica

~~:~r;
..,.;?
...... - ........
I;:
j.?~..:
-

....... ' .
25..
-4 ~~~ [?~~~~~ ..... '? ......e. .
. ..... '
Map

r
i~~~~~~~~~~~~~~~~~~~~~~~~~'
??. '..
' ....."i
'i'' .
:

.... ..~.J..~,
?..... i ~~~~~~+....
....~
......
.. -
i
i~~~~~~~
I t
. .... -? I...... .
.mmN??-~ ....4..; : ?
?- .-...
.. I ?~ ? .: .,.... ...1?
. .~~i....

I '?~~
... .
JIQM/.~~~~~~ ..... ...?..
.~~.....-
.;... .......
-g.
'arn
-~~~~~~~~~~~~~~~~~~?'~~~~~~~~~ -
.... !

i~~~~
ir ~ ~ .....;.~ ,, h . ?
"-r-. ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ I

~'f'
,
??...'
t......4....& ....
i' ...i--?- ~ "' '

'?.. I I ..... '; '?? ???- i ...., ... ..c...

,
' ;
f i ~T""--r----~~~~~~~~...
... t~~~~~~r-??????~~~~~~~~~4?
.. ;...

'C??~~~~~~~~~~~~
..1.....~ -
'5'' ? . . '.? . ? ......
.. . ??? ? ~ ?? ?

. ?i ..
""'.~~~~~~~
'. : ?? ??....
?
? ~. ~ ~ .
~ ... . ...
.,r '~..

M le.. ?25
- ;;~~~~~~~~~~~~~~~~~....
i
1.-
-~?-???I~~. i
.~
P" i
P.L
., ..

>
.. .. . ... : "" ' .... . :~ i
i~~~~~~~~~~~~
i ' .............:
......
~.? ........~
i? . ..... .:.~ :

' ..toe ~ ;I ~;"-~.,~~ ?

.~
' : -? . . .~_~.i. . .
...... ~~,
..~; ?e
: : .
..
I~~~~~~~~~~~~~
... ..... ~... .
...'i~?? '~'
... .'. -~-...-
. . 1 '_ _ i- ' ? O.;.....
i
l .....
??
:? -i :I
' .????I ,*?
_
..???.....i..,,,~ ???t
' ......,......;,er~~ . ?.............

Map 25. Pinus teoc~ote.

brown. Seed wings articulate but effective, held to guateinalensis" (not validly published, with a de-
the seed by two small, oblique claws, obliquely scription in Spanish only) from Quetzaltenangoand
ovate-oblong, 12-18 X 6-8 mm, translucent,yel- "between Huehuetenango and San Carlos Sija,"
lowish with a dark tinge. Number of cotyledons not which was taken up by Critchfieldand Little (1966).
observed. Perry (1991) reported that he was not able to find
Note: Epicormic adventitious shoots often occur in the species in Guatemala,and we, too, have not been
this species. able to locate specimens. Pinus teocote occurs in
Distribution and ecology (Map 25). Mexico: One various habitats,most commonly in ratheropen for-
of the most widely distributedMexican pines, most est or woodland associated with Quercus spp., on
abundantin central Mexico, distributed in Chihua- relatively dry sites with shallow soils. Its altitudinal
hua, Coahuila, Nuevo Le6n, Tamaulipas, Sinaloa, range is (1000-)1500-3000(-3300) m; a few collec-
Durango, Zacatecas, San Luis Potosi, Nayarit, tions are recorded from 700-800 m, but these may
Aguascalientes, Jalisco, Guanajuato,Quer6taro,Hi- be based on incorrectestimates. It often occupies dry
dalgo, Michoacan, Mexico, Distrito Federal, Tlax- ridges, sometimes on calcareous outcrops. Annual
cala, Puebla, Veracruz,Guerrero,Oaxaca, and Chia- precipitationis usually moderate,500-1000 mm, but
pas. Aguilar (1961) describes "Pinus teocote (fo.) much higher in parts of the central highlands of

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SystematicTreatment 183

Chiapas, where it is growing in broad-leaved forest Luis Potosi,"1878, Parry& Palmer844 (E, K, NY).
dominated by Liquidambar and Styrax. In the nor- TAMAULIPAS: Sierra de Tamaulipas, El Adame. 28 Dec
then parts of its range (Durango, Nuevo Le6n) it is 1969, Ortega C. s.n. (MEXU). VERACRUZ:Mendoza, 25
Mar1984,McCarter& Styles336 (FHO);nearPerote,10
reported to grow to a taller tree in favorable sites Mar1983,Nee & Taylor25871(NY);"interCruzBlanca
(Loock, 1950). It occurs with a number of other Nov 1828,Schiedes.v. (HAL);Las Lajas.
et Jalacingo,"
pines in various parts of its range, the most common near Perote,21 Mar 1980, Stead & Styles697 (FHO).
of which are from north to south: P arizonica, P ZACATECAS:
Chalchihuite, 1939. M. Martinez 3425
(MEXU).
engelmannii, P durangensis, P leiophylla, P monte-
zumae, P oocarpa, and P patula. Phenology: Pollen
dispersal is reported from as early as December, in Uses. Pints teocote is of importanceas a timber
Oaxaca, through March, with only a weak correlation tree. Apparentlydue to its moderatesize especially
with altitude (Critchfield, 1966b). in the southern part of its range, resin production
may locally count as the more importantmode of
Representative specimens examined. MEXICO. CHIA- exploitation.
PAS:Mun. Motozintlade Mendoza,21 Nov 1976. Breedlove The type specimen of P teocote was collected by
41699 (MO, NY); Jitotl6, 10 Aug 1965, Roe et al. 1153 C. J. W. Schiede and F. Deppe in September 1828 in
(UC). CHIHUAHUA: Bocoyna, 11 Jul 1971. Bye 1498 "Mexico, ad radices et in adsensu montis Orizabae"
(MEXU). DISTRITOFEDERAL: Eslava, 2 Oct 1904, Pringle
10018 (A, E, F, GOET. K, MICH, MO, NY, P, S, UC). (HAL), and described as a new species by Schlech-
DURANGO: Santiago Bayacora, 35 km from Durango, 2 tendal and Chamisso (1830). Bentham (1842) gave
Apr 1991. Styles, Favela et al. 2 (FHO). GUANAJUATO: On for Hartweg 442 a varietal epithet under P patula,
Cerro Puruagua,28 Oct 1987, Rzedowski45467 (CIIDIR); and Endlicher (1847) briefly characterizedit with
La Ventanilla,1829, Schiede 944 (M). HIDALGO:Real del "Foliis strictis, strobilus minoribus,"but the speci-
Monte, Dec 1837, Ehrenherg4 (Fl), Dec 1837, Ehren,berg
636 (B). Dec 1837, Ehrenberg 1024 (GOET, K, P, US);
mens clearly belong to P teocote, not to P patula,
near Zacualtepan,Jan 1840, Ehrenberg 1080 (Fl); Jacala. nor with P greggii (Martinez, 1945, 1948).
11 Jul 1947, Fassett 27688 (UC); Mineral del Monte, A numberof Roezl's names belong with P patula.
1848, Gregg 628 (NY); Pachuca, 16 Jul 1977, Mittak8498 and most could be [Link] a discussion of
(BANS, FHO, M). MEXiCO:Desiertode Le6n, 2 Apr 1950, Roezl's catalogue and "new species" see under P
Matltud 19497 (FHO, MO); Mun. Villa Nicolas Romero,
22 km NW of Cahuacfin.7 Dec 1975, Lopez 675 (CIIDIR, montezulnae.
MO); Ajusco Mtns., 1897, Pringle 6800 (A, E, GOET,JE, Shaw (1909) describedP teocote [Link]
K, M. MO. NY, P, UC, US). MICHOACAN: Near An- with cones "considerablylarger than those of the
gangueo, 1840, Hartweg 441 (BM, Fl, K, P); between typical variety"and with leaves in fascicles of 3, 4,
Angangueo and La Ventilla, 1828, Schiede s.n. (HAL); and 5. Martinez(1945, 1948) reducedShaw's variety
Cerro San Miguel, near Pomocuran,23 Feb 1980, Stead &
Styles 538 (FHO). NUEvo LE6N: 13 km S of Galeana, 21 to a formabut, at the same time, apparentlyexcluded
km W of Iturbide.26 Mar 1976, Hansen et al. 3856 (LL. part of Shaw's material in referring to a "Pinus
MICH, US); Mun. Villa Santiago, PotreroRedondo, 5 Jul slavae" of Robertsonbut citing Shaw's combination
1935. Mueller 2124 (MICH. MO, P, TEX); Monterrey,23 in synonymy. For the materialShaw had cited from
Jun 1888. Pringle 1964 (A, B, E. K, M, P. UC); Chipinque
National Park, near Monterrey, 10 Mar 1980, Stead & Eslava (Distrito Federal), Martinez suggested the
Styles 602 (FHO). OAXACA: Benito Juarez National Park, epitheton(forma) quiynquefoliata. What Shaw (1909,
20 Mar 1993, Higman. Padilla & Styles 47 (EAP. FHO. 1914) in fact cited underP teocote var. macrocarpa
HEH. MO); Miahuatlin, 22 Mar 1993. Higman, Padilla & are several collections (syntypes), two of which-
Styles 49 (EAP. FHO, HEH. MO); Ixtlan de Juarez. 12
May 1940, Kruger & Gillespie 28 (LL, MO, UC); San Pringle 10013 and 10018-were distributedas P
Felipe, May 1842. Lieb,hilnn14815 (LL, MICH, MO). 13 eslavae (ined.) from Eslava (Shaw, 1909). Perry
Dec 1895, Pringle 6243 (A, BM, E. GOET, JE, K, M, (1991) decided to treat this taxon and P teocote
MICH, MO, NY); San Pablo, 1828?, Schiede & Deppe 407 fo. quinquefoliataMartinez (an invalidly published
(M); along rd. from Oaxaca to Tehuantepec,at Km 72. combination:without Latin descriptionor reference
Mar 1974?, B. T. Styles s.n. (FHO); along rd. from Oaxaca
to Tuxtepec, near Estudiante, 26 Mar 1974, Styles 104 to one other than Shaw's var. tnacrocarpa)as syn-
(FHO);along rd. from Oaxaca to Tuxtepec,forest at Brecha onyms of P teocote and consideredthem occasional
1010, 28 Mar 1974, Styles 110 (FHO, K). PUEBLA: Chi- aberrantindividualsratherthanrepresentativeof dis-
nantla, May 1841, Liebhman14810 (MO, NY); Paso Cor- tinct populationswith consistentlysimilarcharacters.
tez, 17 Jul 1977, Mittak8505 (BANS, FHO, M); Boca del
Not surprisingly,this species shows considerable
Monte, 1907, Purpus 4378 (P, UC). QUERETARO: El Doc-
tor, 1840, Ehrenberg643 (B); San Joaquin, 2 Dec 1981, variation in its wide geographical range, but we
Huerta 722 (ENCB). SANLuts POTOSI: "In region of San agree with Perry that. at least morphologically,no

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184 Flora Neotropica

charactervariationhas been detected which enabled face. Leafanatomy: Cross section semi-circular;epi-
us to recognize any infraspecifictaxa. dermis very thick; hypodermis uniform, with 2-3
layers of cells; resin ducts 2-10(-14), medial, nar-
29. Pinus muricata D. Don var. muricata, Trans. row; stele oval in cross section; cell walls of endo-
Linn. Soc. London 17: 441. 1836. Type. United dermis uniform, thin; vascular bundles 2, distinctly
States. California: San Luis Obispo Co., at Coon [Link] conies crowded near the proximal
Creek, 1832-1833, T. Coulter 712 (isotype, TCD). end of a new shoot (not always distinct due to
Fig. 46 the multinodalhabit), oblong-cylindrical,15-20 mm
long, pink to reddish,turningyellowish brown. Seed
Pinus edgariana K. T. Hartweg,J. Hort. Soc. London 3: cones subterminal, at the base of the subsequent
217. [Link]:Monterey
Peninsula,Hartweg(originalmaterialnot located). shoot, solitary or in whorls of 2-5 on stout, 5-10
Pinus muricata D. Don var. anthonvi J. G. Lemmon, mm long, bracteate peduncles, reflexed, tenacious,
ed. 3. 43. 1895;Pinus
[Link]-bearers, maturecones seemingly sessile, opening only partly.
nmuricataD. Don subsp. anthonvi (J. G. Lemmon) E. Immaturecones ovoid, with prominent spines, ini-
Murray,Kalmia 13: 23. 1983. Type. Mexico. Baja tially purplishred, maturingin two seasons. Mature
A. W Anthony
CaliforniaNorte:"NearSan Quentin,"
cones serotinous, narrowlyovoid to ovoid-attenuate
(originalmaterialnot located).
Pinus relmorata H. Mason, Madrono 2: 9. 1930; Pinus when closed, or strongly asymmetrical, broadly
muricataD. Don [Link](H. Mason)E. Mur- ovoid with long, spiny apophyses, 5-7(-8) x 4-5
ray, Kalmia 12: 23. 1982; Pinus inuricata D. Don var. (-6) cm when (half) open. Seed scales ca. 70-100,
remorata(H. Mason)Silba,Phytologia68: 56. 1990.
SantaCruzIs., summit
[Link]: partingvery slowly except those at the base or lower
of passfromWestRanchto PrisonersHarbor,25 Sep half of the cone, oblong, straightor slightly curved,
1927,Mason4096 (holotype,DH?-n.v.;isotype,CAS). thick woody, purplishbrown,with grey-brownmarks
Pinus muricata D. Don var. stantonii Axelrod, Univ. of seed wings on the adaxial side. Apophysis very
Calif. [Link]. Sci. 127: 77. 1983. Type. United
variable,from slightly raised to extremely elongated
SantaCruzIs., 4 Jun 1983,Axelrod
[Link]:
E-1(lectotype,GH,heredesignated; MO).
isolectotype, (bifacial-conical),especially on the upperside of the
cone, rhombic to pentagonal in outline, sometimes
Shrub or small tree (in Mexico), height to 4-10 curved, transverselykeeled, up to 15 mm wide and
(-15?) m, dbh to 20-50 cm. Trunk monopodial, 20 mm long, from dull, dark brown to lustrous light
often branchingnearthe ground,erect or curved and brown. Uibo dorsal, obtuse to acute, if enlarged
crooked. Bark rough and scaly, exfoliating, on larger forming a flattened, curved spine, armed with a
trunkswith deep longitudinalfissures, darkbrown to sharpprickle in obtuse forms, 2-10 mm long. Seeds
grey, on young trees and branches thin, exfoliating obliquely ovoid, slightly flattened,5-6 x 3-4.5 mm,
in papery flakes exposing pale brown inner bark. grey to black. Seed Vwingsarticulate,effective, held
Branches long, relatively thick, spreading or as- to the seed by two thin, oblique claws, obovate-
cending, in shrubsoften assurging,persistent,form- oblong to dolabriform,14-18 X 5-8 mm, yellowish
ing an open, broad, irregular crown. Shoots brown to grey-brown.Cotvledons4-6.
multinodal,rough with large, short decurrent,persis- Distribution and ecology (Map 26). United
tent pulvini. Cataphylls ca. 10 mm long, subulate, States: In California on and near the coast of the
curved or reflexed, scarious, with erose-ciliate mar- mainland counties of Monterey (Monterey Penin-
gins, brown, early deciduous. Vegetativebuds ovoid- sula), San Luis Obispo, and Santa Barbara;also on
acute, the terminalbuds 10-15 mm long, the laterals Santa Cruz and Santa Rosa [Link] populations
smaller,not or slightly resinous;the scales imbricate, in N California were referredto as P muricata var.
with free apices, subulate,with erose-ciliate margins, borealis Axelrod. Mexico: Baja CaliforniaNorte, in
light brown. Fascicle sheaths 10-14 mm long ini- two localities on the mainlandnear the coast, W and
tially, reducedto <10 mm on maturefascicles, ripped SW of San Vicente. Reports of its occurrence on
at parting of leaves but persistent, hyaline-whitish, Cedros Island by various authors (Epling & Rob-
brown at base. Leaves in fascicles of 2, in rigid inson, 1940; Martinez, 1948, even with a photo-
tufts at the ends of branches, persisting 2-3 years, graph) as well as on GuadalupeIsland (e.g., Mason,
spreadingca. 45? from the shoot, straightor slightly 1932) are erroneous and the reports pertain to P
curved, rigid, (7-)10-14(-16) cm X 1.3-2 mm, with [Link]. In its locations in Mexico it is the
serrulate margins, acute, light green or dark green. only species of Pinus growing there; it is sometimes
Stomata on both faces of leaves, with 6-9 lines on accompaniedby [Link]-
the abaxial, convex face and 4-7 lines on the adaxial sii or Juniperuscallfornica. The largest populations

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Systematic Treatment 185

rp~e~p~lL4li~y*' ir

s$.zais __.' . . i
oy^1"i L .
?a b
FIG. 46. Pints tmurictaivar. munricta (A-Ca, Breedlo'e 4189; Cb, E, High/es & Sidles 178; D. Fa. Bi(ler 205; Fb.
G, Breedlove4176). A. Branchwith leaf fascicles and cone. B. Leaf fascicle. C. Cross sections of leaves. D, E. Ovulifer-
ous cones. F. Seed scales. G. Seed. (Magnifications:A. B, D, E. x0.5: C. x25; F, G. x .)

are on and near Cerro Colorado, an escarpment of have been made because it is next to the road, but
eroded rhyolite on the N side of Rio San Isidro, an to the NE there are small mountainswith stands of
intermittentstream. It is here where most collections this pine, mostly on the N-facing slopes. It occurs

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186 Flora Neotropica

from near sea level to ca. 100 m. It grows within within populations. On Cerro Colorado, cones of
the chaparralzone influenced by (summer and au- both extremes-nearly symmetrical,with flat apoph-
tumn) fog and winterrain,probablyamountingannu- yses and small, prickly umbos (Farjon 260,
ally to ca. 500 mm. Often brushfires sweep the area, Hughes & Styles 178) and asymmetrical,with very
killing stands of pines, but the serotinouscones are strongly developed apophyses and umbos (Broder
adapted so as to open quickly after fire to release 205)-have been collected. Most stands are situated
the seeds when the undergrowthhas been cleared in chaparral,a vegetation type adapted to frequent
away. This often consists of Adenostona, Arctosta- brush fires. Reseeding from one or a few individuals
phylos, Artemisia, Ceanothus, Heteromeles, Salvia, with ripe seed, with much of the gene pool lost, may
and other tall shrubs. have caused a "foundereffect" eliminating interme-
diates (see also Millar, 1986b).
[Link] This heterogeneity seems to have led to the de-
CALIFORNIA NORTE:Near San Vicente, 15 Aug 1962,
Breedlove4176 (FHO, SD), 15 Aug 1962, Breedlove4177 scriptionof numeroustaxa. Only a northernvariety,
(ENCB);8 km W of SanVicente,on slope, 15Aug 1962, occurring outside our region, is currently accepted
Breedlove4189 (MICH);W of SanVicente,NWslopesof by most botanists (Farjon, 1993). Cone types, as
CerroColorado,14Feb 1960,Broder205 (FHO,NY,UC); emphasizedby Axelrod (1983) in orderto be able to
nearSanVicente,[Link] coast,ca. 3 kmfromErendira, classify Pleistocene fossil cones, are of little value
I Oct 1992, Farjon260 (E, U, UC); (Rio) San Isidro in determininggenetic relationshipsin this species.
Canyon,along rd. fromSan Vicenteto Er6ndira,II km
NEof Erendira, 4 Jan 1988,Franklin5843 (NY);nearSan
Vicente,top& sidesof [Link],15Jul 1957,Gentry 30. Pinus radiata D. Don var. binata
16753(US); 15kmW of SanVicente,on N slopeof Cerro (Engelmann)
Colorado, 26 May 1987, Hughes & Styles 178 (FHO);
J. G. Lemmon, [Link] Pacif. Slope, ed.
ArroyoHediondo(ArroyoSanVicente),5 kmSEof Erend- 3,42. 1895. Fig. 47
ira,21 Jul 1979,Moran27871(SD);[Link]
DescansoandErendira,13 Feb 1979,Spjut & Marin5299 Pinus insignis D. Douglas ex J. C. Loudon var. binata
(US). Engelmann in S. Watson,Bot. California2: 127-128.
1880; Piins radiata D. Don forma binata (Engelmann)
J. T. Howell, Leafl. W. Bot. 3: 3. 1941; Pinus radiata
Uses. Due to its bushy habit and very restricted D. Don subsp. binata (Engelmann)E. [Link]
occurrence in Mexico, Pints muricata is not com- 13: 24. 1983. Type. Mexico. Baja California Norte:
mercially used. There has been some local "logging" GuadalupeIsland, 1875, Palmer s.n. (holotype, MO).
in the past, but due to the low populationdensity in Pinus radiata D. Don forma guadalupensis J. T. Howell,
Leafl. W. Bot. 3: 3. 1941. Type. Mexico. Baja California
the region, this use is not extensive. There is evi- Nore: Guadalupe Island, Howell 8185 & 8267 (syn-
dently cattle grazing in the area, but we do not know types, CAS-n.v.).
its effects, if any, on these relict stands of pines. Pinus muricata D. Don var. cedrosensis J. T. Howell,
In a brief series of descriptions of five "new" Leafl.W. Bot. 3: 7. 1941;Pinusradiata D. Don var.
cedrosensis (J. T. Howell) Silba, Phytologia 68: 60.
pines collected by T. Coulter in California, Don [Link]:CedrosIs..
(1836) described Pinus muricata, apparently only 6 Jun 1925,Mason2030 (holotype,CAS;isotypes,K,
with a cone at his disposal. It was collected at Coon NY,US).
Creek nearSan Luis Obispo at ca. 1000 m, according
to Millar (1986b). It is a typically disjunct, relict Tree, small to medium, or a shrub on exposed
taxon which had a wider distributionin the geologic ridges, apparentlyonly a few trees reachinga height
past and is now being reducedprimarilythroughloss to 20-25(-33) m, dbh to 200-220 cm (on Guadalupe
of habitat due to coastal abrasion (Mason, 1932; Is.). Trunk monopodial, often branched or forked
Axelrod, 1983). near the ground, massive in solitary trees. Bark
The populations are often very heterogeneous, thick, rough and scaly, deeply fissuredverticallyand
which is expressed in the variabilityof cone types, horizontally,dark brown, weathering blackish grey,
number of resin ducts in the leaves and stomatal on young trees and branches scaly, exfoliating in
anatomy. Analysis of mtDNA variation (Strauss et small chips. Branches thick, long, often contorted,
al., 1993) and cpDNA variation(Hong et al., 1993) spreading or ascending, forming dense, flat-topped
has revealed a similar heterogeneity,at least among or bushy crowns. Shoots multinodal, the leading
the Californianpopulations,confirmingthe morpho- ones thick, rough with prominent,decurrentpulvini.
logically based [Link] intrapopulationvaria- Cataphylls small, 5-10 mm long, subulate, curved,
tion of mtDNA and cpDNA appears to be limited, scarious,with erose margins,brown,early deciduous.
polymorphy, especially in cone types, occurs also Vegetativebuds ovoid-acute, the terminal buds 10-

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Systematic Treatment 187

15 mm long, the laterals smaller, not resinous; the var. radiata has 3 disjunct populationson the coast
scales imbricate, spreading slightly, subulate, with of California(Griffin& Critchfield,1972). The habi-
ciliate margins, orange-brown. Fascicle sheaths of tat of this species is extremely maritime,influenced
juvenile leaves up to 15 mm long, with separate by the cold watersof the southwardflowing currents,
scales visible, orange-brown,of mature leaves 5-8 bringinghigh humidity in summeras fog. Precipita-
mm long, dark brown, [Link] in fascicles tion as rain is erratic:fog drip probablyaccounts for
of 2(-3 on leading shoots), in tufts on ultimate much of the effective moisture(Libby et al., 1968).
branches, persisting ca. 3 years, straight or curved The climate of the islands is Mediterranean-like,
to twisted, rigid, 8-15 cm X 1.1-1.6 mm, with with cool to cold conditions in winter on Guadalupe
serrulatemargins, acute-pungent,bright green. Sto- Island. Pine stands are restricted to foggy ridges,
mnataon all faces of leaves, in (6-)8-10 lines on the foggy windward slopes, and moist canyons. Here
convex abaxial face and 4-5 lines on the adaxial summer temperaturesare considerably cooler than
face(s). Leaf anatomy: Cross section semi-circular in desert areas nearbywhere fog does not persist. Its
or transverse-triangular; cuticulaandepidermisthick; altitudinal range on sedimentary and metamorphic
hypodermis more or less uniform, I-2-layered; resin CedrosIslandis (200-)300-650 m, on volcanic Gua-
ducts 2-5(-8), medial, occasionally 1-2 internal; dalupe Island 300-1160 m. Associated woody spe-
stele oval in cross section; cell walls of endodermis cies are few. Juniperuscalifornica has been reported
not thickened;vascular bundles 2, [Link] on Cedros Island, and Quercus tomentella is common
cones crowdednearthe proximalend of a new shoot, among the pines on GuadalupeIslandwhere Ervthea
or more scattered on multinodal shoots, subtended edulis forms small groves. Goats are a major prob-
by broad, scarious bracts,oblong to cylindrical, 15- lem, reducingchances of rejuvenationon this island
20 mm long, pink to reddish, turning yellowish (Libby et al., 1968). On Cedros Islandthe vegetation
brown. Seed cones subterminal,sometimes also in a resembles that of peninsular Baja California. Phe-
secondary whorl, usually in whorls of 2-5, some- nology: Time of pollen dispersalnot observedon the
times solitary, on short, stout peduncles, very tena- islands, probablyin early spring.
cious, serotinous. Immaturecones ovoid to broad
ovoid, ca. 15 mm long, on densely bracteate,curved, Representative [Link]
5-15 mm long peduncles, maturingin two seasons. CALIFORNIA NORTE:Cedros Is.. Jul 1896. Anthony77 (A.
NY. US); Cedros Is., Cerros Contiguos, "El Chollal," 21
Maturecones ovoid, nearlysymmetricalor asymmet- Feb 1978,Arteagaet al. 133 (ENCB,MEXU);CedrosIs..
rical with an oblique base, (5-)6.5-9(-13) X (4-)6- 1 kmNEof old mineof Caiadade la Mina,14Apr 1983,
7(-9) cm when open. Seed scales ca. 130-160, part- Dice el al. 320 (SD); Cedros Island, 1.5 km S of lighthouse
ing after a considerable time or only after extreme at Cabo Norte, 19 Jan 1975, Hendrickson14482 (ENCB,
MEXU, NY, SD); Cedros Is., 12 Mar 1911, Rose 16186
heating, oblong, straight, thick woody, purplish (NY, US); Cedros Is., 12 Sep 1952, Littlepage s.n. (SD):
brown, with light brown marksof seed wings adaxi- CedrosIs., Feb 1940,M. Martfiez3467 (MEXU);Cedros
ally, sometimes faintly [Link] slightly Is., "PuntaNorte,"3 Mar 1964, MadrigalS. 1475 (MEXU);
raised, transversely keeled, or gibbous especially Cedros Is., 6 km N of Gill Peak, 27 Mar 1952, Moran
toward the base on one side of asymmetricalcones, 3512 (UC);CedrosIs., nearold coppermineca. 20 km
from N end of island, 27 May 1971, Moran 18434 (SD):
rhombicto weakly pentagonalin outline, with radial Cedros Is., "on highest peak" ICerro Redondo], 18 Mar
lines, lustrous light brown. Umbo dorsal, flat or 1889,Paller 763 (US);CedrosIs., 1939,Stockwell2005
depressed, more or less rhombic in outline, 4-7 mm (MEXU); Guadalupe Is., on NE part of island, Jul 1896.
wide, without a prickle, dark brown to grey. Seeds Anthtony15 (NY, US); GuadalupeIs., crest of ridge on NE
partof island, 23 Apr 1958, Carlquist442 (UC); Guadalupe
obliquely ovoid, slightly flattened, (5-)6-8 X 4-5 Is., 1893, Franceschi2 (F, NY, UC, US); GuadalupeIs., N
mm, grey-brownwith dark spots. Seed wings articu- end of island, 15 Nov 1931, Howell 8183 (NY. SD. US);
late, effective, held to the seed by two thin claws, GuadalupeIs., 16 Mar 1932, Howell 8269 (US); Guadalupe
obliquely obovate or cuneate-dolabriform,14-20 X Is., 23 Apr1958,Wiggins& Ernst51 (US);Guadalupe
Is..
7-9 mm, yellowish brown, semi-transparent,with 2 Mar 1911, Rose 16001 (NY, US); GuadalupeIs.. II Jun
1955, Kuijt & Miller 1018 (MICH. SD. UC); Guadalupe
grey-brown stripes. Cotyledons 5-9 (for mainland Is., Sep 1939, M. Martinez3466 (MEXU); GuadalupeIs.,
var. radiata). NE ridge, 14 Dec 1957, Moran6432 (SD. UC); Guadalupe
Distribution and ecology (Map 26). Mexico: In Is., 1875, Palmer 90 (NY); Guadalupe Is.. 30 Aug 1980.
Baja California Norte, only on Cedros Island near Victoria& Ferndndezs.n. (ENCB).
the Pacific coast and on Guadalupe Island ca. 250
km off the Pacific coast (ca. 29?N, 118?20'W).On Uses. Guadalupe Island is uninhabited;Cedros
these islands it is the only native pine. P radiata Islandhas a small fishing village. There is no exploi-

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188 FloraNeotropica

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.- ?'9CI i - .?????I?L .. -;*i?;: ;?.?. .j.
1 i?,? ?.,"'
?r ?- i?:
?? ..-.i. ?: : :
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i .:?. :*? .,? :::'? ??
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:
?. r:. ?... .? t * r. *
?; r
f
??
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??L.,. _? ? ?-Q..i .,
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C:I:-6:?4'???''bl
e--?i-:
-?
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.r
'''''?-- i?? i:.
??
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?,?''.9: . :. ;??:!:
?!?:
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':r;:;??
.?::-? j* ?-?
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.:.:.

?: . ?:?? ;? .: ??i ??:?':;,??:

:?."''?:' ::?''?I
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?'??.' ??..r''' ': :??? ?''? :?.?: ;' ?: ?:
.. ;?: ; :'t?'?? '?::
L? ? :: ?:. ? ?? , I?
r.. ..r. 1i r: r
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: I iI
::)?: i ?
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, ??- ? ??
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.'.:: : ?? I?. :;i7??.
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.?)-? ???; ??.?:-rlli-;? :P:':* ..
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'' ' '? " ? .' ?? r..i? ; ?,; ; ??;;
i? ::?rl??: ?.?::? -- ?:: ''c';" I
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-.::.. ?i??. ; '''i'
'
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?'; '? ;? ;??I
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"?' u. ? .- -?: ??:
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: ? :? -? ',;..
:. 4:Li? :I
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. .?
:.?:r? :??.iff-'?':??? :? :??. j.
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tai ?? .I... r. ??.!:, ????
;.::i??:.?:??i ''
::' :?:?
'?, :.':?. '.-+:.?i: ...P-???. ;:I
::: -I-ii ?is:. .,:.. ?. ?? ??
:?? :?: :;S ?? :t.. f- ?r
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-- -.-...--..--?-.-.-.-..-i?*;n?,a;-;-;

[Link]
Map26. Pinusrtlrrirrrttr [Link]).
(triangle)l mrlirrrr

tation of the pines of these islands. Overgrazing long). The leaves on lateral shoots are in fascicles
by feral goats seriously restricts rejuvenation on of 2, on leading shoots in 2-3, never 4 as occasion-
Guadalupe Island, where there are not more than ally on the mainland. Howell (1941) found most
400 maturetrees (Libby et al., 1968). cone shapes of specimens from GuadalupeIsland to
In 1875 E. J. Palmercollected a pine from Guada- be intermediateand recognized them as two forms,
lupe Island as "Piins itsignis var.," which was de- one symmetrical (fo. guadalupensis), and the other
scribed as var. binata by Engelmann(1880b). Lem- "intermediate"(fo. binata), of P. radiata.
mon (1892), in pointing out the superfluity of P From the time of its first botanical reference in
insignis Loudon (1838), proposed P. californica 1888, the pines of Cedros Island have been referred
Loiseleur-Deslongchamps(1812) as the earliest valid to as Pintus rmuricata,until Mason, first in part
name for the Monterey pine, but Lemmon (1893, (1932), later entirely (1934) referred them to his
1895) rejected it in favour of P radiata, because new species P renmorata, described from Santa Cruz
Loiseleur's name, lacking a type, could be attributed Island, [Link] species is a taxonomic syn-
to P attenuataor P radiata (see also Millar, 1986a). onym of P. muricata (Linhart et al., 1967; Kral,
The only pine indigenous to Guadalupe Island is 1993; this volume). However, the cone type of the
P radiata var. binata, notwithstanding reports of pines on Cedros Island is that of P. radiata, most
both P muricata and P remoratathere (see Howell, closely those "forms" found on Guadalupe Island,
1941; Libby et al., 1968). It differs only slightly to which it is also comparable in size. Although
from P radiata var. radiata in [Link] cones recognizingthis, Howell (1941) classified the Cedros
are on average smaller and more symmetrical, but Island pines as P muricata var. cedrosensis. With
cones as large as those found near El Carmen in Axelrod (1967) and Perry (1991) we conclude that
California do occur (Rose 16001, US: 10-13 cm they belong with P radiata, which is confirmed by

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Systematic Treatment 189

,-,
iI ''

I / 1 y

FIG. 47. Pinousr(liut var. binat( (A. B. F-H. MornJ 6432: C. E. Hen(lrickson14482: D. Kijit & Miller 1018). A.
Branchletwith leaf fascicles and buds. B. Leaf fascicles. with enlarged (X 10) leaf margin. C. Cross section of leaf. D.
Flushing leaf fascicles and juvenile ovuliferous cone. E. Branchwith ovuliferous cone. F. Symmetricalovuliferous cones.
G. Seed scale (two views). H. Seeds with articulatewings. (Magnifications:A. B. D-H. xO.5; C, x25.)

our examinationof Mason 2030, Howell's type, and size between material from the two islands, with
all other material that we have seen from Cedros those of Cedros Island smaller (6.5 + 1.3 cm vs.
[Link] (1967) observed a difference in cone 8.2 ? 1.2 cm) and treated them as P radiata var.

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190 Flora Neotropica

cedrosensis (without referenceto the basionym, later exfoliating in thin, papery scraps, greyish brown.
corrected by Silba, 1990). This treatment was ac- Branches of first order long, spreadingto assurging,
cepted by Millar (1986a). Since the cone form found along much of the lengthof the stem in free-standing
on Cedros Island definitely occurs on Guadalupe trees, in dense stands only in the upper part; of
Island,there is no convincing evidence for this sepa- higherordersslender,spreading,formingan irregular
ration. Both populations appear to have a similar or rounded, open crown. Shoots often multinodal,
morphology of stomatal accessory cells (Newcomb, rough, with short decurrent,persistentpulvini. Cata-
1959). phylls subulate, scarious, tapering into a caudate
Axelrod (1967, 1980) inferreda possible relation- apex. margins erose-ciliate, dark brown; deciduous
ship between P radiata (mainlandand insular) and before leaf fascicles. Vegetativebuds ovoid-acute,
P oocarpa and P pringlei of Mexico, based on the terminalbuds 15-30 x 10-15 mm, the laterals
(fossil) cone [Link] disagreed with Mason smaller,resinous,brown;the scales appressed,subu-
(1932), who suggested a (possible Pliocene) insular late, scarious, with erose, hyaline margins. Fascicle
origin of the closed-cone pines of California, and sheaths initially 10-18 mm long, papery, orange-
puts their origin much furtherback to the Eocene- brown at base, whitish hyaline distally, persistentbut
Oligocene [Link] increasingarid- reduced to 3-6 mm, turning grey-black. Leaves in
ity during the Miocene restrictedthe temperatetaxa fascicles of 3, rarely2, spreadingobliquely from the
to the coastal fog belt and oceanic islands, while the shoot, persisting 2-3 years, rigid, straight, (8-)10-
"'aridforms" (now P atteenuata)could occupy their 12(-14) cm X 1-1.5 mm, margins serrulate, apex
former locations. The fossil record indicates a range acute, light green or slightly [Link]
along much of the central to southern Californian on all faces of leaves, in 8-12 lines on the convex
coast up to the late Pleistocene (Mason, 1932; Axel- abaxial face and in 3-5 lines on each adaxial face.
rod, 1967). All fossil cone types (P lawsoniana Leaf anatomy: Cross section transverse-triangular;
Axelrod, P. pretuberculataAxelrod, P. radiact D. hypodermis uniform, with 1-2 rows of cells; resin
Don) are within the range of variationobserved in ducts 2-3, medial; stele wide, elliptic in cross sec-
the extant species. The island populationsare in part tion; outer cell walls of endodermis not thickened;
similar to the Pleistocene P lavsoniatna, which has vascularbundles 2, distinctly [Link] cones
been replacedon the mainlandby the moreasymmet- crowded near the proximal end of a new shoot,
rical forms (Axelrod, 1980). Pinus ra(diautvar. bi- subtendedby scarious, subulatebracts,ovoid-oblong
nttaais probablyan insularrelict now extinct on the to cylindrical, 15-20 x 5-7 mm, yellowish to yel-
mainland(Libby et al., 1968). lowish brown. Seed cones subterminal, solitary or
more often in pairs or whorls of 3-5, on short, stout,
31. Pinus attenuata J. G. Lemmon, Mining Sci. recurved peduncles, very persistent on main stem
Press 64: 45. 1892. Type. United States. California: and branches. Immaturecones ovoid-acute, 10-15
Berkeley, J. G. Lemtllmon s.n. (lectotype, UC No. x 5-8 mm, purplish, maturing time two seasons.
A.
338321, designatedby Farjon, 1993: 129; isolec- Mature cones seemingly sessile, reflexed, extremely
48 serotinous, ovoid-oblong to ovoid-attenuate,slightly
totype, K). Fig.
curved, asymmetrical, 8-15 x 3.5-6 cm when
PinuscalifornicaK.T. Hartweg,J. [Link]. London2: closed (up to 8 cm wide when open). Seed scales
189. 1847,nonLoiseleur-Deslongchamps. 1812;Piius ca. 150-180,
tuberculata G. Gordon, J. Hort. Soc. London 4: 218.
parting after fire (extreme heat) but
1849 (nom. nov., basedon P cwlifornlicaK. T. Hartweg). remaining connate at base, thick, rigid, oblong,
non D. Don, 1836. Type. [Link]: straight or slightly curved when parted, purplish
SantaCruzMtns.,H(rtweg,1847?(originalmaterial brown to reddishbrown on the abaxial side, reddish
not located). brown with light brown marksof seed wings on the
Pinus tuberculataG. Gordon var. uc'uttu H. Mayr, Wald.
adaxial side. Apophysis rhombic to pentagonal in
Nordamer. 275. [Link]:
Marr?(originalmaterialnot located). outline, transverselykeeled, from slightly raised on
the lower side of the cone to conical and curved on
Tree,mediumsize, height to 15-20(-25) m, dbh to the upper side, especially toward the base of the
40-50 cm. Trunkmonopodial,straightor sometimes cone (there >10 mm long), ochraceous or yellowish
curved. Bark rather thin, rough, scaly, breaking in brown,slightly lustrous,weatheringgrey. Umbodor-
small, rectangularplates, exfoliating slowly, greyish sal, rhombic, from almost flat to pyramidalor unci-
brown to grey; on young trees and branchessmooth, form in conjunctionwith developmentof the apophy-

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Systematic Treatment 191

FIG. 48. Pinus attenuatai(A-C, E, Meigs s.n.; D. bFirjlons.n., 1973, California. not cited). A. Branch with leaf
fascicles. B. Leaf fascicles. C. Cross section of leaf. D. Ovulifcrous cone. closed. E. Ovuliferous cone. opened. F. Seed
scale (two views). G. Seeds with wings. (Magnifications:A. B. D-G. x0.5: C. x 30.)

sis, curved toward the cone apex, up to 5 x 5 x 3.5-4.5 mm, blackish grey. Seed wings articulate
mm, darker than apophysis; prickle absent. Seeds but effective, held to the seed by two oblique claws,
obliquely ovoid, flattened, apex slightly acute, 5-7 oblong, with one side curved, 12-18 X 5-7 mm,

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192 Flora Neotropica

Uses. Pilns acttenuatais not used as a timber tree

in Mexico and is probably not an importantsource
of [Link] this, it may requiresome protec-
tion statusas the southernmostand disjunctmetapop-
ulation of the species.
Hartweg (1847), who discovered the species in
the Santa Cruz Mountains south of San Francisco,
California. ascribed it to "the doubtful and little
known Pilus californica" (of Loiseleur-Deslong-
champs) and gave an accurate account of it. Both
this name as applied by Hartweg and the substitute
P tiubercularta proposed by Gordon (1849) are later
[Link](1892) renamedthe species in a
nomenclaturalnote, referringto "Lemmonin Herb."
Hartweg's original collection, if it exists, could not
be traced, and instead the collection by Lemmon
in UC has been chosen as the lectotype (Farjon,
1993).
Mayr (1889) described P. tuherculata var. acuta
from southern California, which is now considered
to be a cone form within the range of variationfound
in the species in many populations. Pinus attenuata
is morphologically less variable than its relatives
among the "closed-cone pines" P radiata and P
ilnricata. In trials it readily hybridized with the
former (Millar & Critchfield, 1988). Chloroplast-
DNA variationanalysis confirmedthe close relation-
ship of these pines (Hong et al., 1993).
The fossil record is ratherlimited for this hillside
pine. Mason (1932) published Pitnus linguifornmis
Map 27. Pinuls (anlltht(l(. from Pleistocene clay beds in Los Angeles County,
California. Axelrod (1983) suggested it might still
occur near the Pacific coast, but even for this fossil
yellowish to grey-brown. Cotyledois (5-)7-8(-9). there seems little ground to separate it from the
Distribution and ecology (Map 27). Mainly in present-daytaxon (Millar, 1986a).
United States (Oregon and California), becoming The limited range of the species in Mexico, dis-
increasingly scattered in southern California and just junct from its nearest metapopulationin California
reaching into Mexico (Baja California Norte). where by some 250 km, could be relictual. Its individual
it occurs in a few small populations on the lower W trees are morphologically identical to it. A more
slopes of the Sierra Juirez near Ensenada. It is a fire scattereddistributionoccurs throughoutthe southern
successional tree on dry slopes in chaparral or similar range of the species (Griffin & Critchfield, 1972)
vegetation, also on rocky outcrops with little other and a suggested discontinuityin certaincharacters-
plant growth. In the United States its altitudinal range e.g., cone size (Newcomb, 1962)-is likely related
is 3(X)-12(X) m. In Mexico it is found mainly at 250- to these disjunctions,where in the case of a dispersal
600 m. Phenology: Disperses pollen in late March- event, trees within a population are likely to be
April in most of its range. descendantsof one or a few founder trees only.

Representative specimens examined. MEXICO. BAJA 32. Pinus greggii Engelmann ex Parlatorein Can-
CAIIFORNIA NORTE:Near Ensenada,Sep 1940. M. Martf- dolle, Prodr. 16(2): 396. 1868. Type. Mexico. Coa-
ne:. 3470 (MEXU): 6 km E of Ensenada,Dec 1927. Meigs
s.n. (UC); 6 km NE of Ensenada, along Hwy. 1. 14 Apr
huila: San Antonio de las Alanzanas, near Saltillo.
1968, Mor(a 14855 (MICH, SD, US); hills N of Ensenada. 31 Aug 1848, Gregg 402 (lectotype, MO, here
6 Sep 1981. Moranl29839. 29846 (SD). designated; isolectotype, GH). Fig. 49

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Systematic Treatment 193

Tree,small to medium, height to 20-25 m, dbh to remainingconnate, purplishbrown, with light brown
70-80 cm. Trunkmonopodial,usually straight,terete. marks of seed scales adaxially. Apoplhysisflat or
Bark thick, with deep, longitudinal fissures and slightly raised, rhombicto obtrullatein outline, with
rough, elongated plates, on the upper part of the an undulateor crenate uppermargin,(weakly) trans-
trunk and branches smooth to finally scaly, smooth versely keeled, sometimes gibbous on one side to-
on young trees, greyish brown. Branches long, slen- ward the base of the cone, lustrous ochraceous or
der, spreadingor curved downward, not pendulous, light brown, weathering grey. Unibo dorsal, de-
forming a rounded, dense or more open crown. pressed or flat, rhombic in outline, 3-4 mm wide,
Shoots often multinodal,smooth, ridged with decur- with a minute, deciduous prickle, light brown to
rent bases of pulvini, reddish brown to grey-brown. grey. Seeds obliquely obovoid to ellipsoid, slightly
Cataphylls lanceolate to subulate, straight, erose- flattened, 5-8 x 3-4 mm, grey-brown to blackish
ciliate at margins,darkbrown,soon deciduous. Vege- brown. Seed wings articulate,effective, held to the
tative budsovoid-conical to oval-oblong, the terminal seed by two claws and a thin layer of tissue covering
bud 10-15 mm long, the laterals more ovoid and the seed on one side, oblong, with a straightand a
smaller, not resinous;the scales free or spreadingat curved side, 15-20 X 6-8 mm, yellowish to grey-
apex, subulate, ciliate at margins, brown. Fascicle brown. Numberof cotyledons not observed.
sheaths initially ca. 10 mm long, with chartaceous, Distribution and ecology (Map 7). Mexico: In
yellowish brown to whitish brown scales, persistent extreme SE Coahuila, S Nuevo Le6n, SE San Lufs
but reduced in mature fascicles to 3-5 mm, scaly, Potosi, Quer6taro,Hidalgo,and N [Link] altitudi-
exfoliating, darkbrown, weatheringblackish brown. nal range is 1300-2600 m; in the northernpartof its
Leaves in fascicles of 3, in usually rigid, spreading distribution,2300-2700 m. Annual precipitationis
tufts on the ultimate branches, persisting up to 4 600-800 mm in much of its range, except on the E
years, stiff and straight, (7-)9-13(-15) cm x I- escarpment of the mountain ranges along the Hi-
1.2 mm, serrulatealong the margins,acute-pungent, dalgo-Veracruz borderline, where it is 10(0-16(H)
bright, lustrous green. Stomata on all faces of the mm. In the north it is more often found on slightly
leaves, in 6-8 lines on the convex abaxial face and alkaline soils (pH 7-8); in the south, on acid soils
3-4 lines on each adaxial face. Leaf anatonmv: Cross (pH 4-5) (Dvorak & Donahue, 1992). It is nowhere
section transverse-triangular, with a convex abaxial abundantin its scattered range, and always occurs
side; hypodermis uniform, with a double layer of mixed with, e.g., Querculs, Plat(lons, Liqyidlanbar.
cells; resin ducts 2-4(-6), medial, the two laterals Frraxiils, and other pines, e.g., Piltis pantilt, P
always present; stele transverse-obovateto trans- pseudostrobus, P teocote, P. tlonteulinae. and P
verse-triangularin cross section; walls of endodermal arizonic'a var. stormriae; with P. ceinlbro(idesand Junti-
cells uniformly thin; vascular bundles 2, the xylem pernts flaccida on dry sites; and at higher and more
strandsconnate. Pollen cones crowded nearthe prox- mesic locations with Abies vejarii, Pseudotsuga m,etn-
imal end of a new shoot, subtendedby broad, scari- ziesii, or Cupressus hlsitanica. Phenology: Time of
ous bracts, spreading, ovoid-oblong to cylindrical, pollen dispersal has not been observed.
15-20 x 5-6 mm, yellowish, turning yellowish
Representative [Link]-
brown. Seed cones appearingon very young trees, HUILA:Near Saltillo. 6 Nov 1905. Pringle 10142 (A. BM.
subterminalor lateral, in whorls of 3-8, sometimes E. ENCB. F, GOET. K. M. MICH. MO). HIDALGO: 1.8
more, on short, stout, tenacious peduncles, at matu- km S of Jalameleo. 19(). Acosta S. & Barrioss.1. (ENCB);
along Hwy. 85 NE of [Link] Maguey Verde. 28
rity appearing sessile, long persistent, serotinous. Apr 1994. Farjon& Mejia 330 (E. FHO. HEH. K. MEXU.
Immaturecones on ca. 10 mm long, recurvedpedun- U); Ixtlahuaco. Aug 1940. M. Marteine 3463 (MEXU).
cles, ovoid, light purplish,soon light brown, matur- NUEVOLEON:Las Placetas. 15 km NW of Galeana. 27
ing in two seasons. Mature cones serotinous, nar- Apr 1985. Hlughes693. 694 (FHO. MEXU); Cerro Potosi.
13 Mar 1980. Stead & Styles 629 (ENCB. FHO. MEXU):
rowly and obliquely ovoid to ovoid-attenuate,
along rd. from 18 de Marzo to La [Link] Galeana.
slightly curved or straight when closed, with an 6 Nov 1984. Styles. Hoogh & Favela 16. 17 (FHO).
oblique base often deformed by compression of the PUEBLA:Along rd. between Huanchinangoand Necaxa. 23
expanding branchand scales, (6-)8-13(-15) x (4-) Nov 1960. Madrigal S. s.n. (ENCB. INIF). QUERETARO:
5-7 cm when open (width 3.5-5 cm when closed). Along rd. between Jalpan and Xilitla. at Km 230. 2 Mar
1978, Eguilu:.61 (ENCB. MEXU). SANLuis PO(roi: Las
Seed scales ca. 80-120, remainingclosed several to Tablas. Dec 1939. M. Martinez= 2453 (US); Mun. Xilitla. 5
many years, parting eventually or not, oblong, km N of Soledad de Zaragoza. 28 Jan 1956, Rzedowski
straightor recurvedwhen parted,the proximalscales 7149([Link]).

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194 Flora Neotropica

F o:'

~~~~~~~~~~~~C C

cone.~~~~~
!'La
'sil.C rosscin flae, . rn wth vlfru oeE edsae(w iw) ed
withartiulae wig (hreevies). Magifcaion: AB, -E X0.5 C, 40'

FIG. 49. Pinus gregii (Stead & Styles 629). A. Branchlet with cataphylls, leaf tascicles, and immatureovuliferous
cone. B. Leaf tfascicle.C. Cross sections of leaves. D. Branchwith ovuliferous cones. E. Seed scale (two views). F. Seed,
with articulatewing (three views). (Magnifications:A. B, D-F, XO.5; C, X40.)

Uses. Although locally exploited with other pines, Foresters from abroad are taking an interest in its
it is not specifically in demand as a timber tree in potentialas a forest plantationtree in othercountries.
Mexico. In many areas it has been severely depleted Like another, and probably related, "closed-cone"
by general logging and over-exploitationof forests. pine, P radiata, it seems to grow much faster in

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SystematicTreatment195

trial plots than do several other species (Dvorak & with darker apex, caducous. Vegetativebuds large,
Donahue, 1992). the terminalbuds 15-30 mm long, ovoid-acute, the
Parlatore(1868) based his descriptionof P. greggii lateralsovoid, smaller,resinous,reddishbrown, with
on a manuscriptof G. Engelmann and small parts imbricate, subulate-caudatescales (as cataphylls).
of a specimen collected by J. Gregg near Saltillo, Fascicle sheathlsthick, ca. 20 mm long, with ca. 8
Coahuila, and sent to him by Engelmann. None of imbricate, yellowish white scales, persistent, re-
this materialwas found at Fl, but at MO there is a maining long, weathering scarious, grey. Leaves in
sheet with Gregg402, bearingEngelmann'snote that fascicles of 3, in dense tufts at the ends of upturned
some of it was sent to Parlatore-this sheet is here branches,spreading,persisting3-4 years. very rigid,
designated as the lectotype. In GH there is another straightor curved, slightly twisted, 15-25(-30) cm
sheet of Gregg 402 with a small branch top with x 1.9-2.2 mm, margins bluntly serrulate, apex
leaves which can be consideredan [Link]- acute-pungentto subulate, light green to grey-green,
other of Gregg's collections at MO, originally un- often very resinous. Stomata on all faces of leaves.
named, is a mixturewith P patula; only the closed- with (10-)12-14 lines on the convex abaxial face
cone Gregg 52 belongs to P greggii, and we do not and (4-)5-7 lines on each adaxial face. Leaf anat-
know where it was originally collected. onti: Cross section transverse-triangular, with a con-
An initial investigation into cpDNA variationof vex abaxialside: cuticula and epidermisthick; hypo-
Mexican pines (Perez de la Rosa et al., 1995) indi- dermis multi-layered,pluriform,with intrusionsinto
cated a separate position from some other "hard the mesophyll;resinducts (2-)3-5, medial,occasion-
pines," including P. patula, for P greggii. Morpho- ally I internal:stele elliptic in cross section; outer
logical evidence, both of leaves and of ovuliferous cell walls of endodermis not thickened: vascular
cones, points to possible relationshipswith the Cali- bundles 2, distinctly separated. Pollen conies in a
fornian"closed-cone"pines P attenuate,P muricata, dense cluster near the proximalend of a new shoot.
and P radiata. A preliminarycladistic analysis of ovoid to cylindric. 20-25 mm long, light purplish
22 Mexican "hard pines" using 23 morphological brown, maturingto [Link]
charactersplaced P greggii on a clade with these with peltate distal part minutely striated radially,
species on the strict consensus tree (Farjon,unpubl. darkest in the centre. Seed cones subterminal,at
data). the base of a subsequent leading shoot, solitary or
occasionally in pairs, or in whorls of 3-4(-5) on the
33. Pinus coulteri D. Don, [Link]. Soc. London bole of young trees; borne on thick, short, persistent
17: 440. 1836. Figs. 50-51 peduncles,holding the cone to the branchfor as long
Pinus polnerosa D. Douglas ex P. Lawsonsubsp. coulteri as 25 years (Borchert, 1985) and retaining a few
(D. Don)E. Murray, Kalmia12:23. [Link] basal scales when it falls. Immaturecones (broadly)
[Link]:
[Link]. ovoid, on relatively long, bracteatepeduncles; with
1832-1833, T. Coulter s.n. (holotype, TCD, only an
ovuliferouscone). unguitorm umbos strongly incurvate. the proximal
umbos recurved, yellowish green, maturingin 16-
Tree,mediumsize, height to 15-25 m, dbh to 100 18 [Link] (narrowly)ovoid, massive,
cm. Trunkmonopodial, straight or curved at base. heavy, usuallyslightly oblique or curved, with a very
Bark on the lower part of the trunk 3-5 cm thick, thick (4-7 cm) woody rachis, moderatelyserotinous,
rough, scaly, divided into irregular, longitudinal 2(0-35 x 15-20 cm when open, extremely resinous.
plates and deep, longitudinal fissures, dark brown Seed scales ca. 180-220, flat, thick woody with
with black fissures, on young trees scaly, reddish thin, curved margins, widest toward the apophysis,
brownor greyish [Link] firstordervery graduallypartingover several years, purplishbrown
long, thick, spreading horizontally to more or less on the abaxial side, light brown with conspicuous
ascending, usually persistent along much of the marksof seed wings on the adaxial side. Apophyses
trunk;branches of higher orders sparse, spreading, very strongly developed, on one side of the cone
assurgentor ascending,ultimatebranchesvery thick, largerthan on the other, thick woody. sharplytrans-
upturned,forming a broad, irregular-pyramidal and versely keeled, more or less abruptly(or gradually)
open crown. Shoots multinodal, thick, rigid, rough merging into a long, uncinate umbo, up to 30 mm
with prominent, decurrent pulvini, light orange- wide, light yellowish brown or light caramel-col-
brownor purplishbrown,often glaucous. Cataphylls oured. Umbo dorsal, the strongest developed of all
large, 15-20 mm long, subulate,recurved,with cau- pines, elongate, curved, with keeled sides, 25-35 x
date apex and erose-ciliate margins, orange-brown 10-15 mm at base, ending in a sharpuncinateclaw,

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196 Flora Neotropica

ii i,;: i

i I
/

:-
\$~~~~~~~~~~~~~~~ :"'~'~~'i"":~',!
,,,1111~ ~

\~~~~~~~~~~~~~~~~~~~~~~~~?
-

?i .~~~~~~~~~-?.
c?~~~~~~~.
j"
it-It~~~~~~
A Dt
I?~~~~~~~~~~~~~~, ,'

[Link]
FIG.~
'oter Hghs&[Link]!)!ecns(op..Cos

xO.5; I. ~~~~~~ x25,C, B,

FIG. 50. Pi,nu coulteri (Huglhes& StYles183). A. Branchwith leaf fascicles and buds of pollen cones (top). B. Cross
section of leat. C. Immatureovuliferous cone. D. Basal seed scales of cone attached to branch. (Magnifications:A. D.
x 0.5; B, x25; C. x .)

concolorous with the apophysis or slightly darker. the seed margins, dolabriformto semi-ovate, (excl.
Seeds obliquely obovoid, slightly flattened, 10-18 claws) 18-30 X 12-16 mm, orange-brownto dark
x 7-10 mm, smooth, lustrous dark brown, turning reddish brown. Cotnledotns9-14.
blackish. Seed wings articulatebut effective, held to Distribution and ecology (Map 28). United
the seed by two thin, hollow appendagesenveloping States: Coast Ranges of central and S California,

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Systematic Treatment 197

FIG. 51. Pinus coulteri (Hughes & Styles 183). A. Ovuliferouscone (x 0.3). B. Seed scale (two views). C.
Seeds, with articulatewings. D. Seed and wings from upper scales. (Magnifications:A. x0.3; B-D, X0.5.)

from Contra Costa County south to San Diego it is also a tree of mixed chaparral,together with
County. Mexico: N Baja California Norte. Scattered Quercus chr3solepis, or growing on granite boulder
and rare in the interiorSierraJuarez, isolated in the formations around Laguna Juarez. Its altitudinal
coastal Sierra Juarez (Sierra Blanca), also scattered range in Mexico is from 1200-1800 m in the Sierra
and rare in the Sierra San Pedro Martir (Minnich, Juarez to 1900-2150 m in the Sierra San Pedro
1987). In California, Coulter pine is prominent in Martir(Minnich, 1987). The climate is of a Mediter-
the southernCalifornia Mixed Conifer Forest, espe- ranean type with winter rain and long, hot, dry
cially at the lower limit of this forest type, where it summers. Phenology: Pollen dispersal occurs in
merges into [Link] Baja California May-June.

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198 Flora Neotropica

at an elevation of from 3000 to 4000 feet above its

level," and described by D. Don (1836), together
with four other new species of Pinus (one of them
now recognized in the genus Abies) found by Coulter
in the same general area. The description and plate,
which appearedas intercalatedmaterial in the third
(octavo) edition of Lambert's(1832) Description of
the genus Pinus, has been shown to occur only in a
few copies of the work and to be of a date after
1836 (Little, 1949), disproving a claim of earlier
publicationby Keck (1946). Another but unknowing
discoverer may have been David Douglas, whose
1831-1832 collections of P sabiniana appearto have
become mixed with specimens of P coulteri (Griffin,
1964; Little, 1949; see also Lindley, 1840). While
Douglas's specimens were sent to the Horticultural
Society of London, those of Coulter were lent to
David Don, then librarianwith the Linnean Society
of London, and in TCD, a cone is kept which is
from Coulter's herbarium(Nelson & Probert, 1994:
fig. 43); it was used for the illustrationof the species
in Lambert'sthirdedition (see above).
The collections here studied from Baja California
Norte do not differ from those originating in the
main range of the species in California. Martinez
(1948) gave a description for the Mexican pines
similar to Kral (1993) for the United States. In the
southernCoast Ranges of California,Borchert(1985)
reportedvariationin cone serotiny relatedto vegeta-
tion types, with serotiny being more pronouncedin
Map 28. Pitnuscoulteri.
[Link] may apply to
various populations in Baja California in a similar
pattern.
Naturalhybridshave been reportedbetween Pinus
Representative specimens examined. MEXICO. BAJA
CALIFORNIA NORTE:Sierra San Pedro Martir,n.d., Galle- coulteri and P jeffreyi (Zobel, 1951; Critchfield,
gos s.n. (MEXU No. 95); Sierra Juarez, Laguna Hanson, 1966a) in California. In the Sierra Juarez, both spe-
26 May 1987. H,ughes& Styles 183 (FHO); Sierra Juarez, cies are sympatricin partsof theirranges,but hybrids
National Park"Constitucionde 1857,"6 Mar 1964. Madri- seem to be very infrequent (Zobel, 1951) and no
gal S. 1473 (INIF. MEXU); Mesa del Pinal. "Sierra del
Pinal. Baja California," Sep 1939, M. Martinez 3468 examples have come to our attention.
(MEXU); SierraJudrez,La Sierrita,NW of LagunaJuarez,
1 Aug 1976, Mora,i 23707 (ENCB. NCU. SD). 6 Jun 1985,
Thorrie& Wislira61124 (MEXU). II. Pinus subgen. Strobus
Pinus subgen. Strobus(D. Don) Lemmon, Cal. State
Uses. Pinus coulteri has no particular commercial
Board Forest. Bienn. Rep. 2: 69, 79. 1888; Lem-
value as a timber tree and its seeds are not harvested
mon, Cone-Bear. Trees Pacif. Slope, ed. 3, 20.
for consumption. In Mexico it rarely forms extensive
1895. (spp. 34-47)
stands. Locally, some trees may be harvested where
they occur nearby or mixed with Pinus jeffreyi, with Pinus [group]StrobusD. Don, [Link]. Nepal. 54. 1825;
which Coulter pine may occasionally hybridize (see Pinus [group] Strobus Sweet, Hort. Brit.. ed. 2, 475.
below). 1830, nom. nud.; Pinus sect. Strohus (D. Don) Spach.
Hist. Nat. Veget. Phan. I1: 394. 1842; Pinus sect. Peuce
This remarkable pine was found in 1832-1833 by
Grisebach,Spic. Fl. Rumel. 2: 347. 1844; Pinus subgen.
Thomas Coulter in the Santa Lucia Mountains, near Peuce (Grisebach) Endlicher. Syn. Conif. 138. 1847.
Cone Peak, California, "within sight of the sea and pro syn.: Pinus sect. Haploxylon Koehne. Deut. Den-

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Systematic Treatment199

drol. 28. 1893; Pinus div. TenuisquamaeMasters, J. brown,weatheringgrey; on young trees and branches
[Link]. London,Bot.35: 569. 1904;Pinussubgen. smooth, greenish grey. Branches of first order long,
Haplovxlon(Koehne)Rehderin Bailey,[Link]. slender,
302. 1923;[Link]-Duplan, spreading horizontally, the lower branches
Trav. Lab. Forest. Toulouse T. 2, sect. 1, 4, art. 1: 92. somewhat drooping; of second and higher orders
1950, nom. nud.; Pitus subgen. Malacopit's Ishii, slender, spreadingand/or [Link] pyrami-
[Link]. Sci. Reports2: 112, 124. 1952,sine dal to broadconical, with branchesdown to 2/ or 3/
diagn. lat.; Pinus subgen. Cemlbrapinus de Ferre, of tree
height. Shoots mostly glabrous or current
[Link]. 236: 228. 1953;Pinus shoots brownish puberulent,grooved; pulvini
[Link] de Ferre,[Link] year's
Acad. Sci. 236: 228. 1953. Holotypespecies. Pinus prominent, short [Link] small, 5-10
strohusLinnaeus,Sp. P1.1001. 1753. mm long, subulate,with erose margins,darkbrown,
DucampopinusA. [Link]. Bot. Appl. d'Agr. Trop. soon deciduous. Vegetativebuds small, hidden in leaf
24: 30. 1944; Pinus subgen. Ducampopinus(A. Cheva-
tufts, the terminal buds 8-15 x 5-8 mm, ovoid-
lier)Little& Critchfield,
[Link].U.S. [Link].
991:5. 1966;[Link] Lecomte, oblong, the laterals smaller, ovoid, not or only
[Link].27: 191. 1921. slightly resinous; the scales subulate-linear,acute,
[Link]. Murray, Kalmia13: 11. 1983. free, dark purplish brown. Fascicle sheaths 15-20
[Link] Balfourin A. Mur- (-25) mm
long, with ca. 5-7 loose, oblong, orange-
ray,Bot. Exped.Oregon8: 1. 1853. brown or red-brown scales, deciduous. Leaves in
Pin,s [Link]. Murray, Kalmia13: 13. 1983.
[Link] Wallichex D. Don fascicles of 5, very rarely6, in dense, lax tuftstoward
in Lambert,[Link]. 8, 2: p. s.n. inter144et the upturnedends of ultimatebranches,persisting2-
145, t. 79. 1832. 3 years, straightor slightly curvedor twisted, spread-
[Link].Phytologia76: 77. at 40-60? from the shoot but not drooping,(8-)
1994. Holotypespecies. PinlusnelsoniiShaw,Gard. ing
Chron..ser.3, 36: 122. 1904. 10-15(-18) cm X 0.7-1 mm (-1.5 mm on vigorous
coning shoots), margins (weakly) serrulate, apex
Derivation:From type species, P strobus. acute, green or grey-greenwith two whitish stomatal
Bark thin and smooth up to considerable age in [Link] the two adaxial faces only, in two
several species, eventually variously furrowed and dense bandsof 3-6 intermittentlines. Leaf anatomy:
scaly, exfoliating. Long shoots uninodal, smooth, Cross section triangular,with roughlyequal sides, the
with (very) short decurrent("non-decurrent"of au- abaxial side slightly convex; hypodermis uniform,
thors) leaf bases (pulvini). Ccataphy'lls early decidu- mostly with a single layer of cells; resin ducts 2-4
ous, usually smaller than 10 x 4 mm. Fascicle (-7), external, the largest number usually on the
shleathsdeciduous, rarely persistent (in P. nelsontii). abaxial side; stele terete; outer cell walls of endo-
Leaves epistomatic or amphistomatic, with entire, dermisnot thickened;vascularbundle I. Pollencones
remotely serrulate,or serrulatemargins;resin ducts crowded at the proximalend of a new shoot, spread-
external; stele with thin outer cell walls of endo- ing, subtended by reddish brown, scarious bracts,
dermis, containing a single vascular bundle. Pollen ovoid to short cylindrical, 7-10 mm long, yellow,
'contes ovoid to short cylindrical when mature, ar- [Link] cones towardthe end of
ranged along the proximal part to 2-/ of new long main branches,subterminal,axillary to the terminal
shoots; microsporophyllssmooth distally, with entire bud, solitary or more often in pairs or whorls of 3-
upper margin. Seed contes deciduous, irregular or 4 on short (to 25 mm) peduncles which usually fall
cylindrical. Seed scales few to numerous, with with the cone. Immaturecones elliptic to cylindrical,
mostly simple phyllotaxis; apophysis transversally straightor curved, more or less erect, purple,turning
keeled or not; umbo dorsal or terminal, lacking a green, maturingin two seasons. Maturecones short
spine or prickle. Seeds with an adnate wing shorter pedunculate,pendulous,cylindrical,curved, tapering
to longerthanthe seed, or vestigial/articulate,leaving towardapex, usually>3 times longerthanwide when
a wingless matureseed. open, (10-)15-40(-50) x 7-15 cm when open. Seed
scales ca. 100-150, readily parting to release the
34. Pinus ayacahuite Ehrenbergex Schlechtendal, seeds, spreadingat 40-800 from the axis, thin, flexi-
Linnaea 12: 492. 1838. ble or more rigid, straight, the proximal scales re-
curved, transverselynearly flat or concavo-convex,
Tree, tall, height to 40-45(-50?) m, dbh to 150- long (5-7 cm), with parallel to diverging sides, the
200 cm. Trunkmonopodial,erect, columnar,straight, distal scales narrowing, (abaxially) rugose or
terete. Bark rough, scaly, divided into small, rectan- grooved,dull brownor darkreddishbrownabaxially,
gular scales, exfoliating with small flakes, grey- with light brown marksof seed wings on the adaxial

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200 Flora Neotropica

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

'I~~~~~~~~~~~~~~~?

'A. Ma 9 iu wuha a.(wai~le(ice) ivc/nevr eic/rn? (trianle)

SON

2(i(rn
Map

side. Apophysis thin or thick, irregularly trullate, del Monte, Omitlan de Juarez, near Hacienda de
recurved or reflexed toward the apex, or the apex Guerrero, C. A. Ehrenberg 647, Aug 1837 (holo-
curved up below the umbo, longitudinallygrooved, type, B, destroyed; lectotype, HAL, here desig-
dull light brown, very resinous. Umbo terminal, nated; isolectotypes, GOET, MO, US).
broadly triangular, obtuse, 5-10 mm wide, grey, Figs. 52-53A
usually covered with resin. Seeds in two cup-like
depressions at the base of the scale, (broadly) ob- Pitnus lon-pedrii Roezl. Cat. Grain. Conif. Mexic. 31.
ovoid, laterally flattened, 8-15 x 6-9 mm, dull 1857. Type. Mexico. Mexico: Valle de Mexico, 1857,
brown with black spots, usually equally developed Roezl s.m. (ex herb. Lindley 3155) (lectotype. CGE,
but sometimes of slightly differentsize on one scale. here designated; isolectotype, FI).
Pinns hamuta Roezl, Cat. [Link]. Mexic. 32. 1857.
Seed wings adnate, (2-3.5 x) longer than the seed,
Type. Mexico. "SierraMadre,"Roezl s.n. (lectotype, P.
oblique, with a curved side or broadtruncate,20-35 here designated).
x 8-12 mm, light brown with dark striations or Pinus avicahluite [Link] Schltdl. [Link] Silba,
dark brown. Number of cotyledons not observed. Phytologia 68: 47. 1990. Type. Mexico. Oaxaca: Be-
Distribution (Map 29). Mexico: From central tween San Miguel and Tres Cruces. 9 Feb 1945. [Link]-
der 641 (holotype. NY; isotypes, F, MEXU, MICH).
Mexico southwardto Guatemala,Honduras,and El
Salvador.
Tree, tall, height to 40-45(-50?) m. Leaves in
fascicles of 5, very rarely 6, straight or slightly
Key to the varieties of Pinus ayacahuite curved or twisted, thin, lax, (8-)10-15(-18) cm X
i. Apophysisof seed scales thin. trullate,apex re- 0.7-1 mm; epistomatic, stomatal bands whitish, ab-
curvedbutnotreflexed:seedwing2-3.5 x longer axial face green or greyish green. Seed coptes pendu-
34a. [Link]
thanseed.........................................
I. Apophysisof seed scales thick,extremelyelon- lous, cylindrical, curved, tapering toward apex, (10-)
gated,abruptlynarrowed,reflexed;seed wing up 15-40 cm long, 7-15 cm wide when opened. Seed
to 2 x as long as seed.......................... 34b. var. vieitchii scales ca. 100-120, thin, flexible; apophylsis thin,
trullate, usually recurved, rarely reflexed at apex.
34a. Pinus ayacahuite Ehrenbergex Schlechtendal Seeds 8-10 x 6-8 mm. Seed wing adnate, 20-35
var. ayacahuite. Type. Mexico. Hidalgo: Mineral x 8-12 mm, tapering toward the apex.

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Systematic Treatment 201

F iS
r Bt 1X W

[Link] ja it .

FIG. 52. Pinus auacahuite var. avacahuite (A-C, Stead & Styles 192: D, Tucker1203). A. Branchwith leaf fascicles.
B. (a) leaf fascicle with (b) sheath scales and (c) remotely serrulateleaf margins.C. Cross section of leaf. D. Ovuliferous
cone. E. Seed scale (two views). F. Seeds with adnate wings. (Magnifications:A. B. D-F. x0.5; Be, x 10 C. x50.)

Distribution and ecology. Mexico: In Hidalgo, mala, Honduras,and El [Link]

Puebla, Veracruz,Tlaxcala, Mexico, Morelos, Mi- trees in mixed montaneconifer forest on mesic sites,
choacin, Guerrero,Oaxaca, [Link] in Guate- or growing in groups or small groves; reaching its

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202 Flora Neotropica

. ...,.
~~~~~~~~~~~~~~~~~~.~;..

?: ?
.,?t? !
,,~

?' ,, :?~
.;,.,~~~~~~~~~~~~~~~~~~~~~~~~~~~:',
.:i.Y
.:...----.'......?
;.?.~.~ .

; c ~ ~ L?!/;~~~~~~~~~~~~~~.
t.J~I!~iJ

i- "i]...
:~.I?-?;

..xQ: ,.~..,
':P'IP~~~~~
?~~~~~~~~~~~~~~~~~~~~~~~~.-.':::5i
~~-.. ~ ,
:.1~~~ ~~
, ~ ,,

~~~~~~~~~~~~~~~~~~~~~.:
.
%.

?1G 5
.[Link] coe. P avcau!. ar.'m/# $'/s&Kp//.[Link]'
avachu~v~. v#?h/(Pee. deaRa /88.
.,.(anfctinx

greatest extent in Chiapas and W Guatemala. The tude and altitude; in south-central Mexico it is in
best stands are on loamy, well-drained soils. Its May.
altitudinal range is (1500-)1900-3200(-3600) m.
Representative specimens examined. MEXICO. CHIA-
Phenology:Time of pollen dispersal varies with lati- PAS:Motozintlade Mendoza, Cerro Mozotal. 30 Dec 1972.

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Systematic Treatment 203

Breedlove & Thorne 31157 (MO, NY); Comitan, 13 Mar 34b. Pinus ayacahuite Ehrenbergex Schlechtendal.
1993, Higman, Padilla & Styles 18 (EAP, FHO, HEH, var. veitchii (Roezl) G. R. Shaw, [Pines Mexico]
MO); San Crist6balde las Casas, May 1939, M. Martinlez Publ. Arnold Arbor. 1: 10. 1909.
2336 (NY); Pasital, Jan 1937, Matuda 293 (MICH. NY): Fig. 53B
2 km from Ocosingo towardComitdn,31 Mar 1974, Styles Pinus veitchii Roezl, Cat. [Link]. Mexic. 32. 1857.
114 (FHO); 40 km from San Crist6balde las Casas along Type. Mexico. Mexico/Puebla:E slope of Mt. Popocate-
Pan American Hwy., 18 Mar 1972, Styles & Kemp 117 petl, ca. 3500 m, Roezl s.n. (not found in CGE, Fl. or
(EAP, FHO). GUERRERO: Chichihualco,9 km S of Yerba P). Neotype: "Ridge between Mts. Popocatepetl and
Santa, 16 Jul 1985, Lorea 2100 (FHO, UNAM); Chichihu- Iztaccihuatl,"Nov 1896, Pringle s.. (neotype, A, a
alco, 9 km W of Carrizal, I Dec 1963, Rzedowski 18013 cone illustratedin Shaw, 1909 (t. 5. fig. I). here desig-
(ENCB, MICH). HIDALGO: Omitlande Juarez,near Haci- nated).
enda de Guerrero,Aug 1837, Ehrenberg 645 (MO); La Pinus popocatepellii Roezl, Cat. [Link]. Mexic. 33.
Victoria, 7 km NE of Metepec, 18 May 1982, Herndndez 1857. Type. Mexico. Mexico/Puebla: E slope of Mt.
M. 7258 (MEXU); near Omitlan, Sierra del Lucerno, Jan Popocatepetl,ca. 3500 m, Roezl s.n. (not found in CGE.
1940, M. Martinez2188 (MEXU); El Chico, Jan 1941, M. FI, or P).
Martinez 2341 (NY). Mxlico: Mt. Popocatepetl, 18 Nov Pinus loudoniana G. Gordon & Glendinning, Pinetum
1896, Pringle s.n. (A, F). MICHOACAN: Paracho. N of 230. 1858; Pinus avacahuite Ehrenbergex Schlechten-
Uruapan,Las Palmas, Los Reyes (Palicutin), 15 Feb 1980, dal subsp. loudoniana (G. Gordon & Glendinning) E.
Klaus s.n. (WU). OAXACA: Benito Juarez National Park, Murray,Kalmia 13: 21. 1983; Pinus aalcahuite Ehren-
20 Mar 1993, Higmann,Padilla & Styles 45 (EAP, FHO, berg ex Schlechtendal var. loudoniana (G. Gordon &
HEH, MO); 168 km along rd. from Oaxaca to Puerto Glendinning) Silba. Phytologia 68: 47. 1990. Type.
Angel, in Sierra Madre del Sur, 18 Feb 1985, Hughes Mexico. Mexico/Puebla: Mt. Popocatepetl; possibly
548 (FHO). PUEBLA: Esperanza,1907, Purpus 4381 (UC). based on material collected by Roezl but no original
VERACRUZ: Calcahualco, 3.5 km N of [Nueva] Vaqueria, materialhas been found.
I May 1983, Nee & Taylor27054 (CAS. F, K, NY, XAL);
Cerro Verde, I km N of Carbonera,27 Jan 1984, Nee & Tree, tall, height to 40-45(-50?) m. Leaves in
TaYlor29084 (F, FHO, NY, XAL). fascicles of 5, very rarely 6, straight or slightly
GUATEMALA. EL QUICHE:San Miguel Uspantin,
curved or twisted, thin, lax, (8-)10-15(-18) cm X
May 1892, Heyde & Lux 3156 (A, M, MO, NY). GUATE.
NAI,A: Near Antigua. Jun 1882, Lehmant 1677 (BM). 0.7-1 mm; epistomatic, stomatalbands whitish, ab-
JALAPA:15 km E of Mataquescuintla,4 Mar 1984. axial face green or greyish green. Seed cones pendu-
McCarter & Styles 122 (FHO); Mt. Miramundo.3 Dec lous,
broadly cylindrical, curved, often very large,
1939, Stevermark 32703 (F). QUEZALTENANGO: Quet-
Bernoulli & C(ario 1095 15-50 cm long, 10-15 cm wide when opened. Seed
zaltenango. May 1878, (GOET);
Volcan Santa Maria, 27 Jul 1934, Skutch 868 (A. F). scales ca. 100-150, not flexible; apophysis thick,
SOLOLA:Cumbre de Argueta, May 1896, Bernoulli & trullate, with an extremely elongated, abruptlynar-
Cario 1094 (GOET); between Novala and Nahuala,3 Apr rowed, reflexed distal
part and sometimes an up-
1993, Higman, Padilla & Styles 64 (EAP, FHO, HEH, turned Seeds 10-15 X 7-9 mm. Seed wing
MO). TO1ONICAPAN: Ca. 30 km E of Totonicapan,8 Aug apex.
1958, Beaman3121 (A, ENCB. NY, US); N of Totonicapin adnate,up to 2 X as long as the seed, oblique, broad
at ParajeChaquisis,6 May 1993, Higman, Padlilla& Styles truncateor taperingto a roundedapex.
I (EAP, FHO, HEH, MO); Cerro MariaTecum, Jan 1966, Distribution and ecology. Mexico: In Guanajuato,
Molina R. et ol. 16401 (EAP, F, NY); along rd. from
Quer6taro, Hidalgo, Veracruz,Mexico, Puebla,Tlax-
Totonicapan to Quetzaltenango, 16 Feb 1978, Stead & and Michoacan. Forming emergent
Styles 192 (FHO); Cerro Maria Tecum, ca. 15 km E of cala, Morelos,
Totonicapan, 16 Dec 1962, Williams et al. 23142 (A, EAP. trees in mixed montane conifer forest on mesic sites,
F, NY, US); Cerro Maria Tecum, 5-10 km W of Los or growing in groups or small groves. In much of
Encuentros,24 Dec 1972, Williamset al. 41725 (EAP, F. its range it is more or less sympatric with var.
NY). and hybridizationprobably occurs. The
HONDURAS. LA PAZ:Las Trancas, 15 km from Opa- ayacahlite
toro, Montafa Pacayal, 28 Jun 1982, Hughes 224 (FHO. high-altitude stands are subject to snowfall and frost
K). LEMPIRA:Don Tomias,Celaque National Park. 11 Nov in some [Link] altitudinalrangeis (2000-)2500-
1991, Hawkins 69 (EAP, FHO, MO, TEFH); Filo Seco, 32(X)(-3600) m.
trail from Campo Naranjo to El Sucte, 14 May 1992,
Hawkins 324 (EAP, FHO, MO, TEFH); Celaque National Representative [Link]-
Park, Mt. Celaque, trail from camp Naranjo to El Sucte, NAJUATO:Mtns. ESE of San Jose [Link]. 5 km W of
14 May 1993, Hawkins & Allan 324 (HEH); Celaque Cerro Zamorano,Mesa del Gato, 27 Apr 1949, McVaugh
National Park, Mt. Celaque, II km SW of Gracias, camp 10400 (MEXU, MICH, US). Mexico: 2 km E of San
Naranjo, 14 May 1992, D. Mejia 93 (EAP, FHO, HEH, Rafael, at mouth of Canada del Negro. 23 Sep 1961,
MO, TEFH). SANTABARBARA: Mt. Santa Barbara,4 Apr Beuaman& Andresen 4523 (GH); San Rafael, Cascada
1981, Hughes 4 (FHO), 2 Apr 1982, Hughes 88 (FHO, K). de los Diamantes, 21 Oct 1962, Galicia X. 136 (US);
EL SALVADOR. CHALATENANGO: Los Esesmiles, E Amecameca, Buenavista. 8 Nov 1992, Gardner & Knees
of La Palma, 6 May 1959, Allen & van Severen 7312 (A, 5339 (E); Tlalmanalco, I km E of San Rafael, 12 Dec
EAP, F, K, NY, US), 17 Mar 1942, Tucker1067 (EAP, F, 1991, Pereztie la Rosa 1898 (FHO, NY). Perezde Ia Rosa
K, MICH. NY, P, US), 4 Apr 1942. Tucker1203 (US). 1899 (FHO, MEXU). MORELOS: Zempoala,Jan 1940. M.

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204 Flora Neotropica

Martinez 2189 (US). PUEBLA:Ca. 15 km N of Tiaxco, Rosa, 1993). Variationin leaf length and cone size
near Hwy. 119, 27 Apr 1994, Farjon & D. Mejia 326 (E. can still be great(Molina, 1964), but the morphology
FHO, HEH, K, MEXU, U); San Andr6s, Sep 1939, M. of the seed scales is very similar all through the
Martlnez 3651 (F); Volcan Iztaccihuatl,15.9 km W of Paso
de Cort6son [Link] BuenaVista.26 Feb 1988,Rezlicek& range from Oaxaca to [Link] treat P. aaca-
[Link] 8120 (NY). huite here in this more restrictivesense, well-charac-
terized by the long seed wings which Schlechtendal
Uses. Pinus ayacahuite and its variety P ayacahu- so aptly described in the protologue.
ite var. veitchii are both very importantand highly The observationby Silba (1990) of fascicles of 3
sought-aftertimbertrees as they are the tallest grow- leaves in the type specimen (Alexander 641, NY) of
ing and most regular shaped representativesof the P avacahluite var. oaxacana Silba is an obvious
so-called soft pines in Mexico and Mesoamerica. [Link] distinctcharacterin the cone scales
Especially in Chiapas,easternGuatemala,and north- was mentioned,no cone could be found to belong to
ern El Salvador, many of the original stands have this collection at NY, and thus it is not known from
alreadybeen seriously depleted(Dvorak& Donahue, what specimens Silba derived his cone characters.
1992). Roezl (1857) describedPinus veitchii in his Cata-
Only two varieties of Pinus ayacahuiiteare here logue des grainestde coniferes nexicains, among
recognized, based mainly on quantitativecharacters numerousother "new species," from centralMexico,
in the seed scales and seed wings. Schlechtendal on the eastern slopes of Mt. Popocatepetl at an
(1838) described Pirnusayacahuite from material altitudeof ca. 3500 m. No original materialhas been
collected by Carl Ehrenbergin August 1837 near found among the collections of Roezl's material in
Mineraldel Monte. The large cones are described in CGE, FI, and P. In P there are some large cones in
some detail in the protologue and mention is made glass jars lacking any label, which could well have
of the long seed wings (shaped like the forewings of come from him, but it cannot be proven. A neotype
a narrow-wingedmoth). The holotype in Berlin has seems to be necessary in this unusualcase of typifi-
been destroyed, and with the isotypes seen there are cation of an accepted pine taxon.
no cones. As more materialwas collected duringthe
19th and early 20th centuries, it became apparent 35. Pinus lambertiana D. Douglas, Trans. Linn.
that increasedvariationwas to be found in the cones Soc. London 16: 500. 1827. Type. United States.
of otherwise rather similar trees further west and
Oregon: Aug 1825 (or Oct 1826), Douglas s.n.
north in Mexico. This variation leads to forms ap-
(lectotype, K, 2 foliage specimens numbered (I)
proaching Pinus flexilis of the Rocky Mountains, and with Douglas's remark:"Gum from a Larch
which had been named P strobiformris and P. flexilis tree, eaten by the natives, Douglas," here desig-
var. reflexa(or P reflexa).This led Shaw (1909) first nated). Fig. 54
to the conclusion that P. strobiformisis only a variety
of P ayacahuite with shorter seed wings, and then Pinus laibertiana D. Douglas var. martirensis Silba,
to his (Shaw, 1914) sinking it into synonymy with Phytologia68: 52. 1990. Type. Mexico. Baja California
Nore: Sierra San Pedro Martir,La Encantada,22 Sep
the latter species, leaving only the suggestion to 1930, 1. L. Wiggins & Demaree 5013 (holotype, NY;
recognize two varieties, var. brachyptera and var. isotype, GH).
veitchii, based mainly on a north-southdine in the
length of the seed wing. This latter view was ac- Tree, medium to tall, height to 30-40 m, dbh to
cepted by Martinez(1948), but Perry(1991) retained 80-120 cm. Trunkstraight,terete, the bole often free
the name P strobiformnis for the form with generally of branches '/2 or more of its length. Bark thick,
shorterleaves and shortercones only. rough, scaly, on the lower partof the trunkbreaking
When Carvajaland McVaugh(1992) agreed with into increasingly large, irregular to longitudinal
Shaw (1914), they obviously looked from "Nueva plates separatedby deep fissures,cinnamon-to grey-
Galicia"north,but not south, althoughthey acknowl- brown, on young trees and branches thin, smooth,
edged that in their area (west-central Mexico) most greyish green, turningbrown. Branchesof firstorder
of the "complex of P ayacahuite" is absent, espe- often widely spaced, long, spreading horizontally,
cially the species as to its type. The undoubtedly curved downward toward the ends; branches of
clinal variation in cone morphology diminishes in higher orders numerous, considerably shorter than
the southern parts of its range, especially in the the main branches, spreading or drooping, the ulti-
development of the seed wing, which is invariably mate branches assurgent. The crown is broadly coni-
long and effective in P ayacahuite [Link].(Perezde la cal, lobed, in old trees often flat-toppedfrom wind-

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Systematic Treatment 205

[Link] slender,flexible, puberulentat first but relatively short compared to size of seed scales,
soon glabrous, rough with small pulvini, orange- broadly obovoid to dolabriformor truncate, 20-30
brown turning light grey. Cataphylls small, triangu- x 12-15 mm, light brown. Cotyledons 11-18.
lar,scarious,with thin, hyaline margins,soon decidu- Remarks on morphology. The above description
ous. Vegetativebuds ovoid-oblong, the terminalbuds pertainsto Pinus laitbertiana in Mexico only. Tree
5-10 x 4-6 mm, the laterals smaller, not resinous; size here is considerably shorter (due to climatic
the scales triangular,orange-brownwith hyaline mar- conditions) than in the Sierra Nevada of California
gins. Fascicle sheaths 10-15 mm long on young or the Cascade Range of Oregon, where this species
fascicles, with 5-8 thin, loosely imbricate,orange- grows to be the largestof all extant pines. Maximum
brown scales, soon deciduous and absent when the height in the SierraSan Pedro Martiris given as 40
leaves are full-grown. Leaves in fascicles of 5, in m by Minnich (1987); referencesto tree sizes of 60
small, dense tufts toward the ends of branches,per- m or more in Martinez (1948) and Perry (1991)
sisting 2-4 years, straight, slightly twisted, lax, undoubtedlypertainto the species outside of Mexico.
spreadingat ca. 45" from the shoot, (3.5-)4-8(-10) A maximum cone size of 56 cm has also been
cm x 0.8-1.5 mm, with remotely and sometimes reportedfrom the UnitedStates (Kinloch & Scheuner
scarcely serrulate margins, apex acute-acuminate, in Burns & Honkala, 1990), and such large cones
green to glaucous-green. Stomata on all faces of have not been found in Mexico.
leaves, in 2-4(-5) intermittentlines on the abaxial Distribution and ecology (Map 30). United
face and 3-4 lines on each adaxial face. Leaf anat- States: In Oregon and California mainly along the
omny: Cross section triangular,with equal sides; hypo- Cascade Range and in the SierraNevada, in S Cali-
dermis unilayered or near the margins with 2-3 fornia scatteredin high [Link]: In Baja
layers of cells; resin ducts 2-3, external, rarely I CaliforniaNorte, its only locality is the higher part
internal;stele terete; outer cell walls of endodermis of the SierraSan PedroMartir,which is the southern-
not thickened; vascular bundle single. Pollen cones most occurrenceof the species. Its altitudinalrange
crowded near the proximal end of a new shoot, there is 2200-2800 m, where it occurs frequentlyas
forming more or less elongated "spikes,"ellipsoid- a constituent of mixed conifer forest on sites with
cylindric, 10-15 mm long, yellowish when grown to deepest soil, often along intermittentstreams. Other
full size. Microsporophyllspeltate, rounded at the conifers are Abies concolor, Pinus [Link]-
distal end, ca. 1 mm wide, smooth with entire mar- rayana, and P jeffreyi. Annual precipitation is a
gins. Seed cones concentrated at the ends of main modest 500 mm and winter snows and frosts are
branches,subterminal,solitary or in whorls of 2-4, common. Phenology: Time of pollen dispersal is
erect on 5-12 cm long, 15-20 mm thick, bracteate June-July.
peduncles until the beginning of the second season,
Representative specimens examined. UNITED
becoming pendulous. Immature cones cylindrical, STATES. OREGON:"In Northern California, Lat. 43?,
25-50 mm long at the time of fertilization,growing Long. 123 West in sandy soils." Douglas s.n. (K. original
to twice that size by the end of the first growing material?).
season, at firstpurplish,then greenish, very resinous, MEXICO. BAJACALIFORNIA NORTE:SIERRASANPE-
DROMARTiR:18 May 1893, Brandegee s.n. (UC); La
maturing in two seasons. Mature cones narrowly Corona,31 Oct 1947,
Duffield3 (MEXU,UC); I kmS of
cylindrical when closed, straightor slightly curved, observatory, 2 Oct 1992.Farjon263 (U, UC);La Encan-
broadly cylindrical when opened, then 25-45 x 8- tada,25 Apr 1987,Hughes& Styles172. 173 (FHO):ca.
14 cm. Seed scales ca. 110-130, readily opening, 15 km W of observatory.1 Jun 1974. McGill & Pinkawa
P12325 NW of La Corona,31 Aug [Link]
patent or slightly reflexed, the basal sterile scales 11278 (NY); CerroVenadoBlanco.15 Sep [Link]
often more reflexed, angular-obovateto obtrullate, 15647 (SD);
(SD).
up to 40 mm wide and 4-5 mm thick, purplishbrown
abaxially, reddish brown with light brown marksof Uses. Due to its restrictedrange in Mexico, this
seed wings and shallow seed cavities on the adaxial species is of little economic [Link] remote-
side. Apophysis triangularto obtusely rhombic, 5-8 ness has safeguardedthe Mexican population from
mm thick at base, straight, smooth or radially stri- extensive logging in the past; a road onto the high
ated, yellowish brown, often with yellow resin clus- plateaus has only recently been built. Most of the
ters. Umiboterminal, obtuse-triangular,ca. 10 mm stands are now within the boundariesof the Sierra
wide, concolorous or lighter than the apophysis. San Pedro MartirNational Park.
Seeds obovoid or obliquely obovoid, (10-)12-15 The earliest collection of Pinus lanbertiana south
(-18) x 6-10 mm, dark brown. Seed wings adnate, of the U.S.-Mexican border was made by T. S.

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206 Flora Neotropica

cone. F. Seed scale (two views).G. Seeds, with adnate wings. (Magnifications:A. B, D-G, X 0.5: C. X 30.)

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Systematic Treatment 207

cies in northernBaja California and confirmed the

isolated disjunction of P lambertiana. Silba (1990)
described it as a distinct variety, with "more pec-
tinately arranged,thicker,curved needles 41-67 mm
long by 0.9 mm wide," based on only two studied
collections, the type cited above and McGill & Pin-
kava P12325 (NY). The maximum leaf length mea-
sured on the type specimen is 82 mm; none of the
characters are outside the range found in material
from the United States.
The amphistomaticleaves of P liambertianadistin-
4 guish it from pines of the P ayacahuitelP. strobi-
formis pair and seem to link it with the P montlicola/
P strobus pair, which does not support the scheme
of relationships proposed by Martfnez (1948). Its
cone morphology takes it close to P flexilis, as
does its genetic affinity based on allozyme analysis
(Krutovskii et al., 1994). Further study, including
cpDNA analysis, to investigate these relationshipsis
highly desirable.

36. Pinus flexilis E. James var. reflexa Engelmann

in Rothrock, Rep. U.S. Geogr. Surv., Wheeler, 6,
Bot.: 258. 1879. Fig. 55
Pinusreflexa(Engelmann) [Link].(Craw-
Engelmann,
fordsville) 7: 4. 1882; Pinus aYacahuiteC. G. Ehrenberg
ex Schlechtendalvar. strobiformisJ. G. Lemmon.
Handb.W. [Link]-bearers, ed. 2, 4. 1892;Pinus
acacahuiteC. G. Ehrenberg ex [Link]-
flexa (Engelmann)A. Voss, Mitt. [Link].
Map 30. Pinus lamhbertiana(circles); P. cembroides Ges. 1907(16):95. 1907;PinusflexilisE. Jamessubsp.
subsp. lagunae (triangles). reflexa(Engelmann) E. Murray.Kalmia12: 23. 1982.
[Link]:SantaRita Mtns..E of
Tucson,13 Aug 1874, Rothrock654 ([Link]
3377660(foliage)andMO 1635443(cone),designated
Brandegee in May 1893, about 200 km S of the
neareststands in [Link] species was discov- here).
ered by David Douglas in August 1825, or perhaps Tree, small to medium, height to 10-15(-20) m,
October 1826, in Oregon and was formallydescribed dbh to 100-150 cm. Trunkmonopodial or with 2 to
in a letter by Douglas, read on 6 November 1827 by several stems, straightor contorted, branchingvery
the Secretary on the occasion of a meeting of the low. Bark thin, rough and scaly, breakinginto small,
Linnean Society of London and subsequently pub- scaly plates, brown, weatheringgrey, on young trees
lished in the Transactions (Douglas, 1827). Original and branches smooth and greyish white. Branches
materialis in K, in the herbariaof both J. D. Hooker of first order thick, contorted, or long and slender,
and G. Bentham. Exactly when Douglas collected ascending or spreading, of higher orders slender,
this material is not known, and the location is like- flexible, [Link] initially puberulent,soon
wise vague. At any rate, "Lat. 43?" is not in Califor- glabrous, with short, non-decurrentpulvini, grey-
nia but in [Link] small branchletswith foliage green to grey. Cataphylls 5-10 mm long, subulate,
are preserved and no cones (as cited in the pro- scarious, brown, with erose-hyaline margins, soon
tologue). On the sheet ex herb. J. D. Hooker is deciduous. Vegetativebuds ovoid or subglobose, the
mounted a third, later collection by J. M. Bigelow terminal buds 8-15 mm long, the laterals smaller,
(in 1853-1854), which is not type material. The resinous; with imbricate,appressed,subulate, brown
other two Douglas pieces are here designated as the scales. Fascicle sheaths ca. 15 mm long, with 6-7
lectotype. loosely imbricate, orange-brown scales, deciduous
Minnich (1987) established the extent of this spe- and absent in second year's fascicles. Leaves in

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208 Flora Neotropica

E F <:
FIG. 55. Pinus flerilis var. reflera (A-C, E. W Nelson 6043; D-F. E. [Link] 6736). A. Branch with leaf fascicles.
B. Leaf fascicles. C. Cross section of leaf. D. Ovuliferous cone. E. Apophysis of seed scale (two views). F. Seed (two
views). (Magnifications:A. B, D-F. x0.5; C., x30.)

fascicles of 5, in dense, short tufts toward the ends glaucous to white stomatal lines on both adaxial
of branchlets, persisting (3-)5-6 years, straight or faces. Stomata:Leaves epistomaticor weakly amphi-
slightly curved, sometimes twisted, lax, spreadingat stomatic, with 0-1(-2 intermittent)lines of stomata
30-50? from the shoot, (5-)6-9 cm X 0.8-1.2 mm, in grooves on the abaxial face and 2-4 (intermittent)
with remotely serrulate to nearly entire margins, lines on each adaxial face. Leaf anatoImy: Cross
acute to acuminate, dark green on the abaxial face, section triangular;hypodermis unilayered or some-

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Systematic Treatment 209

times with a few extra cells on the abaxial side and after a cold spring. Seed dispersal is largely effected
near the margins; resin ducts 1-3, external on the by birds.
abaxial side; stele terete; cell walls of endodermis
Representative specimens examined. UNITED
thin; vascular bundle single. Pollen cones crowded STATES. ARIZONA:Sonoita Valley (S Arizona). 1874.
at the proximal end of a new shoot, spreading, Rothrock1001 (MO).
subtended by light brown bracts, ovoid-oblong, 6- MEXICO. CHIHUAHUA: "SierraMadreMts.."Jul 1899.
10 mm long, yellow, maturingto yellowish brown. Nelson 6043 (A, US). COAHUILA: GeneralCepeda. 20 Apr
1902, Nelson 6736 (A, US). NUEVOLEON: Cerro Potosi.
Microsporophyllspeltate, the distal part cordate to 13 Mar 1980, Stead & Styles 250 (MEXU).
rounded,smooth, ca. I mm wide. Seed cones subter-
minal, solitary, in pairs or in whorls of 3(-4), on Uses. Pinus flexilis var. reflexa is not used for
stout, to 15 mm long peduncles which fall with timberdue to its poor shape (from a forester'sview-
the cone. Immature cones erect, ovoid-oblong to point) and its rarity and inaccessibility. No other
cylindrical, greenish or purplish, maturing in two productsare derived from it.
seasons. Mature cones subpendulous, cylindrical, Pinusflexilis and P. strobif)rmis form a polymor-
straightor slightly curved when closed, ovoid-cylin- phic dine in the cordillera of North America, ex-
drical or broad-cylindricalwhen opened, then 10-15 tending from SW Alberta well into N Mexico. At
x 4-6 cm. Seed scales ca. 80-110, parting but the extremities of this large range the two taxa
spreadingonly ca. 50? from the axis, the proximal, are distinct enough. Each has its "sister-species,"
infertile scales slightly recurved, the other scales P. lacmbertianaand P caacahuite, respectively, the
obtrullate,slightly concavo-convex, with seed cavi- former distinct, but the latter forming a species-
ties near the base on the adaxial side, the apex complex (Perez de la Rosa, 1993). The genetic affin-
recurvedat mid-cone, dark(reddish)[Link]- ity (isozyme analysis and cpDNA) of P flexilis ap-
sis relatively thick woody, triangular to rhombic, pears to be with the species in subsect. Strobi. not
with an obtuse apex, recurved but not reflexed at with (primarilyEurasian)species in subsect. Cem-
mid-cone, ochraceous to light brown, very resinous. brae, to which the Rocky Mountainspecies P albi-
Umiboterminal, transverse-triangular, obtuse, 5-10 caulis belongs (Krutovskii et al., 1994). This dine
mm wide, grey-brown,very resinous. Seeds broadly or complex ecologically encompasses the entire
obovoid, flattened, 10-15 X 8-10 mm, darkbrown, range from boreal-subalpine down to subtropical-
sometimes with blackish spots. Seed wings adnate, high montane,of which a good partof the morpho-
vestigial to very small, reduced to a narrow strip logical differences seem to be adaptiveexpressions.
aroundthe distal part of the seed up to 5 mm wide, As can be expected, taxonomies differ mainly con-
or absent when the seed is free from the scale. cerning the intermediateforms, which occur in part
Cooyledons 10-13, 30-40 mm long; according to sympatricallyand possibly hybridizeby introgression
Steinhoff & Andresen(1971) Pinus flexilis [Link]. has in some areas. No genetic barriers seem to exist
10 or fewer cotyledons. between any of these taxa, with the possible excep-
Distribution and ecology (Map 31). United tion of P albicaulis (Steinhoff, 1972).
States: In Arizona, New Mexico, and SW Texas (in Pinusfiexilis was describedby James (1823) from
the United States it is included in P strobiformis). Pikes Peak in Colorado,United States, but no herbar-
Mexico: Identified with certainty from only a few ium specimen was collected on the occasion, and a
collections made in Chihuahua, Coahuila, and S neotype from the most likely location was designated
Nuevo Le6n. Although scattered and restricted to by Andresen and Steinhoff (1971): Andresen et al.
high altitudes, other localities are likely to exist; 2125 (SIU-n.v.; isoneotypes,GH, K). Othermaterial,
specimens may have been collected as P strobi- studied at FHO, agrees with it in its short (4-5.5
forlnis. Piinusflexilis var. reflexa is, like P flexilis cm), entire, amphistomaticleaves, with a variable
var. flexilis, a high-montane to subalpine species width of 1-1.6 mm, a range also given by Kral
occurring on mountain summits or ridges above (1993). The cones are small, 7-15 cm long, with
contiguous forests. It grows in small groves or asso- widely spreading scales that bear short, broad,
ciated with P hartwegiiand P culminiicolaon Cerro straightapophyses.
Potosi, where it occurs generally above the pine Engelmann(1848) described P strobifornis from
forest belt in which P strobifornis occurs. The cli- specimens collected by A. Wislizenus in Chihuahua.
mate is cool temperate, with high winds and frost Mexico (see discussion under that species), "nearly
and snow [Link]:Pollen related to Pinus strobus and Piinusflexilis" but dif-
dispersal is in May-June, or even as late as July fering in its longer, more slender, serrulateleaves,

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210 Flora Neotropica

SON-
i ~~~~ ~ ~~~~~~~
~~~~~~~~~~~.Sl~ ~ ~ {,l
Map31insflxilisva.
;.Cr C ~~~~ .
rea(irces),Pzeowsii(
CY ?' '..
.. n. P . .
I ?

''..... ..

i s.

5... i
~~~~~~~~~~~~~~~~ '?
+~~~.
. . ? - ,.??... _~
'r""Y:~.
?-.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
?~~~~~~~~~~~~~~~. ...

'
? ~~ ~ ~ ~ ~ ?I

"'!? i'.........: ' ~ ' ... r

? ~ IU ... , ... .
n.?~~~ ~~
~~~~~~~~~~~~~~~~~~~,?
.?
Map 31. Pinux flexilis var. reflexa (circles); P rzedowskii (diamonds), P ma rlmarhmnezu (triangles).
:I ??
~.
???~~~~~~~ L...I..
?
,.I ~~~~~.,, ...,..

?.~ .* ~~~ i
,~~~~....
.._; . IL I.

~?* ...l?' ' 1, ..

[Link](ccle)Map31. Pin.
feii P zdwki (daod) [Link](rage)

"supra albo lineatis," and its larger cones (ca. 25 P. avacahuite, placing P. flexilis var. reflexa into
cm) with recurved apophyses. Thirty years later, synonymy with the former and P. strobiformis into
Engelmann(1879) described underP flexilis various synonymy with the [Link](1948) seems to
collections made by J. T. Rothrock in Arizona as have shifted his concept of P flexilis toward P.
varieties of that species. As Steinhoff and Andresen strobiformis in such a way that the former (sensu
(1971) pointed out, the differential characters on Martinez), for which he cites Cerro Potosi as the
which these varieties were based are more or less locality, is var. reflexa and P flexilis var. reflexa
those to be found in P strobiformis ("collectively (sensu Martinez)is P. strobiformis.
approximated his earlier description of P. strobi- Does P. flexilis [Link]. occur in Mexico? Andresen
formis"). But, as the type material shows, each of and Steinhoff (1971) were convinced that it does
them differed in only one character state from P not, despite citations to the contraryin Shaw (1909),
flexilis and not in the combined features by which Martinez(1948), and Loock (1950). The latter two,
Engelmann distinguished P [Link] con- in fact, have copied much more from their predeces-
clusion reached by Steinhoff and Andresen (1971) sors than is generally acknowledged. Our critical
on the basis of a morphometricstudy of collections observation of Shaw's reference (Nelson 6736, US)
from the United States (but not from Mexico)-that reveals that it is P flexilis var. reflexa and thus
Engelmann's varieties of P flexilis are synonymous eliminates the locality of General Cepeda (Coahuila)
with P strobiformis-stretches the morphological from the Mexican map. The populations on Cerro
plasticity of the latter to the limit. Nevertheless, Potosi and Potosi Peak cited by Martinez (1948)
through adoption of this view in the second edition belong in part to var. reflexa and, for a greater part,
of the Checklistof United States Trees(Little, 1979), to P strobiformis (see specimens cited under the
this treatmenthas now prevailed north of the Mexi- latter species). Perry (1991) published a photograph
can border(Kral, 1993). of a cone he collected near the summit of Cerro
Shaw (1914), who looked indeed across the bor- Potosi which indeed looks like typical P flexilis, but
der, had by then widened his species concept to such the foliar characters mentioned in his description
an extent that he recognized only Pinus flexilis and are closer to those found in var. reflexa and in P

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Systematic Treatment 211

[Link] maps in use in the United States spreadinghorizontallyor assurgent,of higher orders
(e.g., Burns & Honkala, 1990), as well as floristic long, slender,flexible, curveddown, ultimatelyassur-
accounts (e.g., Kral, 1993), limit "pure"P flexilis to gent. Crown conical or pyramidal in young trees,
a distance of several hundredkilometersnorthof the becoming more roundedor irregularin older trees.
Mexican [Link] more critical researchof the Shoots smooth, puberulousor glabrous,with promi-
populationson the high mountainsaround Galeana nent, small, short decurrent pulvini, pale reddish
and possibly elsewhere in NE Mexico has been brown. Cataphylls small, 5-10 mm long, subulate,
carriedout, it seems best to follow thatjudgement. brown with erose-hyaline margins, soon deciduous
Related to this discussion, a study of the original (falling well before leaf fascicles). Vegetativebuds
material of Pitnusflexilis var. serrulata Engelmann ovoid to ovoid-oblong, the terminalbud 8-15 x 5-
(1879) established the holotype to be from Mt. Gra- 8 mm, the laterals smaller, not or only slightly
ham, Arizona, Rothrock 783 (MO)-not Rothrock resinous;the scales free, spreading,subulate,reddish
654 (MO), as cited by Andresen and Steinhoff brown with erose-hyaline [Link] sheaths
(1971), which is one of two syntypes (the other 20-25 mm long, consisting of ca. 6-8 loosely imbri-
collection is Rothrock 1001, MO) of P flexilis var. cate, papery,orange-brownscales with hyaline mar-
reflexa. One wonders if their misconception as de- gins, soon disintegratingand deciduousat the end of
scribed above derived in part from this error? Of the growing season. Leaves in fascicles of 5, very
P flexilis var. macrocarpa Engelmann (1879) the rarely 6, in dense tufts toward the ends of ultimate
holotype is likewise from Arizona, Bischoff s.., branches, persisting 3-5 years, lax, straight or
Wheeler's Exped. (MO). These two varieties are slightly curved proximally, sometimes slightly
taxonomic synonyms of Pinus flexilis James var. twisted, spreadingat various angles from the shoot
flexilis. (usually 40-60?), (5-)7-11(-12) cm x (0.6-)0.8-
1.1(-1.2) mm, with remotely (rarely more densely)
37. Pinus strobiformis Engelmann, in Wislizenus, serrulateor nearly entire margins,acute, dark green
Mem. Tour N. Mexico 102. 1848. Fig. 56 to [Link]: Leaves commonly epi-
stomatic, sometimes weakly amphistomaticwith a
Pilus acacahuite C. G. Ehrenberg ex Schlechtendal few intermittentlines of stomataon the abaxial side,
subsp. strobiformis(Engelmann)E. Murray,Kalmia 13:
21. 1983. Type. Mexico. Chihuahua:Cosiquiriachi,Oct rarelyamphistomaticwith ca. 4 lines of stomataon
1846, Wislizenus231 (holotype, MO; isotype, MO). each face. Leaf anatomy: Cross section triangular,
PinusbonaparteaRoezlex G. Gordon,[Link].,ser. the abaxial side slightly convex; epidermis thick;
3. 1858:358. [Link] Mexico: hypodermis thin, unilayered; resin ducts (1-)2-5.
[Roezl s.n. ex herb.LindleY3/54 (lectotype,CGE, here external, rarely I medial, mostly situated on the
designated).
Piulls avacahuite C. G. Ehrenbergex Schlechtendalvar. abaxial side; stele terete; outer walls of endodermal
braclhypteraG. R. Shaw. [Pines Mexico] Publ. Arnold cells not thickened;vascularbundle I, oval in cross
Arbor. 1: 10. 1909. Type. Mexico. Durango:El Salto, section. Pollen cones crowded on the proximal part
12 Jul 1898, E. W Nelson 4555 (lectotype. A, here of new shoots, spreading,subtendedby light brown,
designated;isolectotypes, F, US). scarious bracts,ovoid-oblong to shortcylindrical,6-
Pinus strobiforrmisEngelmannvar. cnarvajaliiSilba, Phy-
tologia 68: 61. 1990. Type. Mexico. Jalisco: Sierra de 10 mm long, yellow, maturingto [Link]
Cuale, R. Lamas49 (holotype, MEXU). cones subterminal,axillary, solitary or in pairs or
[Link] Silba, Phy- sometimes whorls of 3-4 on stout, 15-25 mm long
tologia 68: 62. 1990. Type. Mexico. Nuevo Le6n: As-
cent of Cerro Potosi by N hogback, 27 km NE of peduncles, at first erect, becoming pendulous and
Galeana, 26 Jul 1934, C. H. Mueller & M. T Mueller falling with the peduncle attached. Immaturecones
1244 (holotype, NY; isotypes, A. BM. F, MICH). elliptic to short cylindrical, greenish purple, soon
Pinus avacahuite C. G. Ehrenbergex Schlechtendalvar. light brown, maturingin two seasons. Maturecones
n101ogaliciana Carvajal in McVaugh, Fl. Novo-Galici- variable, mostly cylindrical or broad cylindrical to
ana 17: 48. 1992. Type. Mexico. Jalisco: Sierra de
Cuale, 13 Aug 1984, Carvajal4703 (holotype, CREG). ovoid-oblong when opened, long cones usually
slightly curved, 12-30(-60) X 7-11 cm when open.
Tree, medium to tall, height to 25-30 m, dbh to Seed scales ca. 70-120, spreadingobliquely to pa-
80-100 cm. Trunkmonopodial, straight,erect. Bark tent, the basal scales recurvedor reflexed, the scales
thick, rough and scaly, fissured,breakinginto small, at mid-cone straight(the apophyses usually recurved
irregularplates, dark brown, weathering grey; on or reflexed), obtrullate, with curved margins, thick
young trees and branchesthin, smooth, silvery-grey. woody, of similar shape aroundthe cone but differ-
Branches of first order whorled, moderately long, entiating from base to apex, often longitudinally

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212 Flora Neotropica

i~ ~
??
~, .~.?.".
..,, ~~ ~ ~ ~ 1

~~~~~~~~~~a ~ . ~ V
~~~~~~~~~~~~~~~~~~~~~~~i~i?--.~:.

,,,i..' :-j~'I~.~ ? ....

b~~~~~~~~ .;: . :-
': ' :-::

B Ji
.,,
~~~:., -x,.,~'?
_- D ?.
~",~~~~~~..::,
,.~.
6~~~

F.56? int tbioisA-,Be34:DFGS'es aeaeaL1:DEFSta&Stls65..

[Link].x.5 , 3.)

FIG.56. Piniusstuvbiformis (.A-C, five 3141; Da. Fa, G, Stc/es, Fa~e/a el aii /2; Db, E Fb, Stead &Si~Ies 635). A.
Branch with leaf fascicles and immatureovuliferous cones. B. Leaf fascicle. C. Cross section of leaf. D. Ovuliferous
cones. E. Ovuliferous cone, basal view. F. Seed scales, one abaxial view. G. Seeds, one lateral view. (Magnifications:A.
B, D-G. >1).5; C, x'35.)

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SystematicTreatment 213

furrowedon both sides, with one or two deep seed Spellenberg et al. 10080 (FHO); Cosiquiriachi, 17 Sep
cavities adaxially,reddish brown to purplishbrown, 1846, Wislizenus155 (MO). COAHUILA: Near Saltillo, 10
Nov 1991, Frankis 117 (E. FHO); "12-14 leagues S of
with very small or virtually absent marks of seed
Saltillo," Mar 1880, Palmer s.n. (A, MO); "SierraMadre."
wings. Apophysismore or less cuneate or elongated, 1880, Palmer s.n. (A); Sierra La Cocordia. 26 km NW of
thick at the proximal end, thinningout and straight, [El] Fraile, top of mtn., 16 Jul 1941, Stanfordet al. 456
recurved or reflexed distally, smooth or longitudi- (A, NY); Sierrael Carmen,SentenelaCanyon on Hacienda
Piedra Blanca. 9 Jul 1936, Wvnd& Mueller 630 (A. E. K,
nally furrowed,often very resinous, light yellowish
MO, NY, S, US). DURANGO: Along rd. from Durango to
brown or dark ochraceous. Umbo terminal, broadly El Salto, rocky slopes above Arroyo de Mimbres.31 May
triangular,obtuse, 5-10 mm wide, greyish brown, 1951, Gentry & Gillv 10536 (A. MEXU. MICH); 15 km
usually covered with resin. Seeds (broadly)obovoid, W of El Palmito, along rd. from Mazatlanto Durango,30
slightly flattened,12-18 x 8-11 mm, reddishbrown Apr 1987, Hughes 975 (FHO, MEXU);Otinapa,n.d., Klaus
or brown; commonly one of the seeds on a scale s.n. (WU); La Ciudad, 30 Sep 1960, Little 17865 (US);
San Miguel de las Cruces, I Oct 1960, Little 17882 (K.
aborts. Seed wings adnate, vestigial to very small, US); ca. 5 km W of El Salto, 22 Mar 1963. Little 18944
up to '/2the lengthof the seed in some cones; aborted (K, US); "San Dinias."May 1939, M. Martinez2340 (NY);
seeds usually have relativelylargewings. Cotyledons near El Salto, 1900. M. Martine. 3559 (NY); near El Salto.
11-14, 30-40 mm long. S along lumber rd. toward Pueblo Nuevo, 26 Aug 1952.
Mavsilles 7796 (MEXU, MICH); Mesa de la Sandia, 14
Distribution and ecology (Map 32). United Oct 1905, Pringle 13615 (A, M. MICH, S, US); 10 km E
States: Arizona, New Mexico, rare in Trans-Pecos of La Ciudad at turn-off to San Juan, 27 Jun 1974, Rob-
Texas. Mexico: In Sonora, Chihuahua, Coahuila, erts & Keil 10291 (US); 155 km from Durango along rd.
Nuevo Le6n, Sinaloa, Durango, Jalisco, very local to Mazatlin. 5 Apr 1991, Styles, Favela et al. 12 (FHO).
in Zacatecasand San Luis Potosi. Pinus strobiformis JALISCO: Cuale, 1900, Carvajal 6096 (MICH); Talpa de
Allende, n.d., Santana 892 (MEXU). NUEVOLE6N:Cerro
is a montanepine of mesic sites; its altitudinalrange Potosi, 24 Jul 1935, Mueller 2283 (A, F, [Link], NY);
is 1900-3500 m. It occurs on sites with relatively Cerro Potosi, 1980. Klaus s.n. (WU); Cerro Potosi, along
deep, humus-richthoughoften rocky soils, especially rd. near Km 16, 30 Mar 1963, Little 18984 (K, US); Cerro
on N-facing slopes or along mountain streams. It del Viejo. 24 km W of Dulces Nombres, 16 Aug 1948,
Meyer & Rogers 3002 (A, BM. F, MO); Cerro Potosi, on
grows in small, pure stands within pine or pine- E flank of mtn., 25 Oct 1984. Rushforth415A (E); La
oak forest, but more commonly it is mixed with P Encantada,I Nov 1984. Rushfjrth 557 (E); Cerro Potosi.
arizolica, P engelmnanii, P. leiophylla var. chihua- 13 Mar 1980, Stead & Styles 630. 631. 634, 635 (FHO).
P. durangensis, P. lutholtzii, and/or various
hulctani, SAN Luis POTOSI:Mtn. SE of Canada Grande. 22 May
1950, Robinson & Newmantt106 (US). SINALOA: Sierra
species of Quercus and Arbutus. In a more mesic
Monterey. 10 Mar 1940, Gentry 5881 (MO); Ocurague,
forest type, it is associated with Abies and Pseu-
[Ocurahuil. Sep 1941. Gentry 6401 (MICH. MO. NY).
dotsuga, and at the highest altitudes with P hart- SONORA:'/2 km W of saddle between Cerro Pel6n and
wegii; on Cerro Potosi it grows at the lower limits Cerrode las Flores. 9 Oct 1992, Fishbeinet al. 779 (MO);
of the shrubby pine P cullminicola. The climate Sierra Huachinera.19 Jun 1953, Marshall 141 (UC).
in the Sierra Madre Occidental is characterizedby
summerrains (thunderstorms)as well as winter pre- Uses. As one of the "soft pines," P strobiformis.
cipitation; winter snows are common at the higher often equated with P ayacahuite in the vernacular
altitudes. Phenology: Dispersal of pollen occurs in of N Mexico (Perez de la Rosa, 1993), is sought
May-June. after by lumbermen, who consider its wood of good
quality for carpentry work.
Representative specimens examined. MEXICO. CHI- Engelmann (1848) described P. strobiformis based
HUAHUA: Mun. Guichochi,between Norogachiand La Cie-
on material gathered by F Wislizenus on the "highest
nega, 8 Nov 1972, Bye 3141 (A, MO); "region of the Rio
Bavispe," Dec 1890, Hartmans.n. (A): SW of San Juanito peaks about Cosihuiriachi" in Chihuahua. There are
at La Cien6ga,17 Jul 1957,
along rd. to Maguarichic, two sheets at MO of Wislizenus 231, of which MO-
Kohbloch 76 (BM, MEXU); ca. 55 km W of La Junta, 18 3377668 has been annotated by Engelmann; a third
Mar 1963,Little18910A(US);ca. 3 km W of El Vergel, sheet (Wislizenus 155, only a seedling) has not been
20 Mar 1963,Little18925(K, US); ca. 21 km W of El
determined definitely by Engelmann and is not type
Vergel,20 Mar 1963,Little18928(K, US); San Juanito,
14 Jul 1942.M. Martinez4000 (A, US); [Link],12 material, as stated in Farjon (1993). As some authors
Jul 1892,Mearnss.n. (US);SierraSanLuis, II Jul 1892, (Shaw, 1909; Andresen & Steinhof, 1971) have
Mearnss.n. (US);canyonof ArroyoAncho,7 Jun 1929,
pointed out, Engelmann apparently abandoned this
Mexia2609 (A, BM, E, F. MICH,MO,NY, P, UC);Mt.
name in his later publications, but as he did not
Molinora,I Sep 1898,Nelson4915(A, US);LargoArreco,
6 Jul 1969,Souli 453, 456 (MO);CascadaBasaseachic indicate of what species he would have considered
NationalPark,canyonalongRio Durazno.12 Nov 1989. it to be a synonym, later treatments in synonymy

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214 Flora Neotropica

I : i : r i~~~~~~~~~~~~~

r
4G
'??r.~~~~~~~~~~~~~~~~~
0
*??
5 *

32' I.. Piu

Map io%uii crls;Pamlsvr haess(rage)

with P. avacahuite (Parlatore,1868), or as a variety the range,Arizona and New Mexico. It is likely that,
of it (Shaw, 1909), are to be creditedto these authors in fact, P. strobiformnis sensu Sargent is synonymous
only. in part with P flexilis var. reflexa, as a preliminary
Carvajal and McVaugh (1992) discuss the type survey by A. Farjonof specimens in A and US has
specimen, Wislizenus231 (MO), seen by McVaugh, [Link](1898), in a precursorto Little's
and conclude that its differences with specimens of (1953; 1979) checklists, accepted P strobiformtis
P ayacahuite do "not seem to be enough to justify sensu Sargent,but extended its range to Mexico and
species status for P strobifornis." Unfortunately,of Guatemala,thereby implicitly including P ayacahu-
the cone there is nothing but a single seed scale, ite. Little (1953) at firsttreatedP strobiformisEngel-
mountedwith the abaxial side up, and no seed. Their mann and its synonyms under [Link] var. reflexa,
conclusion is based on the leaves of what amounts but after the work of Steinhoff and Andresen (1971)
to a very incompletecollection. More materialmight and Andresen and Steinhoff (1971), he reversed it
have warranteda different conclusion; however, in and accepted P strobiformisinstead, with P flexilis
this case the type collection does not suffice. var. reflexa in synonymy. As has been argued under
Sargent(1897), after some earlierdoubts, accepted the lattertaxon, the work of Andresen and Steinhoff
P. strobiformisbut based his concept of the species hardly dealt with Mexican trees, and they seem to
primarilyon materialfrom the most northernpartof have ignored Shaw's (1909) [Link] inclu-

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Systematic Treatment 215

sion of P. ayacahuite Ehrenbergex Schlechtendal the Santa Rita Mountainsof Arizona" is an invalid
(1838) in synonymy is certainly erroneous; they argument-a taxon need not be confined to its type
ought to have abandonedthe binomialP. strobiformis locality. Study of more collections from CerroPotosi
Engelmann(1848) for their species on the basis of reveals that there are possibly two relatedtaxa there:
nomenclaturalpriority. P flexilis var. reflexa and P strobiformis. Accounts
As interpretedhere, P. strobifornis, in accordance of cone variationmay be due in partto introgression
with its type, is the taxon Shaw (1909) accepted as involving both taxa. Pinus strohiformis cones are
P. avacahuite var. [Link] is more variable remarkablywide but short (12-17 x 8-11 cm when
than Shaw could have known given the few collec- open; Fig. 56Db) in the materialcollected by Stead
tions he studied, but its ovuliferous cone and seeds and Styles from this mountain in 1980. Study of
are nevertheless distinct from P. flexilis as well as genetic variationand taxonomic relationshipsof the
from P ayacahuite. As pointed out in a recent mor- "White pines" occurring on Cerro Potosi and
phometric study by Perez de la Rosa (1993), the neighbouring mountains [see also Perry's (1991)
longer, usually epistomatic, serrulateleaves and the claim of "typical"P flexilis found there] may turn
larger cones with usually elongated, recurved apo- out to be worthwhile. It seems premature,pending
physes and rudimentaryseed wings distinguish P such a study, to propose any furthernew names for
strobiformisfrom P. flexilis. Pinus ayacahuite has these pines.
similar foliage, though slightly longer still, but its Finally, Perry (1991) treated P strobiformisand
cones have thinner, flexible seed scales without P. ayacahuite var. brachypteraas two distinct but
markedseed cavities and with well-developed seed sympatrictaxa, the formerhaving shorterleaves and
wings. The reducedand ineffective wing of the seed shortercones than the latter. Since both names are
of P. strobiformissuggests a diversion from wind based on the same type, they are nomenclaturally
dispersal to animal vectors, which has been more identical, and Perry should have adopteda different
fully completed in P flexilis and is perfected in P name for the latter instead of following Martinez
alhicaulis, in which not only is wing development (1948), who in turn adopted Shaw's (1909) nomen-
completely suppressedbut the cones remain closed [Link] mentionsfindingoften treeswith interme-
throughripeningas well. diate morphologicalcharactersand ascribes this to
Carvajal(1986) describeda new species, P. novo- hybridizationbetween the two, which, in our opinion,
galiciana, from W Jalisco, citing P avacahuite var. amountsto the conclusion that individualsbelonging
brachypteraShaw, p.p. (excl. type), in synonymy. to one and the same species interbreed.
He failed to provide a Latin diagnosis. The same
taxon was later validly published as a variety of P 38. Pinus strobus Linnaeus var. chiapensis Marti-
avacahuite by Carvajal and McVaugh (1992). As nez, Anales Inst. Biol. Univ. Nac. Mexico I : 81.
was there indicated,Brian Styles would have treated 1940. Fig. 57
"this plant of western Jalisco under the name of Pinus cIhiapenlsis (Martinez) Andresen, Phytologia 10:
Pinus strobiformis Engelm.," which is maintained 417. 1964; Piotils strobus Linnaeus subsp. chiapensis
here. The remarkablelarge cones, given as up to 60 (Martinez) E. Murray, Kalmia 12: 23. 1982. Type.
cm long, have ratherthick woody scales with elon- Mexico. Chiapas:Ocotepec, Jul 1939, M. Martinezs.n.
gated and strongly recurvedapophyses and virtually (lectotype. MEXU No. 97, designated by Andresen.
1964).
wingless seeds. We have not seen cones of that
length, which may be exceptional. Tree, usually tall, height to 30-35(-40) m, dbh
Rushforth(1987) commented on pines of which to 100-150 cm. Trunkmonopodial, erect, usually
he hadcollected specimenson CerroPotosi in Nuevo straight, sometimes bifurcating. Bark rough and
Le6n under the name P reflexa. It was his opinion scaly, with shallow fissures, flaking, greyish brown
that this pine "may not be the same as the one to grey, on young trees and branchessmooth, greyish
Engelmann described as a variety of P flexilis, in green. Brantchesof first order whorled, long and
which case a new name may be required."Silba slender, spreadinghorizontallyor ascending, persis-
(1990) picked up the suggestion, virtuallycopied the tent; branchesof higherordersslender,short,spread-
description given by Rushforth, designated a type ing to subpendulous. Crown pyramidal in young
specimen from the same locality at NY, and named trees, mature and old trees usually open, shaggy.
it P strobiformisvar. potosiensis. That the name P Shoots slender, puberulentwhen 1-2 years old, gla-
reflexa "cannotbe applied here as the type is from brous thereafter,with small, short decurrentpulvini.

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216 Flora Neotropica

Cataphyllssmall, 4-5 x 1-2 mm, subulate,scarious, Guatemala: In the departmentsof El Quiche and
soon deciduous. Vegetativebuds ovoid-oblong, the [Link] small remnantpopula-
terminal bud 5-8 mm long, the laterals shorter,not tions in subtropical/warmtemperatemontaneforests
resinous; the scales imbricate, more or less charta- with abundant precipitation, which varies greatly
ceous, light brownwith [Link]- from site to site and shows no correlation with
cle sheaths 10-15 mm long, with thin, chartaceous, altitude; annual rainfalls of 1500-3000 mm have
imbricatebut soon free and deciduous, light brown been recorded. Rain is concentrated in the period
scales. Leaves in fascicles of 5, in dense or sparse June-September;frequentfog characterizesthe epi-
tufts at the ends of slender branchlets,persisting 2- phyte-richforests at higher [Link] altitudinal
3 years, lax, straight,slightly twisted, spreadingbut range of P strobus is substantial,being (500-)800-
not drooping, (5-)6-12(-13) (commonly 10) cm x 2000(-2200) m, but no frost is known to occur even
0.6-0.8(-1) mm, with (remotely) serrulatemargins, at its upper limit; the species is not frost-resistant.
acute, with minute teeth extending to apex, bicol- The soil is usually deep and loamy and well-drained.
orous, with non-stomatalface light green or yellow- Pinus strobus var. chiapensis occurs in mixed pine-
ish green, stomatal faces whitish green. Stomatlaon broad-leavedforest and in rain forest dominatedby
the two adaxial faces of the leaf, in (2-)3-6(-7) deciduous broad-leavedtrees. Otherassociated pines
conspicuous whitish lines. Leaf anatomy: Cross sec- are P maximinoi, P pseudostrobus, P ayacahuite,
tion triangular,with slightly concave abaxial side; and, more locally, P tecunumanii and P patula.
epidermis thick; hypodermis 1- or 2-layered; resin Dominant or common broad-leavedtrees are Quer-
ducts 2-3, external, large; stele terete;outer walls of cus spp., Liquidambar styraciflua, Magnolia, Plat-
endodermalcells not thickened; vascular bundle 1. anus mexicana, Ulmus mexicana, Carpinus carolini-
Pollen cones crowded, from the proximalend of new ana, Fagus mexicana, Clethra spp., and Brosimum;
shoots extending ca. halfway to the apex after full among conifers, Podocarpussp. is [Link]
shoot elongation, ovoid-oblong, 5-8 mm long, sub- is a definite affinity,especially in Veracruzand Chia-
tended by chartaceous,light brown bracts,yellowish pas, with the tree flora of the eastern United States
green to yellow, turning light brown. Seed cones which is also exemplified by this variety of Pinus
subterminal,usually in whorls of 2-4 on 15-45 mm strobus. Phenology: Pollen dispersal occurs in
long, slender peduncles, initially erect, becoming March.
pendulousin the second growing season before matu-
rity. Immaturecones cylindrical, with obtuse apex, [Link]-
Representative
ca. 15 x 7 mm, purplish blue, turning purplish PAS:Colonia Agr6nomos Mexicanos, E base of Cerro Tres
Picos, near Cerro Bola, 4 May 1972, Breedlove 25047
green to purplishbrownwhen growing. Maturecones (ENCB, MEXU, MO, NY); Selva Negra, 10 km from
cylindrical when closed, opening to ovoid, ovoid- Ray6n Mezcalapa along rd. to Jitotol, 25 Jan 1973.
oblong or cylindrical(dependenton length), variable Breedlove & Smith 32348 (ENCB, MO, NY); Santa Cruz
in size on a single tree, (6-)8-16(-25) x 4-8 cm de los Pinos, Aug 1984, Donahue 140 (NY); mtn. near
when open. Seed scales (40-)50-90(-100), attached Indianvillage of Chalchihuitan,20 Apr 1994, Farjon & D.
to a thin rachis, spreadingwide (usually not patent) Mejia 312 (E, FHO. HEH, K, MEXU, U); Bochil, 14 Mar
1993, Higman, Padilla & Styles 24 (EAP, FHO, HEH.
at maturity,the basal scales imbricate,not recurved, MO); near village of Pueblo Nuevo ca. 17 km N of Jitotol,
thin woody, similarly shaped around the cone, ob- 7 Mar 1989, Hughes 1288 (FHO, K, MEXU); Cintalapa,
long, widest at the apophysis,concavo-convex trans- Jun 1939, M. Marthiez803 (A, F, GH. INIF, MEXU, US);
versally, light brown, with lighter marks of seed Coapilez, Jun 1939, M. Martinez.2382 (A): Santa Maria,
20 km from Cintalapa, Jul 1939, M. Martinez 3419 (F,
wings [Link], irregularlyrhombic, US); 10 km from Jitotol, 20 Feb 1978, Stead & Styles 215
taperingor undulatingtoward an obtuse apex, con- (FHO). GUERRERO: 2.5 km SW of Yerbabuena,along rd.
cave, often furrowedlongitudinally,light dull brown, to Puerto del Gallo, 22 Aug 1984, Lorea 3138 (FHO);
often resinous. Umbo terminal, obtuse to flattened, along rd. between El Paraiso and Filo de Caballos, 16-18
km N of El Paraiso,4 Nov 1979, Kochet al. 79331 (ENCB,
3-4 mm wide, grey, resinous. Seeds obliquely ovoid
INIF, NY). OAXACA:45 km N of Puerto Escondido along
to ovoid, slightly flattened,7-9 x 4-5 mm, light or rd. to Sola de Vega, 12 Nov 1983, Hughes 387 (FHO);
[Link] wings adnate,obliquely ovate, with Teotlaxco, Sep 1986, Donahue 411 (NY); along rd. from
a straightand a curved side, 20-30 X 6-9 mm, light Gueva de Humboldtto San Juan Mazatlan, 6 Mar 1985.
brown, with darkerstripes. Cotyledons6-10, 20-40 McCarter& Styles 349 (FHO);4 km S of Lachao,along rd.
from Oaxacato PuertoEscondido, 13 May 1965, Rledowvski
mm long. 19599 (ENCB, GH, INIF, MEXU, MICH); San Juan Bau-
Distribution and ecology (Map 32). Mexico: In tista Cuicatlan,25 Sep 1973, Hill 1767 (A, NY): between
Guerrero,E Puebla, Veracruz,Oaxaca, and Chiapas. San Juan Zautla and San Juan Tentila, 1900, Schultes 765

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Systematic Treatment 217

/a
GiI
B
/F/,\

FIG. 57. Pinus strobus var. chiapensis (A-C, Stead & Stnles 175; D, Higntan, Padilla & StYles 24: E-G, Stead &
Styles 420). A. Branchwith leaf fascicles. B. (a, b) Leaf fascicles, (c) serrulateleaf margins.C. Cross section of leaf. D.
Shoot with pollen cones. E. Ovuliferous cone. F. Seed scale (two views). G. Seeds, with adnate wings. (Magnifications:
A, Ba, Bb, D-G, x0.5: Bc. x 10; C, x50.)

(A); above Rio Grande, 3 Apr 1946, Sharp & Xolocotzi PUEBLA:Atotocoyan, along rd. between Teteles and Maza-
46237 (NY); 75 km from Puerto Escondido along rd. tepec, 24 Mar 1976, Marquez 714 (F, NY), 24 Mar 1976,
toward Oaxaca, I Feb 1980, Stead & Styles 419. 420. Marquez 721 (F. MO, NY). VERACRUZ:Ca. 6 km (by air)

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218 Flora Neotropica

SSWof TlapacoyLin
towardAltotonga,nearbridge,II Jul Two characters scored higher: leaf serration in P
1982, Nee & Diggs 24885 (F, FHO, NY. SMU, WIS, XAL), strobus per unit leaf length was about one-half as
along Rio Tablazos,20 Mar 1983,Nee et al. 26078 (A, dense as in its variety from Mexico and Guatemala;
CHAPA, F, FHO, INIF, MEXU, MICH, MO).
ELQUICHE: and the proximal cone scales of the former were
GUATEMALA. Along rd. fromNebajto
Quiche, 13 May 1977, Mittak8473 (BANS, FHO,M); usually recurved,while those of the latter were usu-
belowcataractsof Rio Suchun,alongriverbelowNebaj,8 ally straight. These features are confirmed in the
Feb 1946, Sharp 4694 (EAP, F, GH, MEXU, NY); E of materialwe have seen.
Nebaj,II Feb 1978,Stead& Styles175 (FHO);RioAzul, The absence of any discontinuouscharactermakes
along rd. from Nebajto Chajul,II Feb 1978, Stead &
StYles181 (FHO). HUEHUETENANGO: Sierrade los Cuchu- a weak case for recognition of P strobus var. chia-
matines, CerroTolomarac,above Finca San Rafael. 24 Jul pensis as a distinct species. Some of the phenotypic
1942, Stevermark49489 (F); ca. 8 km from Basillas toward variation observed in the leaves between the two
Rio Ibal,I Jan 1946,Sharp461 (EAP,F, MEXU). taxa may be related to the great climatic differences
between their respectiveranges. Rzedowski and Vela
Uses. Although a locally valuabletimbertree, due (1966) reported some differences in the shape
to its scatteredoccurrencein often steep mountainous (length/width ratio) of seeds among populations in
terrainit is not specifically commercially exploited. Mexico, but no consistencies could be discovered in
Many large areas of montane rain forest have been these observationsto supporttaxonomic distinction,
seriously depleted and the species has become rare and they maintainedvarietal rank under P strobus
in large parts of its range. It has been classified by for all Latin American [Link] mentioned
IUCN as vulnerable(Farjonet al., 1993). a locality in Guerrero,the westernmost occurrence
In 1939 Martinez(1940) received some specimens of this taxon, where trees were found with large
of a pine previously unknown from Mexico and cones and short leaves suggesting closer relationship
described it as Pinus strobus var. chiapensis. He with P inonticola of western United States and Can-
recognized its close affinity with the well-known ada. Although the range of cone size they give is
species of NE North America and mentioned only large (9-26 x up to 10 cm when open), it is not
two different characters:his new variety had more much different from that observed in Stead & Styles
slender leaves which showed more often 3 (vs. 2) 18I from Guatemala(10-20 x 5-8 cm). Neither of
resin ducts. This treatment was maintained in his these measurementsare distinctly outside the ranges
monographon the Mexican pines (Martinez, 1945, of cone size reportedfor P monticola and P strobus
1948), although the illustrations of leaf sections (Kral, 1993). Similar range in cone size is given
given in his publicationsdo not supportany of the by Zamora and Velasco (1977) for P strobus var.
two differences [Link] Steyermark chiapensis in Chiapas. This taxon, more than 3000
(1958), Critchfieldand Little (1966) and Little and km distant from P strobus in the eastern United
Critchfield(1969) recognized its varietalstatus. An- States, is a typical vicariant which became isolated
dresen (1966) carriedout a pheneticstudy which led and adapted to a humid-warm temperate climate,
him to the conclusion that this taxon should be probably during the Pleistocene. Over time it may
elevated to species [Link],Gaussen (1960) had evolve to become a distinct species.
likewise classified it as a species, but (as usual)
without referenceto the basionym.
Andresen's phenetic approach, using statistical 39. Pinus rzedowskii MadrigalSanchez & Caballero
methods to evaluate multivariate(i.e., continuous) Deloya, Bol. T6cn. Secr. Agric. Ganad. Subsecr.
characterstates, led firstof all to the observationthat Forest. Fauna26: I. 1969. Type. Mexico. Michoa-
both the width of the leaves and the numberof resin can: "Puertodel Pinabete,"ca. 47 km W of Dos
ducts in them were insignificantlydifferent among Aguas, 25 Nov 1968, Madrigal S. 2202 (holotype.
the three taxa compared in his study (i.e., P mntti- INIF; isotypes, ENCB, GH, MEXU, MICH).
cola, P. strobus, and P strobus var. chiapensis). Fig. 58
However, he maintained that a number of other
characters showed "highly significant"differences. Tree, medium or rarely tall, height to 15-30 m,
Several of these, such as leaf length, numberof seed dbh to 30-60 cm. Trunk monopodial, erect, often
scales, and length of peduncle of the seed cone, not curved or contorted, with a partly clear bole only
only are of an entirely continuous quality but also when growing among other trees in a forest stand.
differ in their sums of means by only 25-30%. Each Bark up to 5-6 cm thick on the lower part of
can be more variable in materialfrom a single tree. larger trunks, rough and scaly, breaking into large,

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Systematic Treatment 219

exfoliating plates divided by deep, longitudinalfis- axis, thin woody, rigid, oblong, with parallel or
sures, dark brown, weatheringgrey; on young trees slightly undulatingmargins, concavo-convex, often
and branches rough and scaly, exfoliating in thin wider (up to ca. 20 mm) than the apophysis, light
plates, more or less smooth (but not as in species brownor orange-brownabaxially,idem with orange-
of sect. Strobus) in highest-orderbranches, reddish brown marks of seed wings on the adaxial side.
brown or grey-brown. Branches of first order long, Apophysis prominent, similarly shaped around the
whorled in young trees, spreading and ascending, cone, transverselykeeled, rhombicor pentagonalin
often contorted,persistent;branchesof higherorders outline, upper margin angular, on proximal scales
slender, flexible. Crown pyramidal in young trees, recurved,yellowish green turningochraceousor light
open and irregular in old trees. Shoots slender, brown,often secretingyellowish resin. Umbodorsal,
smooth, with weakly developed, transverselyridged rhombic-pyramidal,transversely keeled, 5-7 mm
pulvini, greyish. Cataphylls small, ca. 5 mm long, wide, obtuse or with a minute prickle, grey-brown.
fragile, subulate, with erose-hyaline margins, soon Seeds obovoid, (6-)8(-10) x (4-)5-6 mm, dark
deciduous. Vegetativebuds ovoid-oblong, the termi- brown. Seed wings articulate,effective, held to the
nal buds 8-10 X 4-5 mm, the laterals of nearly seed by two claw-like appendages,obliquely oblong,
equal size, not resinous; the scales imbricate,small, with a straightside, 20-30(-35) X 8-13 mm, brown,
subulate,with ciliate marginsand spreading,reddish partly translucent, with darker stripes. Cotyledons
brown apices. Fascicle sheaths of young leaves 7-9 (9-)10-12(-14).
mm long, with soon spreading,yellowish to reddish Remarks. The leaves do not elongate and remain
brown scales; scales of mature fascicles recoiling, as small buds covered by cataphylls in flushing
forming a rosette at the base of the fascicle, finally shoots until the shoots have reachedtheir full length
deciduous before the fascicles. Leaves in fascicles of of 5-7 cm. In the protologue(Madrigal& Caballero,
(3-)4-5, in dense, lax tufts persisting 2-3 years 1969) a leaf width of 0.7-1 mm is mentioned, but
on the branchlets, straight or slightly reflexed, not neither in the type collection nor in any others here
drooping, 6-10 cm X 0.6-0.8 mm, with irregularly studied does the observed width exceed 0.8 mm.
serrulatemargins, acute-acuminate,yellowish green Distribution and ecology (Map 31). Mexico: Mi-
or greyish green, weakly glaucous on the adaxial choacan: In three disjunct localities in the districtof
faces. Stomata on the two adaxial faces only, with Coalcomain,one near the village of Dos Aguas and
3-4 (intermittent)lines of stomata on each face (2 the others approximately40 km to the west. The two
grooves correspondingwith resin ducts on the abax- smaller populations, Cerro Chiqueritas and Cerro
ial face). Leaf anatomy: Cross section triangularor Ocotoso, areon steep talus of large,eroded limestone
transverse-triangular,with a convex abaxial side; blocks, near the summits of small mountainsin the
hypodermisunilayered,thin, rarely with some addi- mainly volcanic SierraMadredel Sur. Each has only
tional cells; resin ducts 2-4, external, 2(-3) on the a few score trees, from old to saplings. The larger
abaxial side and sometimes 1-2 additionallyon one populationis on more level ground,in an areaknown
or both adaxial sides; stele terete; endodermalcells locally as Puerto del Pinabete, also with limestone
largebut thin-walled;vascularbundle 1. Pollen cones bouldersbut interspacedwith other [Link] the
crowded, spirally arranged, after shoot elongation two formerareas, the trees remainsmall, <15 m tall;
forming a 5-7 cm long "spike" leaving the shoot in Puerto del Pinabete trees to 30 m have been
base and apex free, small, 5 x 3 mm, purplish, [Link] altitudinalrange is 2100-2400 m. Annual
turning brown. Microsporophylls peltate, 0.5 mm precipitationis ca. 1500 mm, most of it occurring
wide, with denticulate upper margin. Seed cones from June to [Link] climate is warm temper-
lateralon ultimate branches,solitary or in whorls of ate, with a minimum of -5?C (December) and a
2-4 on 15-30 mm long, slender, slightly curved, maximumof 30?C (April). Although surroundedby
remotely bracteate peduncles which fall with the extensive mixed pine forest with species like Pinius
cones. Immaturecones ovoid, 10-15 x 8-12 mm, pseudostrobus, P herrerae, and P oocairpa, these
with prominent,transverselykeeled umbos, purplish, species do not grow on the limestone talus. There,
turning green when growing, maturingin two sea- Quercus and shrubs such as Clusia salviiniiform an
sons. Mature cones ovoid to ovoid-conical when understoreywith Agave and tall herbs. Fires occur
closed, ovoid when opened, usually symmetrical, frequently, but rejuvenation seemed good at least
slightly flattenedat base, 10-15 X 6-8.5 cm when at Cerro Chiquerita,visited by A. Farjon in 1994.
open, very resinous. Seed scales ca. 80-120, parting Phenology: Pollen dispersal is in April-May, at the
to release the seeds, spreading at 40-50? from the end of the dry season.

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220 Flora Neotropica

II Y. (:

H .K1 ',I '

FIG. 58. Pinu.s rzedowskii(A-E, G-I, Madrigal S. 2202; F. Klaus 277). A. Branch with leaf fascicles and immature
ovuliferous cones. B. Leaf fascicles. C. Serrulateleaf margins. D. Recoiled fascicle sheath. E. Cross section of leaf. F.
Shoot with pollen cones. G. Ovuliferouscone. H. Seed scale (three views). I. Seed with articulatewing. (Magnifications:
A. B. F. G x0.5; C. x 15; D, x3; E. X50: H, I. x 1.)

Representative specimens examined. MEXICO. Mi- S. 3235 (INIF, MEXU, MS); Cerro Ocotoso, ca. 40 km W
CHOACAN: "Puerto del Pinabete," ca. 47 km W of Dos of Dos Aguas, 4 May 1980, Klaus 277 (FHO, WU), 22
Aguas, May 1980. Klaus 276 (FHO, WU), n.d., Madrigal Nov 1968, Madrigal S. 2177 (ENCB, INIF, K, MEXU,

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Systematic Treatment221

MICH,US);ca. 3 kmNWof DosAguas,on CerroChiquer- pine, in which the seeds are large and the wings
itas (fire lookout),7 May 1994,Farjon& Mejia353 (E, vestigial.
FHO,HEH,K,MEXU,U), 28 Mar1978,L [Link]& There is possibly an element of convergence in-
Martinez769 (IBUG,MEXU),1900,Klauss.n. (WU),27
Nov 1968,MadrigalS. 2218 (A, INIF,K, MICH,P, UC, volved in the evolution of the seed characters,and
US), 1900, Madrigal S. 3198 (INIF, MEXU, MS). in conjunctionwith it some cone characters,when it
is taken into account that these charactersare related
Uses. No logging of this rare pine seems to take to seed dispersal strategies, which in the case of
place, probablybecause of the inaccessibility of its seedless wings coevolved with animal vectors, espe-
populationsin remote locations and the poor form of cially corvid birds(e.g., Lanner,1982;Mattes, 1994).
most trees from a lumberman'sviewpoint. Mexican Genetically more-distant species could therefore
botanists and (regional) foresters are apparently have evolved similar cone/seed charactersand vice
aware of the uniqueness of this species, but it has versa. More phylogenetic work, using all available
not received a protectedstatus. Studies in population informative characters, is needed before one can
dynamicsrelatedto environmentalfactors,especially adopt newly proposedclassifications, such as a new
fire and reproduction,are much wanted to ensure subsectionRzedowskianaein (a polyphyletic)section
propermanagementof this evolutionarilyinteresting Parrlanae (Carvajal, 1986), which at this moment
pine. It was classified as endangered (IUCN-SSC, merely serves as a statement that it is somehow
Farjonet al., 1993). distinct.
This remarkablespecies was discovered in 1966
and botanical specimens were collected in 1968 in 40. Pinus maximartinezii Rzedowski, Ciencia
the NW partof the SierraMadredel Sur in Michoa- (Mexico) 23: 17, t. 2. 1964. Type. Mexico. Zaca-
can. Its rarity and the difficult access to the three tecas: Sierra de Morones, "Cerrode Pinones," W
small localities now known, far from any paved of Pueblo Viejo, 25 Jan 1964, Rzedowski 18258
roads, have been the main causes for its late dis- (holotype, MEXU; isotypes, ENCB, F, GH, INIF,
covery. K, MICH, P, US). Fig. 59
Taxonomically, Pinus rzedowskii is one of the
most interestingspecies in the genus. It is classified Tree, small to medium, height to 5-10(-15) m,
as a species in [Link] foliar morphology dbh to 40-50 cm. Trunk monopodial, short,
and anatomy places it close to P. cembroidesand P. branchinglow, often contortedor curved. Bark rela-
maximarrtinezii. Preliminaryresults from the analysis tively thick on the lower trunk, thin elsewhere,
of cpDNA of a limited number of Mexican pine smooth but eventually rough, tessellated into square,
species (Perez de la Rosa et al., 1995) place it and ca. 10 cm wide plates, inner bark orange-brownto
P. cembroides as unresolved clades away from P reddish brown, outer bark grey. Branches of first
maximartineziiand other species in subgen. Strobus. orderoften numerous,long, ascending to erect in the
The elongation of shoots prior to leaf development top, the lower branchesspreading;branchesof higher
is shared with P maximartiiezii; this phenome- orderslong and flexible, those bearingcones becom-
non is more weakly developed in other pinyon pines. ing pendulous. Shoots slender, slow-growing, gla-
The bark of P. rzedowskii is unlike any other in brous or with puberulent bases of fascicles, with
Strobus and at least externally resembles that of small, non-decurrentpulvini, initially glaucous or
species in subgen. Pinus. A more detailed compara- greyish green, turningorange-brownto grey. Leaves
tive study would be desirable here, using more pre- do not grow in flushing shoots until these have
cisely defined methods and terminologythan is cus- reachedtheirfull length,ca. 10 cm. Cataphyllssmall,
tomarily found in barkdescriptions(Junnika, 1994). ca. 5 mm long, narrowly triangular-caudate,re-
The seed cones and seeds are morphologicallymore curved, with erose-ciliate margins and an elevated
similarto some species in subgen. Pinus. The growth midvein, soon deciduous. Vegetative buds small,
of the apophysis, as deduced from the observation ovoid-conical, the terminalbuds 5-8 mm long, the
of cones at some different stages of development, laterals smaller, resinous; the scales imbricate, ap-
and its final shape are reminiscentof P merkusii,a pressed, subulate, light brown. Fascicle sheaths 7-8
very distinct SE Asian pine in subgen. Pinus. The mm long, light brown; the outer scales deciduous,
seeds are likewise small, with well-developed,articu- the innerscales recoiling, semi-persistent,forming a
late wings 3-4 times larger than the seeds. In this small rosette at the base of the fascicle. Leaves in
respect there is no resemblance to either P cem- fascicles of 5, very rarely 3 or 4, in dense tufts near
broides or P maximartintezii, nor to any other pinyon the ends of long, slender branches, persisting two

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222 Flora Neotropica

years, straight, lax, spreading at 30-70? from the and E slopes of the Sierra de Morones, from ca. 12
shoot, 7-11(-13) cm x 0.5-0.7 mm, with entire km SW to a few km W of [Link] main area
(rarely a few minute teeth) margins,acute, glaucous is locally known as Cerro de Piiones and lies a few
green, in some trees green, the two adaxial faces kilometresW of Pueblo Viejo, a small village at the
oten whitish. Stomata only on the adaxial faces, in E base of the mountains. Due to limited botanical
2-3(-4) lines on each face; the slightly convex abax- explorationof this partof Mexico and the likelihood
ial face has two faint grooves correspondingwith that similar ecological conditions occur on other
the underlying resin ducts. Leaf anatom!y:Cross "mesas" in the region, future finds of other popula-
section triangular,with a slightly convex abaxial tions cannot be ruled out. The rock of this mountain
side; hypodermisuniform, with 1-2 layers of cells; is in part sandstoneor limestone and also metamor-
resin ducts usually 2, external, on the abaxial side; phic, the soils are very rocky and shallow. Precipita-
stele terete; outer walls of endodermal cells not tion is probablyca. 700-800 mm annually,virtually
thickened; vascular bundle single, usually with a restrictedto four months in the [Link] maxi-
medianwall of reinforcingcells in the phloem. Pollen martinezii is virtuallythe only pine here, but a few
cones crowded on the proximal half of flushing scattered individuals of P leiophylla var. chihua-
shoots, forming an elongated "spike,"ovoid-oblong, hutanalhave been seen by us. Its altitudinalrange is
8-10 mm long, yellowish. Seed cones lateral,solitary 1800-240() m. Abundant are various large-leaved
on ultimate branches, on a very short peduncle or species of deciduous Quercus such as Q. uacrophy-
almost sessile (a lateralshoot usually grows with an la., which are bare during the long dry season from
angle from the near base of the cone). Immature Septemberto May. Fires occur regularlyin the region
cones large, ovoid-truncate,green until full-grown, in all vegetation types; it is not known whether this
maturingin 18-24 monthsor perhapslonger. Mature species is adapted well to reseed itself after fire.
cones broadly ovoid-truncate,widest slightly below Phenology: Pollen dispersal is usually in May-June;
the middle, (15-)17-25(-27) X 10-15 cm when the the ovuliferous cones take 18-24 months to reach
scales have parted. Seed scales ca. 80-110, parting maturityand perhaps longer for the seeds to ripen
slowly, the distal scales more than the proximal fully, which mostly remain in the cones. Squirrels
scales, usually insufficiently wide to release the are capable of biting off the apohyses to reach the
seeds, very thick woody, rigid, obtrullate, tapering seeds and probably store them. They, and probably
toward the very thick rachis, up to 50 mm wide birds, may play a crucial role in effective seed
below the apophysis, of similar shape around the dispersal, but this has not been investigatedto date.
cone but differentiatingfrom base to apex, with deep
seed cavities on the adaxial side, reddish brown. Representative specimens examined. MEXICO. ZA-
CATECAS:Mun. Juchipila, Sierra de Morones, 3-4 km W
Apophysis very prominent, 35-50 x 20-35 mm, of PuebloVicjo, 18 Feb 1975,Bailey75-78C(NY) Cerro
rhombic-pyramidalat mid-cone, usually straight, Alto,nearPuebloViejo,22 Apr 1976, Bailey76-22(UC),
transverselykeeled, the upper marginangular,those Los Fresnos,1983,Carvajal3650 (MICH),W of Pueblo
on the proximal and distal scales narrowlyconical, Viejo, 1982. Carvajal & Carrawza3350 (MICH), W of
often curved, dull light brown or reddish brown. Pueblo Viejo. I May 1994, Farjon & Mejia 338 (E, FHO,
often resinous. Umbo dorsal, obtuse-triangularor HEH.K, MEXU.U), W of PuebloViejo, 17 Apr 1981,
Godinas.m.(MEXU),W of PuebloViejo,2 May 1987,
rhombic-pyramidal,5-10 x 10-18 mm, sometimes Hughes983, 984 (FHO,K, MEXU),W of PuebloViejo,
with a minute prickle, concolorous or grey-brown. 1980. Klaus & Andrade s.n. (WU), 5 km N of Pueblo
Seeds oblong or ovoid-oblong, slightly flattened,20- Viejo. 1981, Perezdetla Rosa et al. s.n. (MEXU).
28 X (8-)10-12 X 7-10 mm, wingless when the
seed is disconnected from the cup-like depression in Uses. Like other"pinones"(pinyon pines) in Mex-
the scale, but on one side of the seed rudimentsof ico, this species is of local importancefor its edible
a wing may be present. Integumentca. 2 mm thick. seeds, which are harvestedby local people and mar-
very hard, yellowish brown or greyish brown. Seed- keted in the region. Due to its low stature and
lings large, with 18-24 cotyledons; juvenile leaves branchingof the trunk, its timber is not used. The
curved, flattened, ca. 8 cm long, silvery-blue, per- species is consideredendangered(Farjonet al., 1993)
sisting well beyond the developmentof adult leaves, due to fire and grazing hazards resulting in few
sometimes up to 20 years (Muir, 1992). seedlings that succeed in establishing themselves.
Distribution and ecology (Map 31). Mexico: In Although there is an awareness amongst botanists
S Zacatecas, where it is restrictedto a limited area and forestersin Mexico of its importance,currentin
of 5-10 km2 in scatteredstandson the summitridges situ protection (fire prevention and tire fighting)

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Systematic Treatment 223

F 111

II V A

ith

FIG. 59.
FIG. Pinusmarimartinezii
59. Pinus ,naximartinezii (A-D, Huighes983;
(A-D, Hughes E-H, Hughes
983; E-H, 984). A.
Hughes 984). Branchwith
A. Branch with leaf fascicles and
leaf fascicles bud. B. Leaf
and bud. Leaf
fascicle. C. Detail of fascicle sheaths. D. Cross section of leaf. E. Ovuliferous cone. F. Seed scale (adaxial view). G.
Seed.H.
Seed. H. Seedling
Seedlingwith
with20
20 cotyledons.
cotyledons.(Magnifications: B, E-Ga,
A, B,
(Magnifications:A, 0.5; C.
E-Ga, xx0.5; x 3 D,
C, X3; Gb, x 1.)
x40; Gb,
D, x40; 1.)

seems inadequateto ensure long-term preservation of the discovery of Pinus maximartineziiby Jerzy
of this interestingnarrowendemic (Farjon, 1994). Rzedowski in 1963, which was triggeredby an obser-
Carvajal and McVaugh (1992) gave an account vation of abnormallylarge pinyon seeds offered for

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224 Flora Neotropica

sale on a local market. Although it shares several with assurgentbranchesin young trees and a broad,
characterswith P cembroides (pinyon pine)-some irregular,very open crown in mature trees. Shoots
of which are synapomorphiesbut others, such as the elongating before the leaves startto grow, becoming
"wingless"seed, more likely the resultof convergent stiff, assurgent,greyish white, sometimes glaucous,
evolution-it is very distinct in others. Pimiismaxi- with remote,darker,non-decurrentpulvini persisting
martinezii has the highest numberof cotyledons of 2-3 years after the foliage has fallen. Cataphylls
all pines (and of conifers in general). There is the small, 5-8 mm long, subulate, brittle, with erose
peculiar median wall of sclerenchymatouscells in margins, dark brown, soon deciduous. Vegetative
the phloem of the single vascular bundle in the buds ovoid-acute to oval, the terminal bud ca. 10
leaves, noted by Rzedowski (1964), which he inter- mm long, the laterals smaller, resinous; the scales
preted as a trait showing intermediacy between imbricate, appressed, subulate, green, with hyaline
diploxyl (ancestral)and haploxyl (derived) leaves in or yellowish margins, turning to yellowish brown.
pines. Its exceptionally large and heavy seed cones Fascicle sheaths initially 7-9 mm long, consisting
(up to 2 kg when full-grown) are unlike those of P of 5-7 imbricate, red-brown scales, persistent, in
cembroidesand its allied taxa, but resemble those of mature fascicles 5-7 mm long, tight, dark brown,
P gerardiana of Afghanistan and Pakistan, which weathering grey. Leaves in fascicles of 3, rarely 4,
also has similar seeds. Zavarinand Snajberk(1987) connate, appearingas a single leaf, not partinguntil
reportsimilarityin monoterpenecomposition with P the final year of their persistence, in sparse tufts of
pinceana and dissimilarity with P. culminicola, one remote fascicles, persisting 2-3 years, stiff, straight
of the other "pinyons"studied. Pinus culminicola is or slightly curved, ascending or spreading, often
closely related to P cemnbroidesand its allied taxa twisted, 4-8(-10) cm x 0.7-0.8 mm, with serrulate
(Malusa, 1992; this study). A preliminaryinvestiga- marginsand acuminateapex, dark green, sometimes
tion of cpDNA differentiation in Mexican Pinus grey-green? [the latter colour mentioned by Shaw
species (Perez de la Rosa et al., 1995) appears to (1909) and (perhapscopied by) later authors,but not
confirmdissimilaritywith P cemrbroides, as does the observed in our material]. Stomata on all faces of
cladistic analysis of morphologicaldata by Malusa leaves, with (2-)3-4 intermittentlines on the convex
(1992), while it shows that there may indeed be abaxial face and 2-3 intermittentlines on each adax-
some relationship with P pinceana. These recent ial face. Leaf anatomy:Transverse-triangular in cross
investigationsshow thatsect. Parrylanae,as tradition- section, with a convex abaxial side; epidermis thick;
ally circumscribed(Little & Critchfield, 1969; Far- hypodermis unilayered or with a few irregularly
jon, 1984), is a heterogeneousand probablyartificial distributedadditional cells; resin ducts 1-2, large,
group. It also has become increasinglyapparentthat external, toward the margins on the abaxial side;
the "pinyon pines" (subsect. Cembroides) are not stele terete; large but thin-walled endodermalcells;
monophyletic(Malusa, 1992; Perez de la Rosa et al., vascularbundle single. Pollen cones concentratedon
1995; this study) and are in need of a comprehensive the proximal half of a new shoot, forming an elon-
revision. gated "spike," subtendedby scarious bracts, ovoid-
oblong to cylindrical, 7-9 X 3-3.5 mm, pinkish,
41. Pinus nelsonii G. R. Shaw, [Link]., ser. 3, turningbrown. Seed cones rare, often along the top
36: 122. 1904. Type. Mexico. Tamaulipas:Cerro shoot of young trees, near the proximal end of new
La Virgen, near Miquihuana, 10 Jun 1898, E. W. shoots, solitary or in pairs on very long (2.5-6.5 cm
Nelson 4501 (holotype, US; isotypes, A, K). long, 4-9 mm thick), recurved peduncles initially
with long, [Link] cylin-
Fig. 60
drical, 15 x 6 mm, with flattenedscales (the future
Tree, small or bushy, height to 5-10 m, dbh to umbos) subtended by triangular,glaucous bracts,
15-30 cm. Trunkmonopodial,usually straight,erect, purplish, maturingin 2, perhaps 2.5, years. Mature
some trees branching low near the ground. Bark cones irregularlycylindrical,usually falling from the
thin, smooth, only on the lower part of larger trees persistentpeduncle, sometimes curved, (5-)7-12 X
becoming scaly, with thin, small, easily exfoliating 4-5.5 cm when open. Seed scales ca. 60-100, parting
plates, ash-grey,with darker,brownishbandedareas, slightly or more widely but usually not enough to
finally brownish grey, on young trees and branches allow the seeds to fall, ratherweakly attached to a
smooth, greyish white. Branclheslong, slender,assur- thick rachis with a cuneate base (thereby moveable),
gent to irregularlydisposed, persistent, flexible, the thick, irregular,with 1-2 deep, cup-like depressions
foliage branchletsassurgent,forminga conical crown holding the seeds, reddish brown, with dark brown

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Systematic Treatment 225

Cb f

FIG. 60. Pinus nelsonii (A-C. Snles. lHowgh& Fivelt 4; D-G, Stl/es, Hoog/i & Favelia6). A. Branch with leaf
fascicles. B. Leaf fascicles, connate and with parted leaves. C. Cross sections of leaves: (a) three connate leaves, (b)
single leaf. D. Shoot with immatureovuliferous cone. E. Branchwith matureovuliferous cones. F. Seed scales. G. Seeds.
(Magnifications:A, B, D, E, XO.5; C, x35; F, G, x 1.)

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226 Flora Neotropica

seed cups. Apolphysisirregular,usually tetragonal NY); near Hwy. 101, ca. 25 km N of Tula. between Sierra
or pentagonal in outline, prominently raised and El Conche (W) and Sierra Mocha (E), on low limestone
constricted distally, transversely keeled, the distal hills. 29 Apr 1994, Farjon& Mejia 333. 334. 335 (E. FHO.
HEH, K, MEXU. U); 5 km W of [Link] of Jaumave.
part laterally flattened, often rugose, red-brown to 13 Apr 1985, Mars/hall85-120 (FHO); Miquihuana, 31
dark brown. Umtbodorsal, transverselykeeled, ob- Aug 1975. Robert & Passini 90()I (P); Mun. Palmillas.
tuse, up to 3-4 mm high, 6-7 mm wide, sometimes along rd. between San Luis Potosi and Ciudad del Maiz,
resinous, grey to blackish, with a small, triangular 22 Aug 1960. Matlrigal S. 325 (INIF); Sierra El Conche,
5 Apr 1961. King 4486 (MICH, NY, US); between "Her-
spine. Seeds (often one of two on a scale abortive), mosa" I= Valle Hermoso] and Miquihuana,23 Jul 1949.
obliquely obovoid, obtusely angular. 12-15 x 8-10 Stanfr!irei al. 2675 (U. US); 4 km from Miquihuana.25
mm, the integumenthard, 1 mm thick, pale to dark Oct 1984. Styles, Hoogh & Facvel 4 (FHO).
brown. Seed wings absent from seeds, rudiments
Uses. There is no commercial exploitation of this
may remain around the seed cavity on the scale.
Seedlings: Numberof cotyledons not observed;juve- species due to its low statureand its rarityin remote
nile leaves acicular-linear,2-4 cm long, green (not and inaccessible locations. Its seeds resemble those
of the true pinyon pines (Pinus subsect. Cembroides)
glaucous), recurringon seedlings and young plants
and are edible like these, but the seed crop is usually
up to 10-20 years old (in cultivation), stem chalky
white. low comparedwith these species.
Distribution and ecology (Map 33). Mexico: In Pinus nelsonii was described by Shaw (1904c-
S Nuevo Le6n and W Tamaulipasand scattered in 1905a), based on E. W. Nelson's collection in June
San Lufs Potosi. Critchfieldand Little (1966) report 1898 on a mountainabove Miquihuana,Tamaulipas,
"a collection locality" in S Coahuila, and Perry near the borderwith Nuevo Le6n. The very distinct
(1991) mentions the Montanas del Carmen as the connate leaves and the long pedunculate, curious
cones uniquelydistinguishedthis pine from all others
locality in that state, but we have not seen material
from thatstate. Pinus nelsonii is a rarepine occurring Shaw had known. Its connate leaves, held together
in the semi-arid foothills and on mesas of the Sierra by persistent fascicle sheaths, are morphologically
Madre Oriental;the most extensive populationsare and anatomically distinct from any of the other
found around the Sierra Pefa Nevada in Nuevo "pinyon pines." Recent molecular work, analysing
Le6n-Tamaulipas. It is restrictedto sites on rocky cpDNA of several Mexican pines (Perez de la Rosa
limestone with shallow soils. Its altitudinalranle is et al., 1995), revealed the absence of two length
1600-2300(-2450) m. Annual precipitationis 300- mutationsthat respectively defined species grouped
in the two subgenera recognized in Pinlus. This,
6()) mm, falling mostly in the summer during brief
[Link] conifers are Pienuscem- together with a unique cpDNA character, placed it
broides, P remota, and Juniperus spp. Pilnus nelsoanii on a clade separatefrom all other 12 species investi-
occurs in a scrubland zone with deciduous woody gated in this preliminarystudy of cpDNA in Mexican
taxa, e.g., Quercus,Mahonia, Comnarostaph ylis, Bra- pines.
hea. Sophora, and arborescent monocots such as
Yucca and Dasilyrion. At higher altitudes it may 42. Pinus pinceana G. Gordon& Glendinning,Pine-
grade into pinyon-juniper woodland, while lower turn 204. 1858. Type. Mexico. Morelos?: Location
down it is boundedby a hotterand driersemi-desert from protologue:"upona mountainalong the road
scrublandoften dominated by Cactaceae and Yucca to the city of Mexico, at a place called Guernavaca
spp. It, like several other narrowendemic conifers. is [Cuernavaca, . . . alt. 8000-9000 ft.," Ehrenberg
an
probably edaphic relict on limestone. Phenology: s.n. (holotype, not found; isotype, W). Fig. 61
Pollen dispersal occurs in March-April. Pintis lati./isqumaEngelmann,p.p.. [Link]..n.s. 18:
712. 1882. Type. Mexico. Coahuila: "MountainsS of
Representative specimens examined. MEXICO. Saltillo. in the Sierra Madre; 12 or 14 leagues (sheet
NUEVOLEON:Sierra Pefia Nevada. Jan 1942. M. Martine:. I). [ = 40 mi. 165 km] (sheet 2) S of Saltillo," Mar
2/84 (NY); Sierra de San Lorenzo, E of Doctor Arroyo. 7 188). Palmer /299 (holotype, MO. foliage and seed
Nov 1904. Pringle 10016 (BM, E. ENCB. Fl. GH. GOET. scale specimen mounted on 2 sheets. excl. branch on
K. M, MO, NY); 5 km N of San Antonio PeniaNevada. 25 right-handside of sheet I; isotypes. A, K, NY, US).
Oct 1984. Styles. Hoogh & Fati,el 6 (FHO). SAN Luis
PoTosi: Mun. Villa Arista, Sierra de los Li6brillos. 3 Feb Tree, small and bushy, or a large shrub, height to
1983, ElizMhodsx.n.(INIF); Realejo, Sierra La Trinidad.
1980, Klaus s.n. (WU). TAMAUIIPAS: N of Miquihuana.20 6-10(-12) m, dbh to 20-30 cm. Trunk short, often
Nov 1972, Becerril s.n. (INIF); Sierra El Conche, along branchingfrom near the ground, erect, monopodial,
rd. to Bustamante,28 May 1986, Diggs et til. 3759 (MO. contortedor forked. Bark thin, smooth, only on the

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32
Systematic Treatment

1,~~~~~~~~~~~~~~~~~~~~~~~~~~~~,

? ~ ~ ~ ~ ~ ?

... . . ,....'

[Link] eso i

lower partof the trunkand lowest branchesexfoliat- curling and deciduous, only the basal part may re-
ing into irregular plates, eventually longitudinally main for some time in some fascicles. Leaves in
fissured on old trees, with occasional horizontal fascicles of 3, rarely 4, in sparse tufts toward the
cracks, on young trees and branches thin, smooth, ends of drooping or pendulous branchlets, remote,
brownish grey, weathering grey. Branches of first spreading at 30-60? from the shoot, persisting 2-3
order long, thick, contorted,ascending or spreading, years, straight, rigid, 5-12(-14) cm (often variable
of higher orders long, slender, flexible, the ultimate on a single shoot) X 0.8-1.2 mm, with entire mar-
branches drooping to pendulous, forming a broad, gins, acute, greyish green. Stomata: Most leaves
irregular,open crown with branches of young trees epistomatic, with 2-4(-5) lines on each adaxial face,
often drooping to the ground. Shoots long, slender, some leaves amphistomaticwith in addition 2 faint,
flexible, smooth, with small, non-decurrent,widely intermittentlines of stomataon the abaxial face. Leaf
spaced pulvini, brownish grey to grey. Cataphylls antatomy:Triangularto transverse-triangular in cross
small, 3-4 mm long, ligulate to narrowlytriangular, section, with a convex abaxial side; epidermis thick;
with erose-hyalinemargins,light brown, soon decid- hypodermis with 1-2 layers of cells: resin ducts 2,
uous. Vegetativebuds small, inconspicuous, ovoid- external, toward the margins on the abaxial side:
globose to ovoid-conical, the terminal bud 4-6 mm stele terete; cells of endodermis large, with thin
long, the laterals smaller, not resinous; the scales walls; vascularbundle single. Pollet cones crowded
imbricate, appressed, narrowly triangular, light or more remote on the proximalpartof a new shoot.
brown. Fascicle sheaths ca. 10 mm long; the scales subtended by broad, chartaceous, yellowish bracts,
initially imbricate,ligulate-linear,soon spreadingor ovoid-oblong, 8-10 X 4-5 mm, purplishor yellow-

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228 Flora Neotropica

5'
2

FIG. 61. Pinus pinceana (A-E, G, Styles, Burle', Favelci at. 2; F. H. Farjon & D. Mejia 337). A. Branchlet with
leaf fascicles and immatureovuliterous cone. B. Branchletwith leaf fascicles and a few pollen cones. C. Leaf fascicle.
D. Cross section of leaf. E, F. Ovuliferous cones. G. Seed scale (three views). H. Seeds. (Magnifications:A-C. E-H.
x0.5; D, x40.)

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SystematicTreatment 229

ish. Microsporophyllssubpeltate,broadeningfrom a Engard 23110 (ENCB, MEXU, MICH); Puerto Colorado,

cuneate base to a broadly obovate, cuspidate distal (Cafion La Fragua), 2 Sep 1941, Johnston 8735 (GH); W
of Carneros Pass near Cuauhtemoc on dry slopes above
part. Seed cones lateral, not on ultimate branches, cactus desert, 1980, Klaus s.n. (WU); Sierra Parras,9 km
solitary or rarely in pairs on slender, curved and SW of Parras,6 May 1983, Marshall 83-27 (FHO); near
easily breaking peduncles, spreading, maturecones Parras,Mar 1942, M. Marttnez 2375 (NY); near railway
pendent through the habit of the [Link]- sta. El Fraile, Dec 1941, M. Martinez4101 (A); El Cinco,
ture cones ovoid-oblong, obtuse, purplish, maturing 21 Aug 1975, Robert & Passini 5780 (INIF. P); near sta.
El Fraile on Coahuila-Zacatecasrailroad, 1950, Mirovs.n.
in two years. Mature cones ovoid-oblong to short
(CAS, MO, UC); near GeneralCepeda, 1900. Nelson 6140
cylindrical, often irregular,5-10 x 3.5-6(-7) cm (A, NY, US); Careros Pass, 8 Sep 1889, Pringle 2293 (A,
when open. Seed scales ca. 60-80, parting only K, MO, NY, P, US), 12 Nov 1904, Pringle 13207a (A. K,
slightly or sometimes more widely, weakly attached MICH, MO, US), 1903, Shav s.n. (A, K, UC, US); 9 km
to the slender cone rachis with a small cuneate base, S of Parrasin Sierras Negras, 3 Jul 1941, Stanfordet al.
148 (A, DS, MEXU, MO, NY); Cafionde los Palos, 26 Apr
therefore easily moveable, thick woody, concavo- 1985, Styles, Burley & Favela 2, 3 (FHO, K). HIDALGO:
convex, with (1-)2 deep, cup-like depressions hold- Barrancade Tolantongo, 3 km SW of Tolantongo,ca. 40
ing the seeds, puberulent in fresh cones, reddish km NE of Ixmiquilapan,29 Oct 1992, Gardner & Knlees
brown abaxially, dark brown adaxially. Apophysis 5122 (E, U); "near the hacienda del protrero Rosin de
Mextitlan, (along) the road from Mexico (City) to Tam-
irregular, prominently raised, transversely keeled, pico," 1842-1843, Ghiesbreght 54 (P, dupl. s.n. in K);
rhombic to pentagonal in outline, the upper margin Barrancade Tolantongo,27 Nov 1993, Hughes 1852 (FHO,
angularor undulating,lustrousred-brown,sometimes K, MEXU); Sierrade la Mesa, 1900, Rose et al. 9092 (A,
with concentric colour rings or radially striated. NY, P, US). QUERETARO: Vizarr6n,23 Jul 1960, Dia: L.
Umbo dorsal, flattenedat apex, transverselyrhombic s.n. (ENCB); Pil6n, ca. 80 km NE of Quer6taro,along rd.
to Pinal de Amoles, 24 Apr 1949, McValugh10347 (BM,
or oval in outline, up to 10 mm wide, 5 mm high, DS, ENCB, GH, MEXU, MICH, MO). SANLuis POTOsi:
grey, with a minute prickle. Seeds (1-)2 per scale (I Near rd. from Hwy. 101 to Guadalcazar,near village of
often abortive), obovoid, obtuse-angular,11-14 X Aguaje de los Garcia, El Canon Hondo, 29 Apr 1994,
7-8 mm, wingless when detached from the scale, Farjon & Mejia 337 (E, FHO, HEH, K, MEXU, U); 4 km
E of Charco Blanco, on slopes S of rd. to Guadalcazar,
brown, with a darkerside where the seed was adnate 1980, Klaus s.n. (WU); 72 km N of San Luis Potosi, along
to the scale depression. Integumentca. 1 mm thick, rd. to Matehuala, 21 Sep 1979, Robert & Passini 6837
[Link] wings absent from detached seeds, some- (ENCB). ZACATECAS: Pico de Teira, Sierra Zuluago, SW
times rudimentaltissue on the scale aroundthe seed of Cedros, Feb 1908, Lloyd 30 (MO, UC, US); along rd.
to Santa Rosa SW of Concepci6n del Oro, n.d. Passini
[Link] (numberof cotyledons not ob- 20109 (INIF).
served) with chalk-white stems and narrowlylinear,
glaucous leaves appearingfor 5-8 years. Uses. This species is not commercially used; the
Distribution and ecology (Map 34). Mexico: seeds are edible ("pifones") but trees are scarce,
Scatteredin Coahuila,N Zacatecas,San Luis Potosi, inaccessible, and far apart and usually with a low
Quer6taro,and Hidalgo. The locality (along the road crop of cones compared to P cembroides and its
to Mexico City at Cuernavaca) stated in the pro- close relatives.
tologue is probably an error. It occurs often on When Gordon and Glendinning (Gordon, 1858)
calcareousslopes and in ravines(barrancas)at 1400- described Pinus pinceana, it was said to have been
2300 m in arid and semi-arid mountains of NE discovered by "M. Gheisbreght" IA. B. Ghiesbreght]
Mexico. Annual precipitation is lower than in the in 1844 near Mextitlan in Hidalgo (20?36'N,
pinyon-juniperbelt (ca. 300-400 mm, according to 98?45'W) and collected under his "No. 34." In P
Perry,1991) and is concentratedin the [Link] there is a collection Ghiesbreght 54 from 1842-
species sometimes occurs with Pinus cembroides, 1843 made at that locality; an unnumbered specimen
more often with Juniperusfiaccida,and usually with collected by the same author is in K, which is
numeroussclerophyllousor deciduous shrubs-e.g., probably a duplicate. The description is instead based
Prosopis (in arroyos), Mimosa, Karwinskia, Leu- on a collection made by C. Ehrenberg and said to be
caena, Sophora, Quercus, Cercocarpus, Gochnatia, from "upon a mountain along the road to the city of
Fouquieria-as well as Yucca,Agave, [Link] Mexico, at a place called Guernavaca, at an elevation
trees are often widely scattered in this vegetation. of from 8000 to 9000 feet." The city of Cuernavaca
Phenology: Pollen dispersal has been observed in is in Morelos, but as later authors (Shaw, 1905a;
April-May. Martinez, 1948) have pointed out, the species does
[Link]- not occur in that state and it was concluded by Mirov
HUILA:
SierraSan Marcos y Pinos, 10 Oct 1972, Gentry & (1952) to be most likely an error on either Gordon's

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230 Flora Neotropica

f~~~
.L .. ~~ . .

I
.~~~~~~~~~Y

SON

Map-
34.?... Pmnl1s pinccan

or Ehrenberg'spart. The type specimen, according a later collection (Palmer 1299, MO), but as Shaw
to Shaw (1905a), is a single cone at K. No material (1905a) had already observed, this material from
collected by Ehrenberg,or obtained from Gordon's near Saltillo in Coahuila became mixed on the same
herbarium,has been found there, but a fragmented sheet with a branchof what A. Farjon identified as
cone "from [Link]"is at W, sent to Endlicher P flexilis James, probably from the United States.
by Lindley (through the HorticulturalSociety of For this reason, Engelmann, who was not aware of
London);Gordon may well have seen this cone. As this mixture,described his species with short (3.5-5
there may still be some original material in other cm), extremely slender, slightly serrulate leaves in
Europeanherbaria,this specimen is here recognized fascicles of 5 and with a cone identical to Gordon's
as an isotype. It is perhaps due to some lack of species. On the type sheet I of 2 is a very small
memory by Ehrenbergthat Gordonand Glendinning branchletwith ca. 5 cm long leaves in fascicles of
describedthis species as a tree growing 60 ft (20 m) 3, but apparentlythis escaped Engelmann'sattention.
high, but if they really had seen sufficient specimens All that is now left of the cone of Palmer 1299 is a
of the foliage it is ratherstrangethat they described seed scale (sheet 2 of 2), but, in conjunctionwith an
the leaves to be "in threes, but frequentlyin two's." illustrationof a cone on the same sheet, it suffices
Old fascicles may loose a leaf sometimes before they to identify Engelmann'sspecies unambiguously.
fall entirely, but on normal fascicles they would The morphological,chemical, and molecularchar-
be in threes, rarely fours. Clearly, Gordon and his actersof Pinus pinceana place it closer to P maxima-
associate were not at all familiar with this species rtinezii (Malusa, 1992; Zavarin & Snajberk, 1987;
but, following the craze of the day (Roezl, 1858), Perez de la Rosa et al., 1995) than to P cembroides
were eager to describe "novelties for the garden"at and its closely relatedtaxa (e.g., P edulis, P remota,
the slightest [Link] is a remarkablestroke of P quadrifolia). There is ratherconvincing morpho-
luck that they hit upon a unique species. logical evidence that these two species are also more
Engelmann(1882) based his Pinus latisquamaon closely related to P bungeana and P gerardiana of

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Systematic Treatment 231

E Asia than to the P cembroides "complex"of SW lines on each adaxial face. Leaf anatomy: Cross
NorthAmerica(Farjon,1984; Bailey & Hawksworth, section transverse-triangular or half-terete(2-leaved
1988; Malusa, 1992). This could mean, that they are fascicles); hypodermisuniform, 1-, more commonly
to be seen as palaeo-relicts,as opposed to the pinyon 2-layered; resin ducts 2, rarely only 1, external on
pines sensu stricto, which appearto have undergone the abaxial side; stele terete; outer cell walls of
allopatricor parapatricspeciation in relation to Ter- endodermis not thickened; vascular bundle single.
tiary climatic change in the region (Malusa, 1992). Pollen cones crowded on the proximal half of new
shoots, in more or less elongated, spiculate clusters;
43. Pinus cembroides Zuccarini,Abh. Math.-Phys. subtendedby cataphyllousbracts with hyaline mar-
Cl. Konigl. [Link]. Wiss. 1: 392. 1832. gins; small, ca. 5 X 3 mm, the larger ones proxi-
mally; microsporophyllssubpeltate,with erose mar-
Slhrubor tree, height to 3-15(-25) m, dbh to 10- gins. Seed cones subterminal from lateral buds,
80(-120) cm. Trunkmonopodial,very short(shrubs), solitary, paired or more rarely in whorls of 3. on
branchingimmediatelyabove the ground,or short to very short,3-5(-8) mm long peduncleswhich remain
medium size, contorted or more or less straight, with the fallen cones. Immaturecones subglobose to
branchinglow. Bark thick, rough and scaly, breaking ovoid, 8-10 mm long, purplish,or yellowish green,
into small, irregular,shaggy plates, sometimes di- soon light brown to reddishbrown, maturingin two
vided by longitudinalbut shallow and irregularfis- seasons. Maturecones seemingly sessile, irregularly
sures, not easily exfoliating, grey, exposing yellowish globose or ovoid-globose when closed, spreading
to orange-yellow inner bark; on young trees and often wider than long when opened, with a flattened
branches thin, smooth but soon flaking, grey or base, irregular in size and shape, often resinous,
greyish brown. Branches of first order long, usually (2-) 3-5(-7.5) x 3-6(-7) cm when open. Seed
heavy, spreading or trailing on the ground, or as- scales 25-40(-50), parting easily and widely, nar-
cending, irregularlydisposed in most trees;of second rowly and more or less weakly attachedto the rachis
and higher orders long, contorted, spreading,assur- and hence moveable, spreadingor reflexed,only the
gent or ascending. Crown an open or dense shrubor central 10-15(-20) fertile, irregular,up to 15-20 mm
broad, open, irregular. Shoots orange-brown or wide, concavo-convex, with 1-2 deep seed cavities
slightly glaucous at first,soon grey, roughwith small, bordered by membranous seed wing remnants (I
non-decurrentor short decurrentpulvini. Cataphylls seed commonly aborts),ochraceous'to(light) brown
small, 2-4 mm long, subulate or triangular,acute- abaxially, usually darkerbrown with reddish brown
acuminate, with erose margins, light brown, cadu- seed cavities on the adaxialside, weatheringblackish
cous, leaving a small ridge above the pulvini. Vege- brown or greyish black. Apoplhsis raised, trans-
tative buds ovoid-oblong to oval-cylindrical, the versely keeled or radiallykeeled or ribbed, rhombic
terminalbuds 5-8 X 3-5 mm (sometimes up to 10 to pentagonalin outline but irregular,upper margin
mm long), the laterals smaller, not or slightly resin- entire or undulating,often angularor curved, in all
ous, light brown or ochraceous;the scales subulate, shades fromyellowish green or ochraceousto reddish
with spreadingor recurved,darkerbrown apex. Fas- brown, sometimes lustrous. Umlbo dorsal, flat or
cicle sheaths short, 4-6 mm long, loosely imbricate, raisedand curved, 2-4(-5) mm wide, greyish brown
the scales (pale) brown, soon recoiling, then pale to blackishbrown,with a minuteprickle,often resin-
straw-colouredto grey, forming a small rosette at ous. Seeds obovoid-oblique,often acutish,sometimes
the base of the fascicle but deciduous before the faintly ridged, 10-16 X 6-10 mm, greyish brown to
leaves fall. Leaves in fascicles of (2-)3(-4, rarely blackishgrey, or light brown;integumentthick, 0.5-
5), spreadingor assurgent,in small, often regularly I(- . I) mm; macrogametophyte("endosperm"of au-
spaced tufts persisting (3-)4-5(-7) years, curved or thors)pinkishor white when [Link] wings absent
less often straight,lax or sometimes rigid (especially when the seed is detached from the scale, the rudi-
short leaves), (2-)3-6(-8) cm X (0.6-)0.7-1(-1.2) mentsof it remainingwith the scale. Cotyledons(5-)
mm, margins entire, apex acute-acuminateor pun- 6-12(-17), grey-green.
gent; colour variable, dull green to glaucous-green, Distribution (Maps 30, 35, 36). United States:SE
with or without white adaxial faces; sometimes pro- Arizona, SW New Mexico, SW Texas; Mexico: A
ducing a few resin [Link]:Leaves amphisto- wide range running from the interiorslopes of the
matic, or epistomatic in one variety,in 2-3(-4) lines SierraMadresinto the interiorof south-centralMex-
on the convex abaxial face (or none), in 2-3(-4) ico, where it reaches its southernmost point in

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232 Flora Neotropica

[Link] outlier occurs in the Sierrade la Laguna two "pinyonbelts"mergenorthof the "EjeVolcanico
at the southerntip of Baja California. Transversal"and continue SE into Hidalgo. In much
of its range, Pinus cembroidesoccupies a transition
zone between (semi-)desertplateausand valleys and
Key to the varieties and subspecies more mesic coniferous montaneforests. The altitudi-
of Pinus cembroides nal range is extensive (800-)1500-2600(-2800) m,
I. Leaves epistomatic, with two white stomatiferous with highest occurrences in the SE. It grows on a
bands on the adaxial faces, the abaxial face green. variety of substrates,ranging from alluvial bajadas
...............................43a2. subsp. cembroidesvar. bicolor to volcanic
rock, usually on scarcely developed soils.
I. Leaves amphistomatic, more or less monochro-
matous. It forms open woodland alone or mixed with Juni-
2. Leaves commonly relatively long, (2.5-)4-7 perus, Pinus nelsonii, P pinceana, Quercus, Yucca,
(-8) cm, mostly straight,<1 mm wide.............. Agave, Cactaceae (e.g., Opuntia), Arctostaphylos,
.....................................................43b. subsp. laguncle Ceanothus, Arbutus, and other shrubs of dry, hot
2. Leaves commonly shorter, (2-)3-5(-6.5) cm,
areas; at higher and/or moister sites, it forms partof
mostly curved, (0.6-)0.7-1.2 mm wide.
3. Leaves in fascicles of 2-3............................. a mixed pine-oak woodlandor forest including,e.g.,
..............43al. subsp. cembroidesvar. cembroides Pinus arizonica, P engelmanni, P leiophylla var.
3. Leaves in fascicles of 3(-4, rarely 2 or 5)... chihuahuana,and P pseudostrobusin the SE partof
...........................................43c. subsp. orizabensis its
range. The climate is warm and dry, with annual
precipitationin the range of 380-650 mm and a dry
43al. Pinus cembroides Zuccarini subsp. cem- season of 7-8 months. Frost may occur at higher
broides var. cembroides. Type. Mexico. Mexico: elevations in the interiorbut is [Link] is
Near Sultepec, Santa Cruz, Sep 1827, Karnvinski an importantmutualist relationshipwith the corvid
s.n. (holotype, M; isotype, K). Fig. 62 birds Aphelocoma coerulescens and Gymnorhinus
Pinins llaveana Schiede ex Schlechtendal, Linnaea 12: cyanocephalus (Tomback
& Linhart, 1990), which
488. [Link]:BetweenZimapan feed on the seeds and cache them, therebyproviding
and Real del Oro, 1830?, Schiede s.n. (holotype, HAL). an effective [Link]: Pollen
Pinus osteosperma Engelmannin Wislizenus, Mem. Tour is dispersed in May-July, depending mainly on alti-
N. Mexico 89. Apr 1848. Type. Mexico. Coahuila: tude.
Saltillo,"onthebattlefield of BuenaVista"(see Engel-
mann, 1848), 19 Mar 1847, Gregg 321 (holotype, MO).
[Link].
Small tree, height to 10-15 m, dbh to 60-80 cm. HUAHUA:
Along rd. between Babicoraand Yepomera,8 Apr
1977,Bennettet al. 843 (MO,NY); Norogachic,23 Aug
Trunkmonopodial, short to medium size, contorted
1978, Bye 8822 (E, FHO);ArroyoAncho, S of Mesa
or more or less straight, branching low. Leaves in Colorado,28 Aug 1978,Bye 8942 (E); SW of Matachic
fascicles of 2-3, spreading or assurgent, persisting nearkm 9.7 alongrd. Matachic-Cocomoarachic, 29 Aug
3-4 years, curvedor nearlystraight,lax or sometimes 1978,Bye9011 (E);NWof Panalachic, 31 Aug 1978,Bye
cm X 9077 (E); SE of [Link] Hojasichic,4 Jul
rigid (especially short leaves), (2-)3-5(-6.5) 1984,Byeet al. 12872(COLO,MEXU);SierraSan Luis,
(0.6-)0.7-1 mm; colour variable,dull green to glau- 12 Aug 1908,Goldman1408(US);50 km S of La Junta,
cous-green especially on the adaxial faces. Seed above Rio Ancho, 8 Mar 1967, Kimnach & Brandt 866
cones 2-5 X 3-6 cm when open. Seed scales 25- (UC);6 km fromOjito,14 Nov 1967,Kimnach& Brandt
40. Seeds 10-13 X 6-10 mm; integumentthick, 0.6- 1039 (US); 50 km W of La Junta,18 Mar 1963, Little
18908 (K, US); Basuchil,ca. 15 km NW of Mifaca,on
1 mm; macrogametophyte("endosperm"of authors) Cerro la
de Ventana,10 May 1929,Mexia2516 (BM, E,
pinkish when fresh. MICH, MO, NY, US). COAHUILA: Cuautemoc, 10 Nov
Distribution and ecology. United States: SE Ari- 1991,Frankis101 (E, FHO);Mesade las Tablas,10 Nov
zona, SW New Mexico, SW Texas; Mexico: NE 1991, Frankis 115 (E); Careros Pass, 1900, Pringle 2659
CarnerosSta., 12 Nov 1904, Pringle 13207c
Sonora, Chihuahua,Coahuila, Durango, Zacatecas, (A, E);
(MICH,S, US);30 km fromSaltilloat SantaRosa,9 Mar
Nuevo Le6n, W Tamaulipas,San Luis Potosi, Aguas- 1980,Stead& Styles593
(FHO);16 km E of Arteaga.12
calientes, NE Jalisco, N Guanajuato,Quer6taro,Hi- Jul 1958,Straw& Forman1350([Link]);Tejocote.26
dalgo, Mexico, Distrito Federal,Tlaxcala, Veracruz, Apr 1985,Styles,Burlev& FavelaI (FHO);2.5 km S of
and [Link] wide but often scattereddistribution Pico de Cerda,11 Aug 1974, Wendt556B (MEXU).HI-
DALGO: AlongHwy.85, ca. 10 km N of Zimapan,28 Apr
of this most common variety is in the northern
1994,Farjon& D. Mejia327 (E, FHO,HEH,K, MEXU,
part of Mexico concentratedalong the continental U); 3 km NE of Molanguito,5 Aug 1982, Gonzdles-
foothills and lower slopes of the Sierra Madres;the Medrano & Hiriart 12783 (MEXU, MO, NY): Barranca

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Systematic Treatment 233

V V.1 -r CC?....
. . -..

':........ . . ' ; : !
?I "~~~~~~~~~~~~'
??i'?? ? ??- .
r~~~~~~~~~~~~~~~~~~~~~~~~~ '"'
r i ~r, ~? ~ ~ ~: ~ ~~~~~~~~~~~~~~~~~~~~~~~~~~
:r
??,?- ??-?? ? -c ~..
~ ~ ..! ~.~..c...
?
i

~~~~ ~ ~~~~~~~~~~~~~~~~
'

i?~~~
.. . . .

-'4i i
r
I
r,; ??
~ ~~~~~~
~ ....- '

f I ??~~~~~~~~~~
Map3 '
Pb I ' ~? ....'

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30 .: !.......
, . ..........

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insebro d ~~~~~~~a
s 5 v r cerie.

de Tolantongo, ca. 40 km NE of Ixmiquilapan,27 Nov SAN Luis POTosi: San Luis Potosi, "chiefly in the region
1993, Hughes 1853 (FHO, K, MEXU); near Zimapan of San Luis Potosi 22? N, alt. 6000-8000 ft," 1878, Parrv &
("Cardonal& Zimapin"), 1839, Hartweg 440 (BM, FI. K, Palmer 846 (E, F, K, MO, NY, US); 30 km from Zaragosa.
P); 3 km N of Posada del Rey, 7 May 1966, Mears 337E 7 Jul 1955, Rzedowski329 (F). SONORA:Sierra San Jose
(SD); Cerro Santuario, on W slope, 19 Jul 1966, Moran [near U.S. border],3 Aug 1893, Mearns 1623 (US). VERA-
13402 (SD); between Zimapan and Real del Oro, 30 Jun CRUZ:"Mountains near border with Puebla (mapa 36.5
1830, Schiede 955 (M). NUEVOLEON:"74 miles (I 18 km) 21.5)," 5 Feb 1968, Ramnos133 (MEXU); 1-2 km W of
N of Doctor Arroyo," 15 Jun 1970, Fryxell 1341 (NY); mines near La Carbonera,25 Jan 1984, Tailor & Nee 236
DoctorArroyo,Jan 1942, M. Martinez3407 (MEXU); base (BH, CHAPA, F, FHO, INIF, MEXU, MO, NY. U). 25 Jan
of Sierrade San Lorenzo, 7 Nov 1904, Pringle 13207b (F, 1984, Taylor & Nee 250 (CHAPA, F, INIF, MEXU, NY,
K, MICH, S, US); Hwy. 58 E of Iturbide,27 Oct 1984, XAL). ZACATECAS: Mazapil, 6 Aug 1969, Taylor& Taillor
Rushforth 431 (E); along rd. from La Encantada near 5310 (US).
DoctorArroyo, 1 Nov 1984, Rushforth564 (E); 8 km from
Iturbidealong rd. to Linares, 12 Mar 1980, Stead & Styles
615 (FHO). PUEBLA:Crater of Mt. Atitzac, 21 Jul 1966,
43a2. Pinus cembroides Zuccarini subsp. cem-
Boege 177 (MEXU); Acatzingo, between La Candelaria
and El Seco, Gilly et al. 6 (MICH);9 km SW of Alchichica, broides var. bicolor Little, Phytologia 17: 331.
along rd. to Zacatepec, Rzedowski31802 (ENCB, MICH). 1968. Fig. 63F-H

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234 FloraNeotropica

?'rlA . I? '? .I:' ?? ? -I- ?.? ?;r .III'."I;??-

i, ? ?j ""
.`ZIL ?? C rlr??Wr- a?r ????r?
'
I r-
tLL' I ?.....; c??I -.? Is.Y. YI?? ? ?? r
'' r. . 1? i I

"T1--t?- -'-..! i
..... ???I ?? i.; ?ry??? ' :
j04 _I ,. ?C???-.
i. __:'IU' i I '
?*.
?n-
.... ??-.?,..
+
? i
I????i.
-. r. r :i? ,I? ..I .,,
: .. ?h ' i. . r 7.' 1. r. ' - E?"?'l .t..
... "" :? ' ?
'" "' ?r? .. k?.. ?'' ????
? .???? ?" ' i? ? ??
'? ' t.l?Y ??I : I ?? R ?e

?r "~- ???
???J ?,?: ;?' ''' .* ' ?? ?? ;????-
????-?????
;
" i""vrC' ?-...,,,J-ri Y
'I .??-? I?,.. __ 1
a. ; R ?1L' L1
a T J'" ? I?. 2?_i."'
r :?-?
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"V '."?L.L;I','' I'"'"""""""'""'"" i
??I r?
'-? 3C j .?
--??--?j r?-? -?? Y: ;;
?'???'1 r','l??-'
"' ??? ???-??U
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i . i I'??.?? ' I rL J.;1:i .? I ?:
'' ? *::
;M;.'-rr*? ?i?rrr;
'"'' c; ..
?- :?i? i:
?r? ... ' ? . '?.I ?? ?.[Link]'E_.illlr.I : ??.? ' "???iM?1 ?;'
r,
i??YI? '??l?i II . ..?
30N,1. t-ri ;?
1 ...,.
'' ?"'' '?? i? -?? ?r .:
?: r g-.d:??PII
ir ? . ?:
?C "'"?"????? . ?-.;??'?,
?i;:
t ., *t
j *? 1 ..'". .. % I;
?"?1? r
.... -r? ' .1? ? ? '?Luiri?r;':i '"t-??d
...
I;1LLLI YYT 'i;; ?? ..' ?:;
r??-?- i ;?I?-.
;':&'?
? ""- - ' r- ?. 1 ..u?i?? .???; ? ??:.
"':" ??r :? ??.???l:iL! I' : ?;
: ? ?? . ;?. ?? 1I;EZ ?
-??,

irr
" --8:?.....,.. .I,
''
i
"? '?"?

c., ... .3L

..,..
.-.c*.....

'rn?ilu?r,?,?'";"
r
i ?C?
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;rf:???'

;r
:'*

-??i?-?
?? '

Map 36. Pinlrs rmrhTOIC/rs VET. hi?olor (circles); P. rrnhmides subsp, c,ricrhcnsis (trianles).

Pinus discolor D. K. Bailey & F. G. Hawksworth,Phyto- the ground, or ascending. Crown an open or dense
logia 44: 130. 1979; Pinus culminicol Andresen & shrubor broad,open, [Link] oval-
Beaman var. discolor (D. K. Bailey & F. G. Hawks-
cylindrical, the terminal buds 8-10 x 3-5 mm.
worth) Silba, Phytologia 56: 490. 1985. Type. United
States. Arizona: Coronado National Forest, Santa Rita Leaves in fascicles of 3, rarely 2, 4 or 5, persisting
Mtns., Madera Canyon, 20 May 1968. Little 23011 4-5(-7) years, curved or less often straight,more or
(holotype, US; isotypes, A, K, NY). less rigid (especially short leaves), 2.5-5(-6) cm X
Pinus johannis Robert-Passini, Adansonia, ser. 2, 18: 0.8-1.2 mm; bicolorous, dull green to grey-green on
366. 1978; Pinus culninicola Andresen& Beaman var.
the abaxial face, usually glaucous-whiteon the adax-
johannis (Robert-Passini) Silba, Phytologia 56: 491.
1985. Type. Mexico. Zacatecas: Puertoel Dique, 3 km ial faces, divided by a green [Link]:Leaves
W of Concepci6n del Oro, 24 Sep 1975, Robert5936B epistomatic, in 2-3 lines on each adaxial face. Seed
(holotype, P; isotypes, ENCB, INIF). cones 2.5-4.5 X 3-5 cm when open. Apophysis
slightly raised, radially keeled or ribbed. Umbo flat
Shrub or small tree, height to 3-12 m, dbh to or slightly recessed, prickle absent. Seeds 10-12 x
10-80 cm. Trunk monopodial, very short (shrubs), 6-9 mm, integumentthick, 0.5-1 mm; macrogameto-
branching immediately above the ground, or short, phyte ("endosperm"of authors)white when fresh.
contorted or more or less straight, branching low. Distribution and ecology. United States: SE Ari-
Branches of first order long, spreading or trailing on zona, SW New Mexico; Mexico: NE Sonora, Chi-

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Systematic Treatment 235

n E

FIG. 62. Pinus cembrmidessubsp. cemhbridesvar. cembnides (A-C. Styles. Burley & FcavelaI; C, D. Steadl& Styvles
593; E-G, Farjon &dD. Mejia 327). A. Branch with leaf fascicles and pollen cones. B. Leaf fascicles. C. Cross sections
of leaves. D. Branchwith short leaves and ovuliferous cones. E. Ovuliferouscones. F. Seed scale (three views). G. Seed
(two views). (Magnifications:A. B. D,
D-G O.5; C X40.)

huahua,Coahuila, Nuevo Le6n, W Tamaulipas,Du- ably with several populationsstill to be discovered.

rango, Zacatecas, N & W San Luis Potosi. A more Its habitat is similar to it, but var. bicolor occurs
scattereddistributionthan var. cembroides,but prob- more often on exposed, rocky, arid slopes or ridges

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236 Flora Neotropica

and less often mixed with other pines in open forest low or sometimes [Link] in fascicles of (2-)
types. It is commonly mixed with shrubby species 3, very rarely 4, spreading, persisting 3-4 years,
of Quercus, Juniperus, sometimes Pinus cembroides mostly straight,lax, (2.5-)4-7(-8) cm x (0.7-)0.8-
var. cembroides, Agave, Yucca, Opuntia, Arctosta- 0.9 mm, dull green to greyish green. Stomata. Leaves
phylos, Cercocarpus, Garry'a, etc. The altitudinal amphistomatic, in 2(-3) intermittent lines on the
range, although extensive, lies somewhat higher than convex abaxial face, in 2 lines on each adaxial face.
with var. cembroides; in Mexico it was found at Apophysis of seed scales raised, transverselykeeled
1700-3000 m, one collection (Frankis 179) at 3500 or radially keeled or ribbed, mostly reddish brown,
m. The climatic conditions are similarly harsh, with sometimes [Link] 10-16 X 6-10 mm; integ-
annual precipitation usually <500 mm and a 7-8- ument thick, 0.5-0.8 mm; macrogametophyte("en-
month-long dry season. Phenology: Pollen release is dosperm"of authors)pinkish when fresh. Cotyledons
reported to be in June-July (Malusa, 1992); a speci- 5-17, grey-green.
men from Zacatecas with nearly full-grown pollen Distribution and ecology. Mexico: Endemic in
cones (Gonzales G. 77) was collected in June. the Sierrade la Lagunaat the S tip of BajaCalifornia,
where it is estimated to occupy around 20,000 ha.
Representative specimens examined. UNITED The Sierra de la Laguna is a batholitic uprising of
STATES. ARIZONA: CoronadoNational Forest, Santa Rita
Mtns., Madera Canyon, 20 May 1968, Little 23010 (K, granitic rock, flanked by older sedimentary rocks
NY). and rising to 2090 m. Pinus cenbroides subsp. lagu-
MEXICO. CHIHUAHUA: "Soldier Canyon," 26 Sep nae occurs in the highest N part of the range, at
1903, M. E. Jones s.n. (BM, MO, US); N partof San Luis 1600-2050 m. It is best developed on and around
Range, canyon S of Diablo Canyon, 3 Sep 1952, Tucker the
2600 (A); "region E of the ContinentalDivide," Carretas, high meadow (La Laguna) of sandy loam but
21 Aug 1938, White 959 (GH). COAHUILA: Mesa de las occurs also in valleys with coarse gravel or granite
Tablas, 10 Nov 1991, Frankis 114 (E, FHO); Canon de boulders and on granitic slopes. It is commonly
Agua QuatroCi6negas, in Sierrade la Madera,8 Sep 1939, associated with Quercusdevia, which is more abun-
Mueller 3229 (F, GH, MICH, UC); summit of Mt. Jimulco,
dant than Pinus in most areas. No other pines occur
13 km E of Jimulco, 29 Jun 1941, Stanfordet al. 10 (GH,
MEXU, MO, NY, UC). NUEVOLE6N: Los Alamos, II in Baja [Link] dry slopes Yucca,Callian-
Nov 1991, Frankis 123 (E): Santa Cruz, II Nov 1991, dra, and Dodonaea are found in the [Link]
Frankis 137 (E); Cerro Pefia Nevada, on Nuevo Le6n- climate is subtropical; annual precipitation in the
Tamaulipasborder, 15 Nov 1991, Frankis 179 (E); Mun. area is ca. 750 mm, almost all of which comes from
Montemorelos, on Sierra de la Cebolla, 21 Aug 1939,
Mueller 29/6 (GH). SAN Luis POTOSi:21 km S of San summer [Link]: Pollen dispersal oc-
Luis Potosi, 28 Jul 1958, Straw & Forman 1438 (INIF curs in May-June (Passini & Pinel, 1987).
MICH, US). SONORA:Locality unknown, 2 Jan 1890,
Hartman 367 (A, BM, F, GH. K, NY, S, US); locality Representative specimens examined. MEXICO. BAJA
unknown, 2 Jan 1891, Hartman 475 (K); Sierra Madre CALIFORNIA SUR:Sierrade la Laguna,5 km S of Picacho
Occidental, 1900, Schott s.n. (F); Caion de los Metates, IMt.] Laguna, 15 Mar 1979, Bailey 79-09 (E. NY); Sierra
SW of Bavispe, 24 Jun 1940, White 2828 (GH); Rancho de la Laguna,Cerro San Francisquito,18 Oct 1890, Bran-
del Roble, NE of El Tigre, Sep 1941. White4321 (GH). degee s.n. (UC); Sierra de la Laguna,Mt. El Taste. 14 Sep
ZACATECAS: I km W of Concepci6n del Oro, near Ma- 1893, Brandegees.n. (UC); Sierrade la Laguna,La Laguna,
2 Oct 1899, Brandegees.n. (GH, UC); La Laguna, 22 Oct
zapil, 10 Jun 1979. GonzdlesG. 77 (MEXU); near Mazapil,
Nov 1979, Klaus s.n. (WU). UNKNOWN: "Mts. near Jesus 1977, Breedlove & Axelrod 43313 (MO); S of Pico La
Maria,alt. 6500-7500 ft.," Nov 1892, Nelson 7 (US). Aguja, 22 Oct 1977, Breedlove & Axelrod 43356 (MO):
canyon draining Ci6naga de Todos Santos, 17 Dec 1947,
Carter et at. 2422 (GH, K, US); La Laguna, n.d., Diguet
43b. Pinus cembroides Zuccarini subsp. lagunae s.n. (P); La Laguna, 24 Mar 1939, Gentry 4394 (GH. K,
(Robert-Passini) D. K. Bailey, Phytologia 54: 98. MO); La Laguna, 15 Jul 1969, Gilmartin 1515 (MEXU);
Sierra de la Laguna, 20 km ENE of Todos Santos. 21 Apr
1983. Fig. 63I-M 1987, Hughes 966 (FHO, K, MEXU), 21 Apr 1987. Hughes
967 (FHO, K, MEXU); Sierrade la Laguna. 22 Sep 1930,
Pinus cembroidesZuccarini var. lagunae Robert-Passini,
M. E. Jones 27080 (NY, US); La Laguna, 26 Jan 1906,
Bull. Mus. Natl. Hist. Nat., B, Adansonia 1: 64. 1981;
Nelson & Goldman 7457 (A, US); Sierrade la Laguna. 18
Pinus lagunae (Robert-Passini)Passini, Phytologia 63:
Oct 1985, Tenorio L et al. 10554 (MEXU, MO); La
337. 1987. Type. Mexico. Baja California Sur: Sierra
17 May 1959, Thomas 7879 (GH. MEXU, SD,
de la Laguna, 15 Feb 1978, M.-E Rohert-Passini10021 Laguna,
UC, US).
(holotype, P; isotypes, ENCB, INIF).

Tree, small to large, height to 20-25 m, dbh to 43c. Pinus cembroides Zuccarinisubsp. orizabensis
80-120 cm. Trunk monopodial, short to medium D. K. Bailey, Phytologia 54: 89. 1983.
size, contorted or more or less straight, branching Fig. 63A-E

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Systematic Treatment 237

V I

A a //
/

C /

FIG.
FIG. 63.
63. A-E. Pinuscembroides
A-E. Pinus cembroides [Link]
subsp. (A, B,
orizabensis(A, B, Bailey
BaileY83-01;
83-01; C,C, Nee 32941;D,
Nee 32941; E, Nee
D, E, Nee & Taylor
TaYlor28855).
28855).
F-H. P cembmides
F-H. cembroides subsp. cembroidesvar.
subsp. cembroides [Link]
bicolor(F,(F, G, Hartman 367;
G, Hartman 367; H, Frankis 1)4).
H, Frankis I-M. P
114). I-M. subsp.
cembroides subsp.
P. cembroides
lagunae Hughes967;
(I-L, Hughes
lagunae (I-L, 967; M, Brandegees.n.,
M, Brandegee 14 Sep
s.n., 14 1893).A.
Sep 1893). Branchwith
A. Branch with leaf
leaf fascicles
fascicles (type specimen). B.
(type specimen). B. Leaf
Leaf
fascicles. C, D.
fascicles.C, D. Ovuliferous
Ovuliferouscones.
cones. E.
E. Seed. F. Branch
Seed. F. Branchwith
with leaf fascicles.G.
leaf fascicles. G. Leaf
Leaf fascicle,
fascicle, with
with enlarged
enlarged (x 3) detail.
detail.
H. Ovuliferous
H. Ovuliferous cone. 1.
cone. i. Branch
Branchwith
with leaf
leaf fascicles.
fascicles.J. Leaf
Leaffascicle,
fascicle,with enlarged(x
with enlarged ( x 3) detail.
detail. K.
K. Ovuliferous
Ovuliferous cone.
cone. L.
Seedscale
Seed scale (three views). M.
(threeviews). M. Seed
Seed scale
scale and
and seeds.
seeds. (Magnifications 0.5.)
(MagnificationsxX0.5.)

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238 Flora Neotropica

Pinus cembroidesZuccarini var. orizabensis (D. K. Bai- MICH,MO, NY, XAL); FrijolColorado,9 Mar 1980.
ley) Silba, Phytologia 68: 48. 1990; Pinus orizabensis Wentlt& Bailey 2495 (MEXU). TLAXCALA: Santa Maria
(D. K. Bailey) D. K. Bailey & F. G. Hawksworth, las Cuevas, 28 Jan 1982, Lott & Wedt P-134 (MEXU).
Novon 2: 306. 1992. Type. Mexico. Puebla: Along
Hwy. 140, ca. 10 km SW of San Salvador el Seco, 23 Uses. Although not a timber tree in most areas
Feb 1983. Bailey 83-01 (holotype, MEXU; isotypes, due to its low statureand low and heavy branching,
CHAPA, E, INIF, K). Pinus cem,broidesis nevertheless an economically
Small tree, height to 8-10 m, dbh to 50-60 cm. importantspecies of pine in Mexico. Its principal
Trunkmonopodial, short to medium size, contorted value for local economies lies in the edible seeds
or more or less straight, branchinglow. Bark thick, (piiones), which are regularly harvested and mar-
keted. Furtheruse is made of its wood for carpentry,
rough and scaly, with shallow and irregularfissures,
or sometimes for timberwhere there is no other pine
grey, exposing orange-yellow inner bark. Leaves in
fascicles of 3(-4, rarely2 or 5), persisting3-5 years, species available,as in [Link] to its
curved or less often straight, more or less rigid adaptationto semi-arid environmentsand extensive
(especially short leaves), (2-)3-5(-6) cm X 0.7-1.1 range it is also an importantshrubor tree for aspects
mm. Seed cones 3-5(-7.5) x 4-6 cm when open. of land management,such as watershed protection,
Seed scales 30-50. Seeds 10-14 x 6- 10 mm; integu- preventionof erosion, and as a shade tree in agro-
ment thick, 0.7-1.1 mm; macrogametophyte("endo- forestry(see papers in Passini et al., 1988).
The taxonomy of Pinus cembroidess.I. is consid-
sperm"of authors)pinkish when fresh.
Distribution and ecology. Pinus cembroides ered to be complex and still debatable(Styles, 1993).
In recent years several segregate taxa from the "ar-
subsp. orizabensis occurs at the southeasternmost
extension of the range of P cembroides in Mexico. chetypal" P cembroides have been proposed at dif-
It has a relatively restricted distributionin Puebla ferent ranks. It will be argued below that much
and adjacentTlaxcala and W Veracruz,primarilyin of the ensuing confusion has been caused by the
a high basin W of the Pico de Orizaba-Cofre de applicationof methodsmore suitableto study within-
Peroteaxis of high [Link] isolatedpopulation population variation(as practised in forest genetics
occurs ca. 15 km E of the city of Tehuacanin Puebla. research)than variationamong taxa.
The region is semi-arid,with scattered,small extinct Pitnus c'emnbroideswas described by Zuccarini
volcanos. The altitudinalrange is 2100-2800 m; the (1830) from a specimencollected by W. F. Karwinski
soils are mainly of volcanic origin. It forms open in September1827 nearSultepec in Hidalgo,Mexico.
woodland or sometimes dense, small forests with The holotype (M) is only a small branchletwith leaf
fascicles and buds; on the label it is said to come
pure stands or, more commonly, it is mixed with
from near the church of Santa Cruz. From the pro-
Juniperusdeppeana and/orJ. flaccida. Occasionally
Pinus pseudostrobus is present, or the pinyon pines tologue it is evident that Zuccarinialso had cone(s)
are mixed with Quercus sp. Contraryto claims by and seeds. As he mentioned in a second publication
of the species (Zuccarini, 1832), at the time only
Bailey (1983), Bailey and Hawksworth(1988), and
four other Mexican pines were known; a similarity
Perry(1991), P cembroidessubsp. cembroidesdoes
occur in the same area and may be locally sympatric with P cemlbrain its seed scales and seeds led to
with it, e.g., near Ajalpan in Puebla. The climate of its epithet. Shortly afterwards,Schlechtendal(1838)
the region is characterizedby a long dry season describednine Mexicanspecies, seven of which were
from November to May, but relatively abundant true pines, and among these were the four that
mm) occurs during the sum-
Zuccarini had mentioned, but not his new species.
precipitation (800-900
mer. Due to high altitude,frost may occur in Decem- Instead, Schlechtendaldescribed under No. 5 a new
[Link]: Not recorded,but presum- species, P Ilaveana, based on materialcollected by
Schiede between Zimapainand Real del Oro, Hi-
ably pollen is dispersed during May-June.
dalgo. Loudon (1842), who had received material
Representative specimens examined. MEXICO. from Schiede, took up P llaveana as the accepted
PUEBLA:15 km E of Tehuacin, 7 Mar 1980, Bailey 80-05
(COLO): Libres, 10 Mar 1980, Bailey 80-11 (COLO);San name and cited P cembroides as a synonym. End-
Salvador el Seco, 8 Mar 1980, Bailey & Wendt80-06 licher (1847) reversed this, realizing the priority of
(MEXU); Tepeyahualco.22 Jun 1976, Calzada & Fabian Zuccarini'sname, but cited the locality of Schiede's
2487 (F. NY). VERACRUZ: Near Cerro (VolcAin)Pizarroat collection as the correctone and Sultepec as wrong,
Trinidad,21 Jun 1976, Calzada 2460 (F, NY); near top of
pass on Cord6nLa Vigia Alta, 5 km NW of FrijolColorado, perhapsbecause he thoughtthey had come from the
24 Aug 1986. Nee 32941 (F. MO, NY); 2 km NW of Frijol same area. Little (1966), who apparently had not
Colorado, 17 Jan 1984. Nee & Taylor 28855 (F. GH. K, seen Karwinski's material, cited Endlicher for the

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Systematic Treatment 239

type location-an error that also has been made'by than those in many other collections of this taxon,
Parlatore,who omitted Karwinskiand cited Zimapan both from the United States and from Mexico. In a
(Schiede) with P cembroides Zucc. It is a good later treatment,Little (1979) reduced this variety to
example of authorsduplicatingerrorsinstead of go- a synonym of P cembroides, a view currently fol-
ing back to the protologueand type specimen. lowed in the Flora of North America (Kral, 1993).
The species has a very wide range. Early collec- Robert (1978) described a new species of pinyon
tions made in the north of that range-e.g., Gregg pine from Concepci6n del Oro, Zacatecas, P. jo-
321, sent to Engelmann in St. Louis (MO)-were hannis, which she had found in 1969 duringecologi-
not surprisingly also described as new species: P cal field work in NE Mexico. It appearedto grow as
osteosperma Engelmann(1848), found near Saltillo a shrub on a heavily overgrazed mountainside at
and "on the battlefieldof Buena Vista" in Coahuila. 2700 m with scattered P cembroides (var. cem-
As the taxon became gradually better known from broides), which attainedsmall tree size there. After
furtherMexican collections, it was realized that it is a descriptionof its charactersand a table with leaf
indeed a variable species, with sometimes distinct counts followed a discussion in which no critical
allopatric populations, but for a considerable time comparison with the most obvious taxon, P. cen-
proposals of new taxa as segregates from Mexican broides var. bicolor, was made. In the table, Robert
P cembroides were not made. Instead, Voss (1907) noted about 10% of fascicles of P johannis with 2
reduced several species described around the mid- leaves; yet later authors (Bailey & Hawksworth,
nineteenth century from the SW United States to 1979, 1983) never mentioned less than 3. Older
varieties of P cembroides,but his treatmenthas not fascicles may lose a leaf before falling entirely;
generally been accepted, although it was in part counts should be made among new fascicles only
followed by Martfnez(1945, 1948; see also Farjon, for this reason. Apart from the shrubby habit and
1984). After Little's (1968) segregationof two varie- chemical differences(Zavarin& Snajberk,1986), all
ties, one from P edulis and the other from P cem- charactersfall within the range of variabilityfound
broides (var. bicolor), renewed interest led into the in P cembroides var. bicolor. A comparison of the
segregationof more infraspecifictaxa, most of which habit of P johannis with P culmninicola(Robert,
were later raised to specific rank by their respective 1978) is unjustified:their architecture(in the sense
authors. We will deal with these separately in the of Halle et al., 1978) is quite different. Pinus jo-
following discussion. hannis, even when a shrub of 3 m, remains essen-
tially monopodialand exhibits no adventitiousroot-
Pinus cembroides subsp. cembroides var. bi- ing of multiple, decumbent stems. Indeed, as
color- When Shaw (1909) described the taxa he remarkedon the label of Frankis 179 (E), that high-
recognized for Mexico underP. cembroides,he cited altitudecollection (3500 m) which was identifiedas
Hartman367 (A, US, NY) with epistomatic leaves [Link] is a small tree, not a [Link]
in fascicles of 3. Whetherhe would have recognized factors are responsiblefor its shrub-likehabit, and it
this material as taxonomically distinct may seem may be consideredan ecotype on extremelyexposed
doubtful, given his inclusive concept of the species, high-altitudesites. Recently, Passini (1994) reported
but it nevertheless disagrees with his description P. johannis to be dioecious, but this does not occur
"stomatadorsal and ventral."Martinez(1948), who in pines: it may be that certain trees fail to produce
by then had accepted two segregates, occurring one or the other sex of cones in some years, but that
mainly in the United States, as full species following does not mean the species is dioecious. The species
practice established there, followed Shaw for his was then sunk into synonymy with P cembroides
treatmentof P. [Link] firstauthorto recog- var. [Link] the rankof species would,
nize the taxonomic significance of the absence of as was suggested to her by E. L. Little, retain P
stomataon the abaxial face of the leaves in many of johannis against P discolor (see below).
the collections from the United States and N Mexico Bailey and Hawksworth(1979) made a detailed
then known as P. cembroideswas Little (1968), who study of Little's taxon and raised it to the rank of
described materialfrom the Santa Rita Mountainsin species, proposing the new binomial Pinus discolor
SE Arizona as P cembroides var. bicolor. The type because "the epithet bicolor could not be used ac-
collection (Little 23011, K, US) is epistomatic, but cording to the ICBN at specific rank in Pinus."
it lacks the conspicuous white bands of stomata The combination Pinus bicolor (Maxim.) Parl. was
found in much other material,even in dried herbar- merely cited as a synonym (Parlatore, 1868: 418)
ium [Link] leaves are also less wide (1 mm) and therefore not validly published (Art. 34.1c);

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240 Flora Neotropica

therefore, there was no good reason to abandon different monoterpenechemistry based on work by
Little's epithet. Since no name has priorityoutside Zavarin and Snajberk(1985). The chemistry aside,
its own rank (Art. 11.2), Bailey and Hawksworth's all these differences are based on statistical evalua-
choice must be accepted if the taxon is to be recog- tions of morphometricsusing continuous characters
nized as a species. If P johannis and P discolor with highly overlapping states (e.g., Zavarin &
are considered conspecific, the correct name for the Snajberk, 1985, on needle length/"finess index" in
species would be P. johannis because of [Link] fig. 5).
has only a few more distinct characters than the Furtherobservationson growth and habit by Pas-
epistomatic leaves; one is the relative flatness of the sini and Pinel (1987) either had an ecological basis
apophysis combined with the small, recessed umbo. or were not significantlydifferentfrom P cembroides
To recognise these as sufficient for the recognition subsp. cembroides; incidentally,the thickness of in-
of a full species seems prematureuntil we know teguments was now given as 0.5 ? 0.2 mm. The
more about the true genetic distinctions between chemical differences reportedby Zavarin and Snaj-
these taxa. Poorly researched new combinations, berk (1985) and the differences in leaf characters
such as those proposed by Silba (1985b) only help (mainly length and shape) in conjunction with its
to confuse the taxonomyand nomenclatureof pinyon apparentlong-time isolation by a major geological
pines. Still other combinationshave even been seri- event (the separation of Baja California from the
ously proposed without an understandingof nomen- continent) may merit the subspecific recognition,
claturalrules (Zavarin, 1988) and are invalidly pub- which we maintain here. As with other segregates
lished-which does not mean that they may not be from P. cembroides,allozyme and especially cpDNA
taken up in the prolific literatureabout these pines. studies are recommendedto investigate these taxo-
Most morphologicaldifferences between P cein- nomic questions on a more direct genetic basis.
broides var. cemnbroidesand var. bicolor are highly
overlapping, such as leaf number,leaf width [mea- Pinus cembroides subsp. orizabensis- Bailey
sured by Bailey and Hawksworth(1979) as "fascicle (1983) segregated the southernmostpopulations of
width"], and shape of umbo, which can be flat in P. cembroidesas subsp. orizabensis. Apparently,his
some scales of P [Link]. attentionwas first drawn to this taxon when he saw
a cultivatedtree at the Royal Botanic Gardens,Kew.
Pinus cembroidessubsp. lagunae - Althoughcol- As interpreted by Bailey (1983) and Bailey and
lected quite early (e.g., Brandtegees.n., 18 May 1893, Hawksworth (1988), P cembroides sensu Gordon
UC; E. W Nelson & Goldman 7457, 26 Jan 1906, (1846; 1858), based on a now apparentlylost collec-
US) as Pinus cembroides,and fairly regularlysince, tion by C. T. Hartwegmade "in the cold districts of
this taxon was only recently recognized as distinct the mountain Orizaba, near the village of Chi-
by Robert-Passini (1981), who classified it as a chiquila, ... at an elevation of 10.000 feet above
varietyof P cembroides.A studyof seed and seedling the sea," belongs to this subspecies. The village of
charactersled her to this conclusion; however, two Chichiquila is some 25 km NE of the volcano, at
main differencescited in the diagnosis are erroneous. only ca. 2000 m. Gordon describes P cembroides
The integumentwas incorrectlygiven to be 0.1-0.3 with rathershort (2.5-3.5 cm) leaves in fascicles of
mm thick, and there is no significant difference in 3 and with large cones, of which the paperpublished
the mean numbersof cotyledons between the species in 1846 gives an [Link] recent collections
and its variety (Robert-Passini,1981: table 3). With- from that location are available (Bailey, 1983, and
out any explanation,Passini (1987), presenting"sev- our extensive database). His observation that the
eral additionalcharacters"to justify raisingthe taxon cone is large and resembles that of P cembra L.
to species rank, then gave the thickness of the seed (hence cembroidess")is, in fact, taken from Zuccar-
coat as 0.2-0.9 mm, which nullifies the distinction ini, but in 1858 Gordon makes the epithet his own
earlier given with P cembroidessubsp. cembroides. and synonymizes P cembroides Zucc. with P lla-
Material we measured ranged from 0.5 mm to 0.8 veana Schiede ex Schlechtendal,thereby creating a
mm. In the meantime, Bailey (1983) had raised the later homonym. Not having seen the material on
taxon to subspecific rank, without noticing Robert- which P. cembroides Zucc. was based, Gordon
Passini's errors, on the ground of "additionaldistin- (1846) referredto Hartweg440 from Zimapan,cited
guishing characters"such as longer leaves (based on in Bentham (1840). However, this collection (BM,
averages) and fewer stomatallines per needle (based K, P) represents P cembroides subsp. cembroides
on averages), plus a few even more minor ones and and not the taxon Gordon described, so that his

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SystematicTreatment241

description may indeed be the earliest reference to tomatic), with 4-5 lines on each face. Leaf anatomy:
P cembroidessubsp. orizabensis. Cross section triangular,with a convex abaxial side;
The differentiatingcharacters Bailey (1983) and hypodermis monomorphic, with 2 layers of cells;
Zavarin and Snajberk (1985) have mentioned to resin ducts 1-2, external, on the abaxial side; stele
separatethis taxon from P cembroidesare all quanti- terete; outer cell walls of the endodermisnot thick-
tative and very much of a continuous nature, and ened; vascular bundle single. Pollen cones crowded
none statisticallymore significantthan leaf numbers on the proximalpart (ca. '/2) of a new shoot, ovoid-
per fascicle has been identifiedto date. Furthermore, oblong when mature,5-8 mm long, yellowish, turn-
the claim that P cembroides [Link]. does not occur ing yellowish brown. Seed cones subterminal,soli-
within the area occupied by subsp. orizabensis must tary or in pairs on short, stout peduncles which are
be [Link] collections cited above under covered with subulate-caudatecataphylls. Immature
P. cembroides subsp. cembroides var. cembroides cones subglobose, resinous, purplishbrown, matur-
from Puebla are from this area. With this evidence ing in 2 years. Maturecones subglobose when still
it is at least prematureto raise this taxon to the rank closed, opening with a flattened base and remote,
of a species, as Bailey and Hawksworth (1992) spreadingfertile seed scales, then 3-4.5 x 3-5 cm.
recently proposedwithout presentingany conclusive Seed scales ca. 45-60, only 10-20 fertile, partingand
evidence. spreadingwide except the smaller,infertileproximal
scales, irregular,often curved, thin, concavo-convex,
44. Pinus culminicola Andresen & Beaman, J. Ar- with 1-2 deep, cup-like depressions holding the
nold Arbor.42: 437. 1961. Type. Mexico. Nuevo seeds, borderedby membranousseed wing remnants,
Le6n: Cerro Potosi, near summit of mtn., I Jul margins irregular,erose, yellowish on both sides,
1959, Beaman 2675 (holotype, MSC; isotypes, A, seed cups brown. Apophysis slightly raised, trans-
MONT, US). Fig. 64 versely keeled, rhomboid to pentagonal in outline,
radiallyrugose when dry, marginscrenateor angular,
Shrub, usually decumbent,height 1-5 m, diam. of colour yellowish brown, often resinous. Umbo dor-
stems to 15-25 cm; multi-stemmed, or very low- sal, slightly raised, rhombic in outline, up to 5 mm
branched,individualsforming a spreading,in places wide, with or without a minute prickle, darkbrown,
dense vegetation("matorral").Barkthin, scaly, exfo- often very resinous. Seeds obliquely obovoid, 5-7
liating in small, irregularplates, grey-brown, soon x 4-5 mm, with a I mm thick integument,brown.
weatheringgrey. Branches numerous,the first-order Seed wings absent on seeds partedfrom the scales.
branches often prostrate to assurgent, the higher- Cotyledons8-1 1.
order branchesassurgentto erect, short, thick, rigid Distribution and ecology (Map 37). Mexico: Re-
but resilient (snow cover), forming a dense, rounded stricted to mountain summit areas in extreme SE
to flat crown, usually matted with other individuals. Coahuila and south-centralNuevo Le6n. First de-
Shoots short, thick, rough with decurrent,persistent scribed from Cerro Potosi, Nuevo Le6n; it has more
pulvini, light brown turninggrey. Cataphylls small, recently been found on other isolated summits in the
narrowlytriangularto subulate, apex caudate, mar- region, from where it was identified long ago as P
gins erose, persistentin part,light brown,weathering cembroides. Its altitudinal range is 3000-3700 m,
blackish grey. Vegetativebuds broadly ovoid, the which includesthe highest ridgesof these mountains.
terminal bud 6-10 mm long, the laterals smaller, Its habit is very similar to other mountain "dwarf
slightly resinous; the scales imbricate,free at apex, pines," e.g., P mugo in Europeand P pumila in NE
subulate-caudate,light brown. Fascicle sheaths ini- Asia. Adaptationto blasting,ice- or sand-ladenwinds
tially 6-8 mm long, consisting of 4-6 imbricate, and a short growing season is responsible for this
translucent,ciliate-marginedscales; in maturefasci- habit. Soils are mostly rocky and calcareous. Cli-
cles the scales separateand recoil, forming a rosette matic conditions are not well-known due to a lack
at the base of the fascicle, straw-colouredto grey, of weather stations at these summits, but precipita-
semi-persistent but mainly falling before the leaf tion, some of it as snow, is [Link]
fascicles. Leaves in fascicles of 5 (very rarely 4 or Cerro Potosi, the species forms extensive monocul-
6), in dense tufts on the ultimate branches,erect to tures of close-packed individuals. It occurs there
assurgent, persisting 2-3 years, curved, rigid, 3-5 with scattered,stunted P hartwegii,which indicates
cm x 0.9-1.3 mm, very remotely serrulateto entire, that the climatic tree line is not reached there at
obtuse, greyish green, whitish or glaucous adaxially. around 3700 m. Somewhat lower, on the Sierra La
Stomatarestrictedto both adaxial faces (leaves epis- Marta,Coahuila, P culminicola has been found in a

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242 Flora Neotropica

B a

FIG. 64. Pinus culminicolai(A-C, Stead & Styles 633; D-F, Frankis 155). A. Branch with leaf fascicles and remnant
cataphylls. B. Leaf fascicles. (a) with enlarged(X 10) detail of recoiled scale of fascicle sheath. C. Cross section of leaf.
D. Ovuliferous
D, Ovulifcrous cone. E, Seed
cone. E. scale (two
$ccdscale views). F,
(two vicws). F. Seeds.
Seeds. (Magnifications: A, B,
(Magnifications:A, B, D-F, 0.5; Ba,
D-F, x 0.5; 10; C,
Ba, x 10: C, x 30.)
30.)

scrub-communitywith Quercusspp., Arctostaphylos. ported growing scattered with P culmniicola. Phe-

Ceanothus, Agave, and grasses; on the Cerro La nology: Pollen dispersal has been reportedon Cerro
Viega and the Sierrade Arteaga,Coahuila, a similar Potosi to occur in late July, at 3690 m, which indi-
vegetation, but also Abies and Pseudotsuga, are re- cates a late fertilisationand short growing season.

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SystematicTreatment 243

L IL
??r
1
ii 1, 9
?' ?? ''? ??
t ??. il?
I

I?L., ?. 'I i
Ir I, i. -t ?
. ?"t; r? 1 1 .
.

I ??
r r-- ? ? r?
"- .?',.? tr ?, A:

ri" I ,_ 'ljl ,.?II, I

r t
:?e '?.I??'? ?J
??'' ?I
Ij ?' \ I ,.Zc. ?? I I -L? ?i ?II
I? ? '. ?? .. ?.?":
?IIru . . ' E. .
?r. I 1 ?1 GF ? ??i; ??
? t.
Y. j? ? 1?I? --L- -.
I- ? I ? ' 1 r
r 1 _ 1. ? ?1 .1 71 ???llr??-l.._.:?.?.
r d
I- I i ?'? ?': ?-? ;?
;?..?.? r-? ? ?
.,
?1 ? ? .? ?? rC ? . ? r, .L? ??Y????. ? *??
?'?d
*1 ?r .I. ?? ?
,. ?I:? f.-? ?;.I- .? : ?.?".II
?I I .r.
r ?

?''
rll? ? ?'

r,.r ?I. 'IcJ?'

..
????

?,. ? ?Y
-I
.?..3*i?
-?
i
? ?? ?? '?

QC1 ' "" I -? - ? .I+

m 1 ? ?.
?? 1. ?. ? ? I ??: ? ??: -' 1 -. +??L?.:
" ? ??
'' , .r'C??: " ..
1 ??
'' i.?1:f. ?
9? 'I
? ,? ;::??
Ii i s ? .n 'lc??ll ..... ?.?

Map 37. Pinus culmirlicolu.

Representative specimens examined. MEXICO. COA- vulnerable to fire during long dry periods (Perry,
HUILA:Sierra de Arteaga. 5 May 1908. Villareal & Car- 1991).
ranza 4242 (MEXU); Cerro La Viega. 4 Sep 1975, Rob- Pinus culminicola was described from near the
ert & Passini 6416 (P); Sierra La Marta, 31 Mar 1976,
Pattersons.n. (INIF, MEXU. SD. US); on summit of Sierra summit of Cerro Potosf (3670 m), based on a collec-
La Marta, 18 Jun 1985, Yanez 175A (MEXU). NUEvo tion made on I July 1959 (Andresen & Beaman,
LEON:CerroPotosi. 30 Jan 1975. Bailey 75-46 (NY); Cerro 1961). However, it had been collected much earlier
Potosi, 19 Sep 1961, Beaman & Andresen4511 (B, ENCB, on that mountain by C. H. Mueller in 1934 and
INIF, NY, UC, US); Cerro Potosi, 2 Nov 1973, Castillo &
Villar s.,. (ENCB. INIF, MEXU); Cerro Potosi, 21 Oct
1935. A still older collection of the species (Vil-
1984, Favela 3 (FHO); Cerro Potosi, 13 Nov 1991, Frankis lareal & Caranza4242), from the Sierrade Arteaga
155 (E, FHO); Cerro Potosi, 30 Sep 1968, Medrano & in Coahuila, had likewise been overlooked. Mueller
Delgadillo 1755 (MEXU); Cerro Potosi, 26 Jul 1934, referred his collections to P. flexilis, and Villareal
Mueller & Mueller 1241 (A. F, MICH, MONT,NY. P. US); and Carranza identified their material as P cenit-
Cerro Potosi, 23 May 1982. Sdinchez79 (MEXU); Cerro
Potosi, 22 Mar 1970, Rzedowski& Hinton 17275 (ENCB,
broides. Since Beaman'scollections, it was rediscov-
MEXU, MICH); Cerro Potosi, 16 Jul 1945, Sharp 45779 ered in the Sierra de Arteaga, and also found on
(MEXU); Cerro Potosi, 13 Mar 1980, Stead & Styles 633 other mountains in the Sierra Madre "arch" S of
(FHO. MO); Cerro Potosi. 10 Aug 1982, Torres C. 1081
Monterrey.
(CIIDIR, MEXU). Silba (1984, 1985a, 1985b, 1986) greatly "ex-
Uses. This species is not used commercially, al- panded"the range of this taxon by first including as
though locally it may be used for firewood. It is synonyms (but soon thereafterclassifying as varieties

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244 Flora Neotropica

of P culminicola) P. discolor and P johannis (= crowded near the proximal end of a new shoot,
P cembroides var. bicolor), apparently based on subtendedby subulate,deciduous bracts,subglobose
"similarityof description"in Bailey and Hawksworth to ovoid, 4-5 mm long, pink or purplish, turning
(1983). light yellow. Microsporophylls sub-peltate, with
erose margins. Seed cones subterminal,solitary or
45. Pinus remota (Little) D. K. Bailey & F. G. sometimes in pairs on 5-8 mm long, slender,curved
Hawksworth,Phytologia 44: 129. 1979. Fig. 65 peduncles with semi-persistentcataphylls. Immature
cones ovoid, ca. 5 mm long, purplish,when growing
Pinus cembroides Zuccarini var. remote Little, Wrightia
globose, the apophysaltissue "bulging"between the
3: 183. 1966. Type. United States. Texas: Val Verde
County,Loma Alta, I Apr 1963, Little & Correll 18991
purplish pink umbos, becoming light yellowish
(holotype, US; isotypes, GH, NY).
brown to orange-brown,maturingin two years. Ma-
Pinus catarinae Robert-Passini,Bull. Mus. Natl. Hist. turecones subglobose or globose when closed, open-
Nat., B, Adansonia 1: 70. 1981. Type. Mexico. Nuevo ing with a flat base to form an irregularrosette, then
Le6n:CasaBlanca,nearSantaCatarina,10 Feb 1978, wider than
Robert-Passini 10011 (holotype, P; isotypes, ENCB. long, (2-)2.5-4 X 3-6 cm when open.
Seed scales ca. 25-35, of which ca. 10 are fertile,
INIF).
these partingwidely, the basal scales remainingcon-
Small tree or shrub, height to 3-9 m, dbh to 15- nate, all scales loosely attachedto the rachis, irregu-
40 cm. Trunkshort, contorted,soon [Link] larly shaped,concavo-convex, with incurvedmargins
thick, rough and scaly, exfoliating with thin, scaly and 1-2 deep seed cavities, dull brown abaxially,
plates, in old trees on lower part of trunklongitudi- dark brown to blackish brown on the adaxial side.
nally furrowed,grey to blackishgrey, on young trees Apophysis raised, transversely or radially keeled,
and branches thin, smooth but soon flaking, grey. irregularlyrhombic to pentagonal in outline, upper
Branchesof first order thick, contorted,spreadingor marginangularand irregular,(lustrous) light brown
ascending; branches of second and higher orders to reddish brown. Umbo dorsal, flat or indented,2-
spreadingor ascending, the ultimate branchesstout, 4 mm wide, dark brown, soon grey, with a minute,
[Link] stout, rough with prominent,non- deciduous prickle. Seeds obovoid-angular or ellip-
decurrent pulvini, which leave circular scars after soidal, slightly flattened, 12-16 X 8-10 mm, light
the fascicles have fallen. Cataphyllsca. 5 mm long, ochraceous with a grey tinge; integumentvery thin,
subulate, reflexed, grey. Vegetativebuds ovoid to 0.1-0.4 mm; macrogametophyte ("endosperm")
ovoid-cylindric, the terminalbud 5-7 mm long, the white. Seed wings vestigial, remnants remain on
laterals shorter, acute, not resinous or sometimes the scale when the seeds are dispersed. Number of
slightly resinous; the scales imbricate, appressed, cotyledons not observed.
[Link] sheaths initially ca. 5 mm Distribution and ecology (Map 38). United
long, with imbricate, thin, light brown scales with States: In Texas on the Edwards Plateau and in
hyaline margins, soon reflexing and caducous, not extreme W Texas along the Rio Grande, where the
recoiling to form a rosette, deciduous before the populationextends into adjacentChihuahua,Mexico.
fascicles themselves have fallen. Leaves in fascicles Mexico: In NE and SE Chihuahua,Coahuila, and
of 2(-3), numbers mostly consistent or plants with extreme W Nuevo Le6n. The populations of Pinus
2-3, spreading around assurgent branchlets, per- remota are almost all highly disjunct; the species
sisting 4-5 years, slightly curved to falcate, rigid, has obviously retreated to isolated mountains. Its
(2-)3-4.5(-5.5) cm X 0.8-1.1 mm, margins entire, altitudinal range is 1200-1600(-1850) m (on the
acute-acuminate, abaxial face dull light green or Edwards Plateau in Texas it occurs considerably
yellowish green, adaxial face(s) slightly glaucous, lower-the type collection is from 450 m). It is
whitish stomatal lines [Link] all restrictedto canyons or rocky mountainslopes, often
faces of the leaves, in 2-4 often intermittentlines on on calcareous soil or limestone rock, on dry sites
the abaxial face and (2-)3-4(-5) lines on (each) where pinyon-juniperwoodland is not well-devel-
adaxial face (lower numberspertainto leaves in 3's). oped and the pines often remain shrubby. Annual
Leaf anatomy: Cross section semi-circular or (3- precipitationis 300-500 mm but is extremely vari-
leaved) transverse-triangular, with a convex abaxial able from year to year. Frostis common in December
side; hypodermis uniform, with 1-2 layers of cells; and [Link] species occurs occasionally with
resin ducts 2-5, external, on the abaxial side; stele P cembroides and more rarely with P arizonica var.
terete;outer walls of endodermalcells slightly thick- stormiae,common are Juniperusmonospernma and J.
ened; vascularbundle single. Pollen cones numerous, ashei (in the northernpart of the range), Quercus,

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Systematic Treatment 245

E Y?jEwaa. 41n -

FIG. 65. Pinus remota (A-D, Styles, Burleg:Faveli et al. 4; E, G, H, Little & Correll 18991; F. Frvxell 1334a). A.
Branchwith leaf fascicles and flushing shoots with pollen cones. B. Branchletwith leaf fascicles and immatureovulifer-
ous cone. C. Leaf fascicles. D. Cross section of leaf. E, F. Ovuliferous cones. open and closed. G. Seed scale (two
views). H. Seed (two views). (Magnifications:A-C, E-H, XO.5; D. X40.)

Cercocarpus,and semi-desertplantse.g., Agave lech- April (start of flushing and growth of pollen cones
eguilla, Opuntia, and Fouquieria splendens. Phe- was observed in the type specimen collected I April,
nology: Time of pollen dispersalnot certain, perhaps but at the lowest altitude of the species).

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246 Flora Neotropica

N~~~~~~~~~~~~~~ *

:~~~~ 'i.?
*..... ~. .. .' .. -' ' ":..

Map 38. Pi. us remota.

Representative specimens examined. MEXICO. CHI- of mtns., 18 Jun 1936, Wvnd& Mueller 166 (K, MO, NY,
HUAHUA: Sierradel Pino, 8 km from Rancho La Parra,n.d., S, US). NUEVOLE6N:[Link].6.5 km E of
Bailey 442 (UC); Canon Huahuatan,10 mi SE of Madera, borderwithCoahuila.17Aug 1978,Bailey78-50 (COLO,
24 Sep 1939, Mueller 3477 (MICH, UC); Majalca,24 Aug [Link]);betweenMonterrey and [Link]
1935, LeSueur207 (MO, UC); Cainondel Rayo. toward N 45 on Hwy. 40, 14 Jun 1970. Fr.:rell 1334( (NY).
end of Sierra del Diablo. 25 Jul 1941, Stewart 933 (BM,
GH). COAHUILA: Sierrade Encantada,along Hwy. 53, n.d., Uses. Any use of this species (firewood, edible
Bailey 6 (UC); Sierra de la Madera,Canon del Agua, n.d., seeds) is probably incidental or at most on a very
Bailey I11 (UC); Sierra del Pino, Cafada La Vaca, n.d., small scale due to its scattered occurrence in often
Bailey 423 (UC); La Cuesta, along Hwy. 53, 13 Jan 1975. remote places.
Bailey 75-11 (COLO); Sierra de San Lazaro, [Puerto del
San Lazaro,30 Aug 1939. Mueller 3064 (GH, MICH, UC); Little (1966), in studying the pinyon pines of SW
19-25 km NE of Saltillo along rd. to Monterrey, 10 Jun Texas as part of an ongoing systematic investigation
1963, Gentry et al. 20021 (US); Sierra del Pino, near La of the U.S. representativesof Pinius subsect. Cem-
Noria, 1940, Johnston & Mueller 523 (GH); Cuatro Cie- broides, described P cembroides var. remota as a
negas Basin, near Los Fresnos, 4 Apr 1969, Keil et al. new variety,differing from P [Link]. in its
5994 (US); W slope of Sierra de San Vicente, Canon
Espantosa, 1941, Schroeder 95 (GH); Canon de San Lo- very thin seed integument (0.1-0.4 vs. -0.6 mm)
renzo, along rd. to Saltillo, near Monterrey,26 Apr 1985. and its fewer leaves per fascicle (2, sometimes 3, vs.
Styles. Burley & Favela 4 (FHO); Sierra de la Madera, 3, sometimes 2). According to Little, morphological
Cation del Agua. 13 Aug 1980, Wendt& Lott P16 (ENCB. charactersplaced it with P cembroides; this judge-
MEXU); N part of Serraniasdel Burro. Cation del Colo-
rado, 1974. Wendret al. 116 (TEX); Sierra de la Madera.
ment is based primarily on a comparison with P
near mouth of Cation del Agua, 16 Mar 1977, Wendtet al. edulis (with thick leaves 1-1.4 mm wide vs. 0.7-1
1944 ([Link]); [Puertodel San Lazaro,on NW slopes mm, and a few other,minordifferences that are more

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Systematic Treatment 247

continuous). Little did not study material from NE neitherand, to "avoid [adding] to the complexity of
Mexico, where the greater part of the range of this the cembroidesgroup,"proceededto describe it as a
species is now known to be. Mexican trees appear new species. Its only difference with P. remota is
to have slightly thicker leaves (0.8-1.1 mm) thanthe its shrub-like habit at the type locality; the author
specimens Little studied in Texas. Perry's (1991) apparentlyconsidered the habit of pinyon pines a
citation (Bailey & Hawksworth, 1983) of width of taxonomically [Link] it is not,
leaves (1.3-1.8 mm) is an error; the latter authors unless "habit"were based on an unusualarchitectural
described"fascicle width"-i.e., the total widthwhen model in pines (not Rauh'smodel), which is certainly
the leaves are still connate-thereby introducingan not the case here. "C'est un petit arbre,haut de 3-9
unusualand, indeed, confusing character. m, a couronne 6tendue arondie" (Robert-Passini,
Bailey and Hawksworth(1979) raisedLittle's vari- 1981: 70) can only be read as a description of an
ety to specific rank. Although Little (1966) men- environmentallydeterminedperformanceof a small
tioned the distinct difference in the thickness of pine, which undermore favorableconditionscan and
the seed integument between P cembroides and P will grow to a differentstatureand crown shape. We
[Link],he did not give this character would not have to comment on such spurious taxa
due credit. It is very constant in all Mexican plants if ecologists could appreciatethat taxonomy is also
here considered to belong to that taxon. Based on a serious science.
this character,his variety could have been placed
with P edulis, for which Little gave a thickness of 46. Pinus monophylla J. Torrey& Fr6montin Fre-
the integumentof 0.3-0.4 mm. In addition, Bailey
mont, Rep. Exped. Rocky Mts. 319, t. 4. 1845.
and Hawksworth(1979) mentioneda higher number
Fig. 66
of resin ducts in the leaves of P remota compared
with P cembroidesand P edulis (2-5 vs. 2). There Pitnusfremonltiaitna Endlicher, Syn. Conif. 183. 1847:
seems to be a reduction of this number northward, Caryopitysnmonophylla (J. Torrey& Fremont)Rydberg,
so that the EdwardsPlateau populationof P remota Bull. TorreyBot. Club 32: 597. 1905; Pinus cembroides
Zuccarini var. tnlnolphylla (J. Torrey & Fremont) A.
approachesthe other two taxa in this [Link]- Voss, [Link]. [Link]. 1907(16):95. 1907;
otherdifferencebetween P cemnbroides and P remota Pinus cemntroidesZuccarinisubsp. mtonoph!llt(J. Tor-
is found in the fascicle sheaths, which curve back rey& Fr6mont) E. Murray,Kalmia12:22. [Link].
and form semi-persistentrosettes in P cembroides [Link]:
"Collectedin northernCali-
but only spread while disintegrating in P remota. fornia,"1843-1844,Fremont 367 (holotype,NY;possi-
ble isotypes,GH (Fremonots.n., "secondexpedition"),
Both characterstates are found to be distinctbetween K (Fre!montrs.n., "exped. to California 1843-44")).
species; the recurvedscales also occur in P culmini- PinusedulisEngelmann [Link].r Little,Phytologia17:
cola and in P rzedowskiibut are absent in P quadri- 331. 1968;Pinus californiarum D. K. Bailey subsp.
folia and P pinceana. In these pines, a thick-walled fallax (Little)D. K. Bailey, Notes Roy. Bot. Gard.
seed integument is plesiomorphic; in the species- Edinburgh44: 279. 1987; Pinus no,noplhlla J. Torrey&
Fremont var. fillax (Little) Silba, Phytologia 68: 54.
complex of P cembroides s.l., recurved fascicle 1990. Type. United States. Arizona: Sierra Ancha Ex-
sheath scales are the primitive state. Pinus remota perimentalForest, 4 Jul 1961. Little 18581 (holotype.
US: isotype, A).
appears to be characterized by the loss of both;
Pinus calliJbrrniaruD. K. Bailey. Notes Roy. Bot. Gard.
accordingto Nixon and Wheeler (1992), such losses Edinburgh44: 278. 1987; Pinus imonopihllaJ. Torrey&
are the significant events of speciation. Since P [Link](D. K. Bailey) Silba, Phyto-
remote is distinct in the combinationof these charac- logia 68: 54. 1990. Type. United States. California:
ters compared with all other pinyon pines, while it Anza Borrego Desert State Park, Vallecito Mountains.
is also largely allopatric,and when sympatric,does 2.5 km NW of summit of Whale Peak. 27 Mar 1981.
Bailey 81-08 (holotype. COLO-n.v.: isotypes. E, NY).
not hybridize with P cembroides, its recognition as Pinus edulis [Link](lJ. Torreyin J.C.
a species seems to us to be justified. Ives. Rep. Colorado River 4: 28. 1861. nom. nud.
Robert-Passini(1981) described Pinus catarinae
as a new species from near Santa Catarinaalong the Tree, small and bushy, or a large shrub,height to
roadbetween Monterreyand Saltillo in Nuevo Le6n. 15-20 m, dbh to 40-50 cm. Trunlkmonopodial,
Comparisonof this populationand its characteristics usually short, low-branched, straight or contorted.
was primarily made with P edulis and P remota, Bark thick, rough and scaly, with shallow, longitudi-
and the author,disagreeing with Bailey and Hawks- nal but irregularfissures, exfoliating in small plates,
worth's (1979) inclusion of this population in the inner barkorange, outer barkreddishbrownto grey,
latter species, equated the shrubby pinyon pines to on young trees and branches thin and smooth.

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248 Flora Neotropica

Branches of first order thick, long, spreading or cavo-convex, with 1-2 deep seed cavities bordered
ascending, long persistent,of higher orders numer- by thin remnants of vestigial seed wings (I seed
ous, curved down, spreadingor assurgent,forming a often aborts), ochraceous to light or dark brown.
wide-spreading, rounded but irregular,open crown Apophysis thick woody, raised, pyramidalor obtuse
often extending low above the ground; in young conical, transversely keeled, up to 15 x 20 mm,
trees the crown may be more regular, dense and recurvedor straight,lustrous,ochraceous to yellow-
"beehived" in shape. Shoots short, stout, rigid, ini- ish brown, often resinous. Umnbodorsal, flat or ob-
tially rough with short decurrent,soon weathering tuse-pyramidal,rhombic in outline, up to 10 mm
pulvini, orange-yellow, turninggrey after 1-2 years. wide, centrallyindented,concolorous with apophysis
Cataphvlls short, triangular, rigid and spreading, or lighter, often resinous. Seeds obliquely obovoid,
margins erose, orange-yellow, deciduous before the slightly flattenedon one side, 13-18 x 8-12 mm,
leaf fascicles. Vegetativebuds ovoid-conical, acute, integument thin (0.3-0.5 mm), greyish brown to
the terminal bud 10-15 X 5-7 mm, the laterals grey, macrogametophyte("endosperm")white. Seed
smaller, (slightly) resinous;the scales imbricate,ap- wings rudimentaryon the scale, absent from the
pressed, narrowly triangular,yellowish green, with developed, free seed. Cotyledons6-10.
shortciliate [Link] sheathsof young fasci- Ecology and distribution (Map 39). United
cles 5-8 mm long (up to II mm in N partof range States: Widespreadin California, Nevada, S Idaho,
outside our region), with imbricate,hyaline-margined and partsof Utahand NW Arizona,where the species
scales, soon disintegrating in recoiling but almost is peripatric with P edulis to the east. Mexico:
simultaneously deciduous scales of which only the Restricted to Baja California Norte, mainly in the
basal parts persist (not forming rosettes). Leaves in Sierra Juarez and lower slopes of the Sierra San
fascicles of 1, rarely 2, remote, persisting4-8 years, Pedro Martir;an outlier populationin the Sierra La
assurgentto spreading,rigid, curved at least near the Asamblea forms the southernmostoccurrenceof the
base, (2-)2.5-6 cm X 1.2-2.2(-2.5) mm, in rare species at 29?17'N, 114?03'[Link] Mexico, P mono-
fascicles of 2 with entire margins, acuminate-pun- phylla has an altitudinalrange of (950-)1200-1700
gent, usually lustrous, green, grey-green, or glau- (-2000) m. It is locally common in, or just above, a
cous-green, with a thick, often exfoliating cuticula. high chaparral zone but does not form extensive
Stomata around the leaves, with 12-25 lines (some pinyon-juniperwoodland. Other common pines are
intermittent)of stomata in grooves. Leaf anatomy: P quadrifolia and P jeffreyi, the latter species ex-
Terete, often with 2 grooves markingpotentialdivi- tending to higher [Link]
sion into two leaves; epidermis thick; hypodermis Quercus turbinellacommonly occur with it in areas
multi-layered,with small intrusions into the meso- with granite rock outcrops and on steep slopes. An-
phyll; resin ducts (2-)5-12, occasionally >12 (not nual precipitationis a moderate300-600 mm, con-
observed: see Bailey, 1987), large, some smaller if centrated in the winter and part of it as snow at
there are many, external; stele terete; outer walls higher altitudes. Phenology: Pollen dispersal occurs
of endodermalcells not thickened; vascular bundle in April-May.
single, [Link] cones crowded on the proxi-
Representative specimens examined. MEXICO. BAJA
mal end of new shoots, subtended by triangular, CALIFORNIA NORTE:Sierra La Asamblea, 20 Mar 1982,
chartaceous bracts, ovoid-globose to short cylindri- Bailey82-01 (E, NY);"64 mi. (102 km)SE of Ensenada.
cal, up to 10 mm long when sheddingpollen, initially 3 mi. (5 km) WNW of Santa Catarina,"25 May 1961,
Broder472 (MEXU,US);"E-endof SierraSanLuis"I=
purplishred, then yellowish. Microsporophyllssub- Sierra La Asamblea], 3 Apr 1950, Gentry & Cech 8978
peltate, with an incurvatedistal part, smooth. Seed (MEXU, MICH, SD, US); between La Rumorosa and
cones subterminal,solitary or in whorls of 2-4 on Laguna de Hanson, 26 May 1987, Hughes & Styles 184.
slender, 5-10 mm long peduncles which are decidu- 185 (FHO); 10 km E of La Rumorosa, 22 May 1966,
ous with the cones. Immaturecones globose, ca. 5 Moran 13150 (SD); Agua de Gambucinos,on dry slope E
of Lagunade Hanson,4 Sep 1966, Moran 13496 (SD); 12
mm, green, yellowish green when growing, with km S of La Rumorosa,24 Sep 1966, Moran 13586 (SD);
partly overlapping,more or less flat apophyses, ma- El Toruno. 16 Mar 1968, Moran 14834 (SD); CerroChichi
turing in two years. Maturecones ovoid-globose to de la India,31 Aug 1968.Moran15561(MICH,SD);Tajo
globose when closed, irregularwhen opened, often Canyon, E slope of Sierra Juarez, 13 Feb 1971. Moran
somewhat wider than long, more or less roundedat 18243 (SD); "SierraSan Luis" [= Sierra La Asamblea],
13 km N of Yubai, 18 Apr 1973, Mortrta20540 (SD, US):
base, 4-6 x 4.5-7 cm when open. Seed scales ca. CerroMatomi,near summiton N slope, 3 May 1973,
30-50, only the central 6-12 fertile, partingwidely, Moran 20762 (MEXU, MO, SD, US); 9 km W of La
spreadingbut not reflexed, moveable, irregular,con- Rumorosa, 14 May 1977. Moran 24063 (SD); ArroyoAgua

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Systematic Treatment 249

E .....'?"'

FIG. 66. Pinus monophivllI (A. C, Moran 15561: B, D, E, Hughes & StIles 185; F. G. Little 18581 (sheet I of 2); H-
J,.Little 18581 (sheet 2 of 2)). A. Branchwith leaf fascicles and immatureovuliferous cone. B. Branchwith leaf fascicles
and vegetative buds. C, D. Leaf fascicles. E. Cross section of leaf. F. Branch with leaves and pollen cones. G, H.
Ovuliferouscones. I. Seed scale (two views). J. Seed (two views). (Magnifications:A-D, F-J. X0.5; E. x 18.)

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250 Flora Neotropica

to extend "from long. 111? to 120?, and through a

considerable range of latitude." After the description
of Pinus quadrifolia by Sudworth (1897), for a long
time these two species (and an outpost of P edulis,
which has a range to the east) remained the only
pinyon pines recognized in U.S. and Mexican Cali-
fornia (Munz, 1974).
This situation changed drastically when forest ge-
neticists and chemists began to investigate the bio-
systematics of these pines by means of sampling
usually large amounts of material in a standardized
scheme from several to many populations throughout
much of the often discontinuous ranges of these
species (Lanner, 1974a, 1974b; Zavarin et al., 1980;
Trombulak & Cody, 1980; Lauriers & Ikeda, 1986;
Bailey, 1987, 1988).
What these studies have in common is that they
investigate characters of a continuous nature (mor-
phological, sometimes also chemical, and then the
latter are compound as well as continuous), calculate
mean values per population sample (1-10 trees) and
apply multivariate analysis to describe the variation
among these samples. Simple statistical significance
39. Map
onopyll. Pis tests are employed to determine which of the popula-
tion samples group together as similar and which are
to be considered distinct. Among the morphological
characters, leaf number per fascicle is most fre-
quently used and, indeed, often cited as an important
diagnostic character (Lanner, 1974a; Trombulak &
Cody, 1980) or implicitly assumed to be so (Lau-
Map 39. PininsmonophYlla. riers & Ikeda, 1986; Bailey, 1987). This character is
seldom constant, even on a single branch. Scepticism
about conclusions based on leaf numbers ["none of
America's pinyons has a consistent number of nee-
Grande. 9 km S of La Rumorosa. 30' May 1982, Moran dles per fascicle" (Roof, 1978)] has not been heeded.
30828 (SD); "N of Cerro San Luis" [= Sierra La Asam-
All papers seem to imply that "diagnostic" pertains
blea]. 2 Mar 1963. Moran & Henrickson 10319 (MEXU,
MICH, SD, US); Sierra Juirez, La Tete de la India, 14 km to species delimitation; in neither of these papers is
S of La Rumorosa,7 Oct 1961, Raven 16809 (GH, MICH, there any discussion about the significance of these
SD); "along the road between Mexicali and Alaska, in the characters for taxonomy. Instead, apart from infer-
Sierra Pinal, on the summit of Cuesta Alduncin"[ = Sierra ences about
Juarez, near La Rumorosa], 4 Oct 1946, Wiggins 11239 introgression and hybridization, statisti-
cal significance has been automatically translated
(GH, MEXU. MO).
into taxonomic significance, leading to the descrip-
tion of new species or the elevation in rank of
Uses. Due to its irregular shape and slow growth, formerly named varieties (Lanner, 1974a; Bailey,
this tree is not used for lumber. Locally it is used as 1987). Species are based on "statistically significant"
firewood. The edible seeds are locally harvested and divarications of continuous characters, intermediate
sold at village markets or alongside the highways. samples (rarely individuals as these disappear in the
Pinus monophylla was the second pinyon pine means) are explained by various grades of genetic
described in the botanical literature (Torrey & Fre- introgression in the direction most suitable to the
mont, 1845), after P cembroides, and the first from character states observed (Lanner, 1974a, 1974b;
territory later to become part of the United States. Bailey, 1987). More problematic still, individuals
The location of the type collection is given as "north- wholly at variance with the tabulated character states
ern California" and the range of the species is said of samples (populations) can disappear from the data

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Systematic Treatment251

presentedas taxonomic evidence due to the methods 47. Pinus quadrifolia Parlatoreex Sudworth, [No-
used (e.g., Bailey, 1987). mencl. Arb. Fl. U.S.], U.S.D.A. Div. Forest. Bull.
Implicitin this methodis a (returnto a) typological 14: 17. 1897. Fig. 67
or "essentialist" species concept (Mayr, 1988), in
Pinus Ilaveana J. Torrey, non Schlechtendal (1838) in
this case defined by an ideal "type"describedby the
Emory,U.S. [Link].2(1):208, t. 55.
outcome of multivariatestatistics applied to a set of 1858;Pinusparrnana Engelmann, non G. Gordon&
arbitrarilychosen numerical characters. Intermedi- Glendinning(1858), Amer.J. Sci. Arts, ser. 2, 34:
ates are explained away as "hybrids,"and "aberrant 332. 1862; Pinus cembroides Zuccarini var. parryanra
individuals" are (often?) excluded from the data. (Engelmann)A. Voss, Mitt. Deut. [Link].
1907(16): 95. 1907; Pinus cembroidesZuccarinisubsp.
There is hardly a need for a biologist to emphasize parrvana(Engelmann) E. Murray,Kalmia12:22. 1982:
that this kind of species does not exist in nature. Pinus cembroidesZuccarini var. quadrifolia (Parlatore
What mattersinstead is to extrapolatefrom observed ex Sudworth)Silba, PhytologiaMem. 7: 49. 1984.
variationin apparentpopulationsto species delimita- [Link]:"MountainsE of San
tion. To do this with any confidence,one has to have Diego,"1850,Parry1390(holotype,US; isotypes,NY.
P).
a knowledge of this variation(not its mean values), PinusjuarezensisLanner,Southw.Naturalist19(1):77.
an understandingof the evolutionarysignificanceof 1974;Pinus [Link](Lan-
characters,theirheredityand their plasticityor rigid- ner) Silba, Phytologia68: 48. 1990. Type. Mexico.
ity in relationto the environment. BajaCaliforniaNorte:AlongHwy.2, 2 km W of La
Rumorosa, 4 Aug 1969,Lanner2016(holotype,UTC-
Bailey (1987) segregated Pinus californiarum n.v.;isotypescitedto be in NY andUS arenotthere).
from P monophylla, and recombined P edulis var.
fallax Little as a subspecies of it. His table I gave a Tree, small and bushy, or a large shrub,height to
conspectus of characterscomparingthese three taxa. 10-15 m, dbh to 30-50 cm. Trunkshort, erect, low-
For some characters, the ranges are biased since branched or with a short, clear bole. Bark thick,
exceptions disappearedfrom the data as "indetermi- rough and scaly, with deep lateral and longitudinal
nate trees.""Discontinuous"characterstates, such as fissures,reddishbrown, innerbarkyellowish orange,
fascicle sheath length, may thereforehave been less outer bark weathering grey, on young trees and
discontinuous than they are presented. Resin duct branchesthin and smooth, reddishbrown. Branches
numberis presentedto be 2-7 for P. monophylla,9- numerous, long, spreadingor ascending, persistent;
16 for P californiarumnsubsp. californiarum and the higher-orderbranchesslender,spreadingor assur-
again 2-7 for its [Link], introducinga reversal. gent. Crown rounded, wide and open in old trees,
But Lanner(1974b) reportedto have found a correla- more dense and regular("beehived")in young trees.
tion for this number to site differences, whereby Shoots stout, rough with short decurrent pulvini.
the number increases from mesic to arid sites. The Cataphyllssmall, rigid, broad triangular,with erose
southernCalifornianto Baja Californianpopulations margins, yellowish orange, soon grey and breaking
(all identified as P californiarum) partly occur in off, leaving a small basal [Link]-
less arid environment than many of the northern conical, the terminal bud 6-10 X 4-5 mm, the
populations(identifiedas P monophylla)represented laterals acute, <8 mm long, orange-brown,slightly
by Bailey's samples. Even if there is evidence for resinous; the scales imbricate,triangular,appressed
genetic determinationof resin duct numbers (Lau- but with free, acuminate apex and erose-hyaline
riers& Ikeda, 1986), therestill remainsthe likelihood margins. Fascicle sheaths initially 5-8 mm long,
that under equal test conditions resin duct numbers soon breakingup in recurvingbut early falling scales
would exhibit overlapping states. None of Bailey's (not or rarely forming rosettes). Leaves in fascicles
other characters show true discontinuities between of (3-)4(-5), rarely a few fascicles 2 or 6; the
populations,independentof environmentalgradients. fascicles often disintegratepartly before falling; re-
Morphological characters of the seed cones and mote on foliage branches, persisting (3-)4-7 years,
seeds, presumablyinconclusive, have not been used spreading,curved or sometimes straight,rigid, often
in any of the studies cited here; because of this connivent for some time, (1.5-)2-4(-5) cm x (0.8-)
and other limitations, they cannot even be taken as 1-1.5(-1.7) mm, entire, acute-pungent,often with a
phenetic evaluations of multivariate characters. A partly exfoliating cuticula giving the margins an
more rigorous species delimitation-more appro- erose appearance,dull or lustrousgrey-greento glau-
priate from a theoretical, as well as a pragmatic, cous-green, with whitish stomatalbands. Stomata in
viewpoint (e.g., Stebbins, 1993)-is applied here to two bandsof 4-5 irregularlines on the adaxialfaces,
the "single-leaf pinyon"of both Califorias. separatedby a prominentmidrib, rarely with a few

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252 Flora Neotropica

D - '.".v..
^*tS.s4NSi*y C :.

FIG. 67. Pinus quadrifoliu (A-F, Hluglhes& StYles 168). A. Branch with leaf fascicles and terminal buds. B. Leaf
fascicles. C. Cross section of leaf. D. Ovuliferouscone. E. Seed scale (three views), the bottom view of the adaxial side
with seed depressions and seed wing remnants.F. Seeds. (Magnifications:A, B, D-F, x 0.5; C x 25.)

stomata near the apex on the abaxial face. Leaf layers of cells towards the margins; resin ducts 2,
anatomy: Triangular in cross section; epidermis rarely I or 3-4, external on the abaxial side; stele
thick; hypodermis uniform, with 1-2, or locally 3 terete; the outer walls of the endodermal cells not

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Systematic Treatment 253

thickened; vascular bundle I, terete. Pollen cones moderate300-500 mm, but it is quite variable;most
crowded at the proximal end of new shoots in short of it comes during winter cyclonic storms and there
clusters, subtended by triangularbracts, ovoid-glo- is a long dry season from spring through summer.
bose to short cylindrical, ca. 10 mm long when Phenology: Pollen dispersal is in March-April.
sheddingpollen, purplishred, turningyellowish. Mi-
[Link]
Representative
crosporophylls subpeltate, with an incurvate distal CALIFORNIA
NORTE: SierraSan PedroMartir,6 May 1893.
part, smooth. Seed cones subterminal,solitary or in Brandegee s.n. (UC);"64miles(102 km)SE of Ensenada.
whorls of 2-4 on short, slender peduncles which fall 3 mi. (5 km)NE of SantaCatarina,1.5-2.5mi. (2.5-4km)
with the cones. Immaturecones globose, ca. 5 mm, upstream fromRinc6n." 29 May1961,Broder499 (MEXU,
with rhombicapophyses, yellowish green, soon yel- US); N of Ojos Negros,betweenArroyoLos Chivosand
RanchoCasa Verde.28 Apr 1978. Broomte et al. /998
lowish brown, maturingin two years. Maturecones (NY);47.5 km E of Tecate,20 km W of La Rumorosa.
ovoid-globose to globose when closed, irregular along Hwy. 2, 22 Mar 1958, Carter & Chisaki 1031
when opened, often somewhatwider thanlong, more (MEXU,UC); E of San Jose, SierraSan PedroMartir.
or less roundedat base, 4-6 X 4.5-7 cm when open. belowPotreroLaEncinal,31 Oct [Link] (MEXU.
MO);SierraSanPedroMartir, [Link]
Seed scales ca. 30-50, only the central 6-12 fertile,
MelingRanchandobservatory, 2 Oct 1992.Farjon261 (E.
partingwidely, spreadingbut not reflexed,moveable, U, UC);5 km W of La Rumorosa alongHwy.2. 16 Oct
irregular,concavo-convex, with 1-2 deep seed cavi- 1992,Farjoni275 (E. U, UC);SierraSan PedroMartir.22
ties bordered by thin remnants of vestigial seed km W of observatory,29 Nov 1989, Franklin& Chandlcer
5682 (NY); Lagunade Hanson,2 Jun 1905, Goldmanl1132
wings (1 seed often aborts), ochraceous to light or (A, US);45 km NWof Trinidad
de la Valle,7 Oct 1966.
dark [Link] woody, raised, pyrami- Hastings & Turner66-57 (SD); Sierra San Pedro Martir.
dal or obtuse conical, transverselykeeled, up to 10 along rd. to observatory,
25 Apr 1987,Hughes& Styles
(-15?) x 20 mm, recurved or straight,ochraceous /68 (FHO);San Ram6nCanyon,betweenMissionSan
to yellowish brownor reddishbrown,often resinous. PedroMartirandSantoTomas.20 Sep 1938,Wiggins9055
(US);ca. 3 kmS of NejiRancho,16Sep 1929,Wiggins&
Umnbodorsal, flat or obtuse-pyramidal,rhombic in
Gillespie4161 (MICH,MO, SD. US); La Rumorosa.4
outline, up to 10 mm wide, centrally indented,con- Aug 1969,Lanner2025(ENCB);NationalPark"Constitu-
colorous with apophysis or lighter, often resinous. ci6n de 1857,""northboundaryof nationalpark,"25 Feb
Seeds obliquely obovoid or elliptic, slightly flattened 1964, Madrigal S. 1472 (ENCB, MEXU); E of Lagunade
on one side, 12-18 X 8-12 mm, integument thin Hansonat Ranchodel Conde.24 Mar 1974,ManceraO.
266 (MEXU);ElToruno,[Link] 14835(SD.
(0.3-0.5 mm), greyish brown to grey, macrogameto- UC, US); ArroyoCopal, 24 Aug 1968, Mor(an15442
phyte ("endosperm")white. Seed wings rudimentary (MICH,SD, UC); 1.5 km N of RanchoLa Hechicera,6
on the scale, absent from the developed, free seed. Sep 1969,Moran16591(SD); 10 km N of El Alamo.31
Cotyledons6-8. May 1970,Moran17742 (SD); "Centralplateau.4500-
6000 ft.," 16Apr 1882,Pringles.n. (P. US);CerroChato,
Distribution and ecology (Map 40). United on NE slope of mtn., 3 Jun 1963, Thorne.31988 (MICH).
States:California,in Riversideand San Diego Coun-
ties. Mexico: In Baja California Norte, extending Uses. There is no use of this species for [Link]
from near the U.S. border in the Sierra Juarez S to is locally used for firewood. The seeds are edible
the SW foothills of the SierraSan Pedro Martirat ca. and are harvestedto be sold in local [Link]
30?30'[Link] altitudinalrangeof Pinus quadrifoliais may be tappedon a small scale as well.
900-2400(-2700) m. It grows between the semi- Duringthe Mexican BoundarySurvey by the U.S.
desert and the chaparral scrub zones (and partly Army under direction of Major W. H. Emory in
within the latter)and the mixed coniferous forest on 1850, C. C. Parrycollected a new species of Pinus
the highest partsof the [Link] is more widely in the "mountainseast of San Diego" which was
distributedandoften morecommon thanP. monophy- called P quadrifolia by Parry (msc.) and cited in
lla in the pinyon-juniperwoodland, but occurs often synonymy under P. parrvana Engelmannby Parla-
with it. Pinus jeffreyi is the only other pine with tore (1868). Pittus parryana Engelmann is a later
which it occurs in Mexico. Juniperuscalifornica and homonymof P parrvana G. Gordon& Glendinning,
Quercus turbinella are common; in the chaparral which is a synonym of P. ponderosa. Engelmann
zone many shrubsdominate,e.g., Adenostoma,Cea- (1862) based his descriptionon the same collection
nothus, Artemisia, Cercocarpus,Rhus, Eriodictyon, made by [Link], Torrey(1858) had used a
Arctostaphylos,and [Link] of these mountains later homonym, P. llaveana, to name Parry's pine.
are granitic, but in the south of the ranges more Sudworth (1897) corrected these nomenclaturaler-
volcanic rock is found. Pinus quadrifoliaoften grows rors and took up Parry's manuscriptname for the
in cracks among [Link] precipitationis a correct binomial. The type collection has leaves in

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254 Flora Neotropica

fascicles had 2. Using mean values to demonstrate

the hybrid origin of individuals (multivariateinter-
mediacy) seems to be the wrong approachto infer-
ring hybridity(Wilson, 1992). Hybrid taxa are char-
acterized by the coincidence of various parentaland
some intermediatecharacterstates, and in addition
also by such which do not occur in a parentaltaxon
(Rieseberg, 1995). Presenting data as mean values
may show "overall intermediacy,"not character-by-
characterintermediacyin the samples (individuals).
This is not to deny that some of the trees may be
hybrids between P monophylla and P quadrifolia,
as specimens with a consistently intermediatenum-
ber of leaves (2-3) have been found (Lanner, 1974a:
tables 1, 4), or that no introgressionhas taken place
anywhere. On the other hand, fascicles of 5 leaves
are present in various quantities on P quadrifolia
brancheswith 3-, 4-, and 5-leaved fascicles through-
out its range. This character is extremely plastic,
since it varies not only among trees in an apparent
population,or on single trees, but also from year to
year on a single branch as well (see also Lanner,
1974a).
Apart from these considerations,the inference of
hybridity of P quadrifolia sensu Lanner (1974a) is
based on a limited number of characters,of which
leaf number and position of stomata appear to be
intermediateonly in trees with 2-3 leaves per fasci-
cle. Back-crossing in diploid pines may, of course,
obscure morphological intermediacy. The 2(-3)-
Map 40. Pinus quadrifolia. leaved individuals at La Rumorosa, growing with
P monophylla and P quadrifolia, were thought to
represent Fl hybrids (Lanner, 1974a). Comparison
with artificiallyderived hybridsseem to confirm this
fascicles of 3-5, with ca. 15% in 3's, 50% in 4's, assumption.
and 35% in 5's (see also Lanner, 1974a). In P juarezensis there seem to be no fixed charac-
Lanner(1974a) studied several populationsof Pi- ters, only attributes(sensu Davis & Nixon, 1992),
nus quadrifolia in San Diego County,California,and which may define individuals but not populations
in the SierraJuarez,Baja CaliforniaNorte. Based on (and in this case not taxa either). Lanner's explana-
his findings, of which he believed the number of tion-that the few trees (in two widely separated
leaves in a fascicle to be taxonomically the most localities) with predominantly5-leaved fascicles rep-
informativecharacter,he proposeda new species, P resent the last individuals of a species "being intro-
juarezensis. It was described as a 5- (rarely 4-) gressed out of existence"-is nothing but specula-
leaved pinyon pine with 2 (rarely 1-3) resin ducts per tion.
leaf. Only a few trees remain among "populations" The ovuliferous cones of P quadrifolia are mor-
otherwise dominated by hybrids between P mono- phologically very similar to those of P monophylla,
phylla and the new species; these putative hybrids and usually quite [Link] have very
were designated as P x quadrifolia. Furtherevi- pronouncedapophyses, contrasting with those in P.
dence for this hypothesis was given in a (rather cembroides and its subspecies. The seed integument
weak) correlation between mean resin duct number is thin in both (<0.5 mm), whereas it is thick in
and mean leaf number,whereby2-3-leaved fascicles P cembroides. The fascicle sheaths of both Baja
had an average of 2-3 resin ducts and 3-5-leaved Californian species disintegrate quickly, without

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SystematicTreatment 255

forming the more persistent small "rosettes"found The following names were published when the
in P. cembroides s.l. and some other pines more manuscriptof this monographhad been completed;
distantly related to it. Therefore, P. quadrifolia (in- there has not been an opportunityto study the mate-
cluding P juarezensis) appears to be related to P rialon which these new taxa are [Link] identity
monophylla, and it is unlikely that P. juarezensis needs to be furtherinvestigated, and only tentative
could merely be a variety of P cembroides,while P affinities can be suggested here.
quadrifoliais to be maintainedas a separatespecies,
as Silba (1990) suggested. Pinus yecorensis Debreczy & Racz, Phytologia
78(3): 15. 1995. [Possibly a variety of P. pseu-
dostrobus,described from SE Sonora, well to the
Doubtful Names and Excluded Taxa
NW of its known range.]
DOUBTFULNAMES
Pinus yecorensis Debreczy & Rdcz var. sinaloensis
The following validly published names cannot
Debreczy & Racz, Phytologia 78(3): 18. 1995.
be assigned to any known taxon nor be identified
[Described as similar to the above, but with nar-
unambiguouslyas a distinct taxon. The descriptions rower leaves and smaller seed cones.]
given are too vague and/or general to allow certain
identification, and no original material has been Pinus oaxacana
(Martinez)Mirov var. diversiformis
found that could help in identificationand serve as
Debreczy & Rdcz, Phytologia 78(3): 19. 1995. [P
a type.
oaxacana = P. pseudostrobus Lindl. var. apul-
censis (Lindl.) Shaw; described with green to
Pinus chalmaensis Roezl ex Carriere,Trait6 Gen. brown branchlets,not pruinose, and oblong cones
Conif., ed. 2, 2: 583. 1867. with less protuberantapophyses:see the discussion
of the variabilityof var. apulcensis in this mono-
Pinus depauperata Roezl ex Carriere,Traite Gen.
graph.]
Conif., ed. 2, 2: 573. 1867.
Pinus lawsonii Roezl ex G. Gordon & Glendinning
Pinus endlicherianaRoezl [Link] Roezl, Cat. var. gracilis Debreczy & Racz, Phytologia 78(3):
Grain. Conif. Mexic. 20. 1857. 19. 1995. [Described as differing from the species
in its narrower leaves (measurements cited fall
Pinus heteromorphaRoezl, Cat. [Link]. Mexic. within the range given here for the species) and
28. 1857. smaller, more pedunculatecones.]

Pinus macrocarpaLindley, Edwards'sBot. Reg. 26: EXCLUDED NAMES

62. 1839. Pinus contorta Douglas ex Loudon var. contorta,
[Link]. Brit. 2292. 1838.
Pinus ntontezumaeG. Gordon& Glendinning,Pine-
tum 232. 1858, non Lambert, 1832. Pinus contorta Douglas ex Loudon var. latifolia
Engelmannin Watson, Botany (Fortieth Parallel):
Pinus prasina Roezl, Cat. Grain. Conif. Mexic. 24. 331. 1871.
1857.
Neither of these varieties, despite referenceto the
Pinus skinneri Forbes, Pin. Woburn.223. 1839. latter in Martinez(1948) for the trees in the Sierra
San Pedro Martir,Baja California,occur in Mexico.
The disjunct population in Baja California is by
Pinus subpatula Roezl ex Carriere, Trait6 Gen.
all contemporaryaccounts identical with the Pacific
Conif., ed. 2, 2: 584. 1867.
Coast taxon var. murravana.
Pinus tumida Roezl ex Carriere,TraiteGen. Conif.,
Pinus edulis Engelmannin Wislizenus, Mem. Tour
ed. 2, 2: 585. 1867.
N. Mexico 88. 1848.
Pinus zamoraensis Roezl ex Carriere,Trait6 Gen. Despite referencesin the literatureto the contrary
Conif., ed. 2, 2: 585. 1867. (Martinez, 1948; Mirov, 1967; Little, 1979; Perry,

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256 Flora Neotropica

1991), we have not found a single collection from Pinus quichensis Aguilar, [Pinos de Guatemala32.
Mexico to substantiatethe reports of its (putative) (1953) 1961] Cat. Mus. Natl. Ci. Nat. La Aurora
occurrence there. It does occur along the borderof 64. 1942, n.v.
New Mexico and Chihuahua in the United States
(Critchfield& Little, 1966). This species is cited to be published in the above-
mentioned catalogue; it was republished in "Pinos
Pinus flexilis E. James [Link], Account Exped. de Guatemala"by Aguilar,issued in 1953 and 1961,
Pittsburgh2: 27, 35. 1823. with a description only in Spanish. It seems most
Both Martinez (1948) and Perry (1991) mention likely to be a formof P pseudostrobuswith "predom-
P flexilis [Link] from the summit area of Cerro inantly" 6 leaves per fascicle (how many trees were
Potosi, Nuevo Le6n, at around 3800 m. Martinez investigated?). No type specimen was designated. If
also cites Sharp 45779 from "Cima del Cerro del the material was deposited in GUAT, it may now
Potosfi (Potosf Peak), near Galeana,Nuevo Le6n, at be incorporatedin AGUAT, but we have seen no
3420 m. The material from Potosi Peak (Sharp specimens.
45779, MEXU) is P. culminicola. Some material
from the first location is betterclassified as P flexilis Pinus radiate D. Don var.
radiata, [Link]. Soc.
var. reflexa, despite the rather small cone with London 17: 442. 1836.
straight and more or less rounded apophyses illus-
tratedby [Link] collections are P strobiformis
Martinez(1948) describes the pines on Guadalupe
(see discussion under these species).
Island, Baja California, under this name, but men-
tions the variety hinata in his discussion. He ex-
Pinus ponderosa Douglas ex Lawson [Link],
presses doubts about the validity of the variety, not
Agric. Man. 354. 1836.
having studied these pines in their naturalsurround-
Martinez(1948) erroneouslyrefersto P ponderosa ings. Later studies have established the identity of
Dougl. when describing trees in northernBaja Cali- the insular taxon as var. binata. Thus, if one recog-
fornia (near Tecate and elsewhere) which clearly nizes Pinus radiata var. binata, P radiata var. radi-
belong to P jeffreyi. ata does not occur in Mexico.

ACKNOWLEDGEMENTS
The study of neotropical pines has been a major essential reference collection of pine specimens at
researchfocus of the Oxford ForestryInstitute(OFI), FHO. Supportand collaborationwere also received
Universityof Oxford, for nearly 30 years. Following from the forest authoritiesand staff of the national
an initial emphasis on CentralAmericanspecies with tree seed banks and herbariain Belize, El Salvador,
potential for forestry in tropical countries, Brian T. Guatemala, Honduras,Mexico, and [Link]
Styles extended and changed the scope of research field base for all collection activities (including the
to a taxonomic study of all species occurring south last one carriedout by Aljos Farjonin 1994) was at
of the U.S. [Link] are due to Ron the Escuela Nacional de Ciencias Forestales (ES-
Kemp, initially at OFI and later at the Overseas NACIFOR),and laterat the headquartersof Proyecto
Development Administration(ODA), who first rec- de Conservaci6ny Mejoramientode Recursos Fore-
ognized the need for a taxonomic monograph stales (CONSEFORH) in Siguatepeque, Honduras.
of these pines, enlisted Brian's support in early We are grateful to many of the staff of these institu-
OFI fieldwork in CentralAmerica, and was instru- tions for the logistic support they provided to our
mental in obtaining sustained ODA support for the expeditions, and thank in particularOscar Ochoa
work. Mendoza, Marcus Robbins, the late George Gibson,
Many field trips were undertakenby Brian and Tony Wolffsohn, Graham Chaplin, and Ernesto
co-fieldworkers,and we thankTony Greaves, Justin Ponce. Field assistance was provided by technicians
Stead, Graham Chaplin, Colin Hughes, Peter and tree climbers from ESNACIFORand by Alejan-
McCarter, Kevin Wilkinson, and Sophie Higman, dro TrujilloCorzo from San Francisco,Pet6n,Guate-
mainly from OFI, who helped greatly in buildingthe mala.

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Systematic Treatment257

Many of the staff at MEXU helped on numerous BM, CAS, CGE, CHAP, CHAPA, CIIDIR, COLO,
occasions; Brian particularlythanks Teresa German CREG, CU, DS, E, EAP, ENCB, , FHO, Fl, GH,
and FernandoChiang for their support,while Aljos GOET,GUADAL, HAJB, HAL, HEH, IBUG, IFGP,
is grateful to PatriciaDavila for her help in various INIF, JE, K, LAGU, LL, M, MEXU, MICH, MO,
ways duringhis visit in 1994. Brianalso thanksWilli NCSC, NY, P, PRF,S, SD, SYR, TCD, TEFH,TEX,
Mittakfor provision of additionalbotanical material U, UC, US, W, WIS, WU, XAL.
from Guatemala,and Bill Dvorakand Jeff Donoghue Many colleagues in Oxfordhave helped in various
at CAMCORE, University of North Carolina, for substantial ways in the compilation of the mono-
their ideas and discussion about pine genetics and graph. Use of the botanical database BRAHMS,
[Link] are also due to Rogers McVaugh developed by Denis Filer, and computer mapping
for his collaborationanddiscussions with Brianwhen facilities developed by William Hawthorne,greatly
he visited Oxford in 1988. facilitated the compilation of synonymy, specimen
In Oxford, Anne Garton provided technical sup- citations, list of exsiccatae, and [Link]
port in the early years of Brian's study of pines. In Hughes helped in many ways, and he and Robert
1986, Chistine Surman took over and her help has Scotland critically read drafts of introductorychap-
been invaluable in many ways, especially with han- ters. Others helped by trying out identificationkeys
dling and editing of specimen data. She has also on herbariumspecimens. All these colleagues are
helped with sorting the collections and papers after thanked for their time and efforts put in to make
Brian's death, compiling the bibliographyand much things happen. Michael Frankis from Newcastle-
more. We owe it in large part to her diligence and upon-Tyne is also thanked for his constructive re-
patience that the monograph could be completed. markson several chapters,as are reviewers of vari-
The illustrationsby RosemaryWise contributeenor- ous partsof the monograph.
mously to the value of the monograph,and we would We would like to thank Cynthia Styles for her
like to thank Rosemary for all her hard work, her considerablehelp, concern, and determinationto see
professionalism, and her understandingwhen ex- the completion of the monograph. Finally, Aljos
isting drawings had to be changed. Alison Strugnell would like to thank the members of the Advisory
is thankedfor her help in facilitatingthe many loans Committee,GhilleanPrance(chair),Jeff Burley(sec-
of specimens requiredfor this work. We also thank retary),the late FrankWhite, TerryPennington,and
Sean Turnerfor her assistancein mountingnumerous Colin Hughes for their encouragement,support,and
collections for the [Link] supportwas guidance with the production of this monograph
also received from two students: Richard Smith following Brian's death in 1993.
helped with checking the geographicaldata of speci- Financial support was provided by the Overseas
mens for mapping and Neil Brummitthelped with DevelopmentAdministrationof the U.K. government
sorting the large volume of herbariummaterial for througha series of projects within the ForestryRe-
distributionto other herbaria. searchProgrammeover a 20-year [Link]
We are grateful to the curators of the following support for herbariumwork was provided by the
herbariafor allowing access to collections and provi- Royal Society.
sion of loans, often of types: A, B, BAB, BANS,

LITERATURE CITED
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1987. Typification of Juniperus barbadensis L. and J. chiapensis. Phytologia 10: 417-421.
bermudiana L. and rediscovery of J. barbadensis from - . 1966. A multivariateanalysis of the Pinulschlia-
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Aguilar, G. J. I. 1953, 1961. Pinos de [Link]. 2 - & J. Beaman. 1961. A new species of PimLus from
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Aitchison, J. 1986. The statisticalanalysis of compositional flexilis and P. strobifomiis. Phytologia 22(2): 57-70.
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Anderson, A. B., R. Riffer & A. Wong. 1970. Chemistry cone pine forest. Pages 93-149 in R. N. Philbrick(ed.).
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. 1976. History of the coniferous forests, California oocarpa/Pinus patula subspecies tecunumanii. Com-
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. 1980. History of the maritimeclosed-cone pines, & P. J. Kanowski. 1988. Interpretationof the
Alta and Baja [Link]. Sci. Publ. 120: 1-143. composition of coniferous resin. Silvae Genetica 37:
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Critchfield, W. B. 1957. Geographicalvariation in Pinus Engler & K. A. E. Prantl (eds.), Die natirlichen Pflan-
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. 1966a. Crossabilityand relationshipsof the Cali- 115 in: F. A. Wislizenus, Memoir of a tour to northern
fornia big-cone pines. U.S.D.A. Forest Serv. Res. Paper Mexico. Tippin & Streeper,Washington,DC.
NC-6: 36-44. -- . 1862. Supplement1. Coniferae. In: C. C. Parry&
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NumericalListof Taxa 267

NUMERICAL LIST OF TAXA

la. Pinus leiophylla Schiede ex Schlechtendal& Chamisso 21. Pinus praetermissa Styles & McVaugh
var. leiophylla 22a. Pinus patula Schiede ex Schlechtendal & Chamisso
lb. Pinus leiophylla var. chihuahuana(Engelmann)Shaw [Link]
2. Pinus tropicalis Morelet 22b. Pinus patula var. longipedunculataLoock ex Martinez
3. Pinus contorta Douglas ex Loudon var. murravana 23. Pinusjaliscana Perez de la Rosa
(Balfour) Engelmann 24. Pinus tecunuimaniiEguiluz & J. P. Perry
4. Pinus herrerae Martinez 25. Pinus durangensisMartinez
5a. Pinus caribaea Morelet var. caribaea 26. Pinus lawsonii Roezl ex G. Gordon & Glendinning
5b. Pinus caribaea var. bahamensis (Grisebach) W. H. 27. Pinus pringlei Shaw
Barrett& Golfari 28. Pinus teocote Schiede ex Schlechtendal& Chamisso
5c. Pinus caribaea var. hondurensis (Seneclauze) W. H. 29. Pinus muricata D. Don var. muricata
Barrett& Golfari 30. Pinus radiata D. Don [Link] (Engelmann)Lemmon
6. Pinus elliottii Engelmann var. densa Little & K. W. 31. Pinus attenuata Lemmon
Dorman 32. Pinus greggii Engelmannex Parlatore
7. Pinus cubensis Grisebach 33. Pinus coulteri D. Don
8. Pinus occidentalis Swartz 34a. Pinus avacahuite Ehrenberg ex Schlechtendal var.
9. Pinus ponderosa Douglas ex Lawson var. scopulorum ayacahuite
Engelmann 34b. Pinus avacahuite var. veitchii (Roezl) Shaw
10a. Pinus arizonica Engelmannvar. arizonica 35. Pinus lambertianaDouglas
lOb. Pinus arizonica var. cooperi (C. E. Blanco) Farjon 36. Pinusflexilis var. reflexaEngelmann
lOc. Pinus arizonica var. stormiae Martinez 37. Pinus strobiformisEngelmann
11. Pinus jeffreyi Balfour 38. Pinus strobusLinnaeusvar. chiapensis Martinez
12. Pinus engelmanniiCarriere 39. Pinus rzedowskiiMadrigal& M. Caballero
13. Pinus hartwegii Lindley 40. Pinus maximartineziiRzedowski
14al. Pinus pseudostrobusLindley var. pseudostrobusfo. 41. Pinus nelsonii Shaw
pseudostrobus 42. Pinus pinceana G. Gordon & Glendinning
14a2. Pinus pseudostrobus var. pseudostrobus fo. protu- 43al. Pinus cembroides Zuccarini subsp. cembroides var.
berans Martinez cembroides
14b. Pinus pseudostrobusvar. apulcensis (Lindley) Shaw 43a2. Pinus cembroides subsp. cembroides var. bicolor
15a. Pinus montezumaeLambertvar. montezumae Little
15b. Pinus montezumaevar. gordoniana (Hartwegex Gor- 43b. Pinus centbroides subsp. lagunae (Robert-Passini)
don) Silba D. K. Bailey
16. Pinus devoniana Lindley 43c. Pinus cembroidessubsp. orizabensis D. K. Bailey
17. Pinus douglasiana Martinez 44. Pinus culminicolaAndresen& Beaman
18. Pinus maximinoiH. E. Moore 45. Pinus remota(Little) D. K. Bailey
19. Pinus lumholtziiB. L. Robinson & Fernald 46. Pinus monophyllaTorrey& Fremont
20a. Pinus oocarpa Schiede ex Schlechtendalvar. oocarpa 47. Pinus quadrifolia Parlatoreex Sudworth
20b. Pinus oocarpa var. trifoliata Martinez

LIST OF EXSICCATAE
Abarca y Vazquez,M., 39 (20a). AlejandroZarate,L., s.n. (37).
Abbott, W. L., 1947 (8). Alexander,E. J., 641, 848 (34a); 876 (14al).
Aceves, M., 4 (13). Allard, H. A., 14823 (8).
Acosta S., P., 303 (19); s.n. (22a). Allen, P. H., 6417, 6497 (5c); 7186, 7267, 7268 (20a); s.n.
Acosta S., P. & M. A. Barrios, 232 (32); 232a (22a); 298 (34a).
(22a); 299 (22b); 334 (20a); s.n. (32). Allen, P. H. et al., 6096 (13); 6097 (34a).
Acosta S., P. & E. Neri, s.n. (22b). Allen, P. H. & M. L. van Severen, 7312 (34a); 7324 (14b);
Acuna, J. B., 12364 (7). 7325 (24).
Adams, R. F., 73 (la). Almazon, J., 100 (la).
Aguirre,0., 106 (22a). Almazon, J. et al., 919 (25).
Alain, Brother[A. H. Liogier], 6160 (5a). Alonso, L., Ill (15a).
Alain, Brother[A. H. Liogier] & E. P. Killip, 2135A (2). Alvarado,S., 11051 (43a2).
Alain, Brother[A. H. Liogier] & P. Liogier, 23237 (8). Alvarez A., A., 371, 372, 373, 374, 404 (16); 423 (38);
Alava, R. 0., 1258, 1268, 1310 (24); 1311 (20a); 1321, AA436 (14b).
1326, 1327 (24); 1332 (14b); 1354 (16); 1356 (14b). Alvarez, G. & P. Alaya, s.n. (7).
Alava, R. O. et al., 2439A (11). Alvarez, P. L., s.n. (13).
Alcorn, J. B., 3182 (32). Andrade,R., s.n. (43a2).

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268 Flora Neotropica

Andresen,J. W., 666 (44). Baylis, W. B. H., 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12. 13.
Andresen, J. W. & R. J. Steinhoff, A2(X)2 (20a); A2008 14, 15, 16, 17, 18, 19, 20. 21, 22, 23, 24, 25, 26. 27, 28
(14al); 2014, 2023 (38). (5b).
Andrle, R. F., 17, 98 (20a). Beaman, J. H., 1723, 1789, 2213 (13); 2421 (15a); 2422
Anshapanek,D., 245 (28). (la); 2675 (44); 2866 (15a); 2960 (13); 3063 (34a); 3111
Anthony,A. W., 15. 77 (30). (13); 3121 (34a); 3122, 3173. 3208, 3274, 3297, 3352,
Aquino, A. C., 153 (4); 328, 360, 514 (13). 3642 (13); 3677 (34a); 3856, 4093, 4207, 4306 (13);
Araquistain,M., 3536 (24). 4400 (15a); 4434 (la); 4516 (44); 4519. 4520 (13); 4523
Arce [Rico-ArceJ,L., s.n. (13). (34b); 4526 (22a).
Arce [Rico-Arce], L. et al., 1280 (43al). Beaman, J. H. & J. W. Andresen. 4511 (44); 4520 (13);
Armenta,A.. s.n. (la). 4521, 4522 (ISa); 4523 (34b); 4524 (la); 4527 (14al);
Arnason,T. & J. Lambert, 17134 (5c). 4528 (la); 4529 (16).
Arnold. S.. 117 (26). Becerra.M. E., s.i. (34a).
Arredondo,A., s.n. (16). Becerril, V. M., s.n. (41).
Arsene, Fr&re[G. Arsenel, s.. (la, 28). Beetle. A., M6132 (Ib).
Arsene. G., 1064, 3552, 5141, 5369 (la); 5477 (16); 5581 Bell, C. R. & J. A. Duke, 16552 (43al); 16691 (la).
(la); 5707 (18); 5792, 5793, 5796, 5797, 5820, 5821, Bello, A., s.n. (41).
5822, 5823, 5824 (la); 6034 (18); 6039 (28); 6040, 6041 Benitez, A., 503 (43al).
(18); 6042 (15a); 6683 (28); 6967, 6974 (Ia): 6975 (18); Bennett et al., 843 (43al); 844 (12).
7259 (la): 8574, 8769 (15a); s.n. (la). Bernal, O., s.n. (IOc).
Arteaga, B. et al., 133 (30); 135 (11). Bernal S., D., s.n. (41).
Arteaga,C. & C. Villarreal.s.n. (44). Bernardo,T. A., 6 (27).
Ascencio, D., 35-82 (28). Bernoulli,C. G. & R. Cario, 1079 (24); 1081 (14al); 1085
Asprey, G. F., 3694 (5c). (16); 1086(18); 1089(13); 1090 (5c); 1091 (20a); 1094,
Atwood, J. T., 3118 (5c). 1095, 1096 (34a); 1097 (18).
Avila. J. A. et al., s.n. (43a2). Betancourt,A., s.n. (5a. 7, 8).
Avila, M. G., s.n. (4). Bisse, J., 731 (2).
Avila, M. H., s.n. (5c, 27). Bisse. J. & J. Duek, 9118 (7).
Avila, M. H. & R. Rodriguez,s.n. (4, 38). Bisse, J. & E. Kohler,5930 (7).
Avila H., M. & R. Mufoz, s.n. (38). Bisse. J. & H. Lippold, 12756. 15040, 19318 (7).
Axelrod, D. I., E-l (29). Bisse. J. & L. Rojas, 1359 (2); 1965. 3188, 3605, 3762,
Bailey, D. K., 6, 111, 211, 414, 423, 442 (45); 74-11 3846 (7).
(43al); 75-11 (45); 75-37 (43al); 75-46 (44); 75-78C Blackmore,S. & G. L. A. Heath, s.n. (20a).
(40); 75-79, 75-130A, 75-130B, 75-137 (43al); 76-22 Blake, S. F., 7443 (24).
(40); 78-50 (45); 79-09 (43b); 80-05, 80-07, 80-08, 80- Blakley, E. R., 7150 (47).
11 (43c); 80-12 (43al); 81-08, 82-01 (46); 83-01 (43c). Blanco, C. E., s.n. (12).
Bailey, D. K. & B. E. Berger, 81-17 (43al). Blanco, R. & S. Rodriguez,s.n. (18).
Bailey, D. K. & F. G. Hawksworth, 78-34, 79-04. s.n. Blumer,J. C., 1825 (12).
(43al). Boege, W., 177 (43al); 532 (22a); 776 (34a); 1335 (38);
Bailey, D. K. & K. Snajberk,sn. (43al, 43a2). 1342 (13); 1361a (14al); 1693 (34a); 2019 (14b).
Bailey, D. K. & T. Wendt,80-06 (43c). Boeke, J. D., 99 (13).
Bailey, D. K. et al.. 81-18 (43al); 7901 (43c). Bonpland,M. A. & A. Humboldt,3913 (15a); s.n. (18).
Baldemaret al.. 139 (46). Borgo, G., s.n. (5c).
Ballesteros, s.n. (20a). Boshier, D. H., s.n. (5c).
Balls, E. K., 4115, 4395 (13): 4431, 4653, 4797, 4798, Boullon, F. & F. Brandt,2877 (37).
4799 (22a); 5191 (la). Bourgeau,E., 902 (15a); 904, 905 (22a).
Balls, E. K. & W. B. Gourlay,4230 (13). Bourgeau, M., 902 (15a); 903 (la); 904 (22a); 905, 1016
Balls, N. et al., B4394 (13); B4431 (la); B4654 (28). (la); 2815 (14al).
Bancroft,G., s.n. (3). Boutin, F., 3378 (41).
Banda, R. et al., s.n. (28, 37). Boyas, J. C., 400 (la); 403 (14b); 638 (15a); 643 (13); 647
Banda-Silva, R., s.n. (10c). (34b); s.n. (la, lb, 22a).
Banda-Silva, R. & J. E. Elizondo, s.n. (13). Boyd, C. E.. 81 (Ia).
Bangham,W. N., 317 (5c). Brace, L. J. K., 1634, 3670, s.n. (5b).
Barclay,G. W., 2698 (20a). Bracomontes,M. R., 253 (43al).
Barkley. F. A. et al., 48 (13); 7284 (43al). Brand, D. D., s.n. (20a).
Barlow, B., s.n. (43al). Brandegee,T. S., s... (I , 35, 43al, 43b. 47).
Barr. R. J. & C. T. Mason, 2370 (15a); 2372 (28). Brattstr6m,H., s.n. (47).
BarreraH., R., 179 (22a). Braud,D., s.n. (18).
Barrett,W. H., 77582 (5c); s.n. (22a). Bravo, J. 0., 7 (19).
Bartholomew,B. et al., 2572 (4); 2831 (la): 3595 (43al). Breckon, G. J., 865 (13).
Bartlett,H. H., 11200, 11934 (5c); s.n. (la, 28, 43al). Breedlove, D. E., 1682 (4); 1683 (20a); 1693 (19); 1704
Battstrom.s.n. (47). (4); 4176, 4177, 4189 (29); 6288 (20a); 6424 (18); 6523
Bauml, J. et al., 1130 (21); 1253 (19); 1254 (15a); 1255 (38); 7053, 7113 (24); 7114 (14al); 7116 (14b); 7117
(28). (34a); 7534 (14al); 7558 (18); 7792 (38); 8609 (13);

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List of Exsiccatae 269

8643 (15a); 8788 (34a); 9222 (15a); 9354 (24); 9506. 94, 95, 96 (28); 98 (14b); 097, 098 (43al): 103 (14b):
9546 (14b); 9654 (16); 9868, 9950 (20a); 10059, 10120 111. 114. 133 (32): [Link].(43al, 44).
(24): 10174(38); i0819(14al); 11000(20a); 11318(24); Carbajal.R. M.. 50. 66 (15a); 73 (18); 74 (43al).
14273 (20a); 14274 (34a); 16521 (4); 18726. 18770(1a): Carillo. A., s.n. (20a).
18771, 18826(25); 18860 (1a); 19182 (20a); 21763 (18); Carlquist.S.. 442 (30).
24140 (20a); 24282 (13); 25045 (18); 25047 (38); 25076 Carlson. M. C., 281. 413. 449, 612 (24); 801 (20a): 860
(16); 33414 (15a); 33452 (20a); 39665 (16): 41699 (28): (18): 860 (24): 1421 (28): 1617 (24): 1690 (34a): 1843
41821 (20a); 44282 (19); 44407, 44429 (28). (20a): 1911 (16): 2172 (28): 2348 (20a): 2899 (34a):
Brecdlove, D. E. & F. Almeda. 45504 (19). 3924 (13).
Breedlove. D. E. & D. 1. Axelrod, 43313. 43356 (43b). Carranza,M. A. & A. Rodriguez. 2370 (43al).
Breedlove, D. E. & F. S. Kawahara. 16831 (19); 16832 Carranza.M. A. et al., 1151 (44).
(4); 17000 (20a). Carrera,J., s.n. (42).
Breedlove, D. E. & A. R. Smith, 21763 (14al); 31304 (28): Carrillo,A.. s.n. (20a).
31932. 32105 (24); 32348 (38). Carter.A. & F. Chisaki. 1031 (47).
Breedlove, D. E. & R. F. Thorne. 18163 (20a); 18279 (19); Carter.A. et al.. 2422 (43b).
18361 (4); 20592 (20a); 21330 (18); 30944 (20a); 30968 Carvajal.S., 3231 (19): 3314 (4): 3389 (la); 3444, 3456.
(18); 31129 (24): 31156 (15a); 31157 (34a). 3478, 3481, 3482. 3490 (23): 3507 (lb): 3540 (19): 3650
Briones. O., s.n. (14b). (40): 3841 (37): 4018 (17): 4027 (19): 4031 (la): 4588
Britten,W. L., 101 (6). (23): 4651. 4703 (37); 5602 (19): 5796 (12): 6096 (37).
Britton. E. G., 6589 (5b). Carvajal.S. & M. A. Carranza,3350 (40).
Britton.N. L., 25 (5b). Carvajal,S. & M. C. Luce, 3411 (15a).
Britton,N. L. & C. F. Millspaugh,2469 (5b). Carvajal.S. & A. Z. Meza, 4607 (15a): 5967. 5975 (20b);
Britton, N. L. & J. F. Cowell, 12751 (8). 6311 (16).
Britton.N. L. et al., 6457 (2); 6766 (5a); 7319 (2): 14416 Castellanos, 177 (19).
(2). Castelo N.. E.. 446 (16).
Brizicky, G. K. & W. L. Stern. 378 (6). Castillo & Villar.5 (32): s.r. (44).
Brizuela, F. V.. 764 (43al). Castillo, C. G.. s.n. (21).
Broder,R. E.. 205 (29); 472 (46); 498 (11); 499 (47). Castillo. G. & R. Ortega O., 418 (28); 426 p.p. (15a); 426
Broome. C. R. et al., 1998 (47). p.p., 427 (14b): 433 (15a): 437. 442 (13).
Bruff, I. R.. 1215. 1815 (20a). Castillo Q.. D.. s.. (lb. 28).
Brunce & Hansen. 1582 (20a). Castorena,I.. s.n. (14al).
Brunt, M., 2301 (24); 2316 (5c). Cedillo-Trigos,R. & D. Lorence,454 (20a): 540 (24): 1248
Bucher,G., 513 (7). (14al); 1259 (38).
Bunting. G..S. 1306 (20a). Cedillo-Trigos,R.. et al.. 680 (14b): 975 (la): 1252 (27).
Bunting. G. S. & L. Licht. 506 (5c). Cervantes.N.. s.n. (la).
Burch, D., 5508, 5509 (5c); 5549 (34a); 6012 (20a): 6021 Cervantes.N. & R. G6mez. 11 (37).
(14al); 6174 (5c). Cervantes.N. et al.. 31 (16).
Burgers. T. F., 15733 (20a); 15734, 15735. 15736 (24); Chable, A. J.. s.n. (34a).
15738, s... (20a). Chaplin.G.. c28(509) (20a); c60, c61 (5c); c312 (18); c413
Bye. R. A., 1498 (28): 2376 (Ib); 2377 (37); 2380 (10a); (5c).
2407 (37); 3002 (12); 3141 (37); 3156(12); 3175 (43a1); Chargoy,A. E. & M. T. Enriquez,SIA-5A (14al).
6011B (43al); 6107. 6173, 7501 (Ib); 7520 (12); 7580 Chase, V. H.. 7368 (28): 7649 (O1a).
(37); 8592, 8822 (43al); 8823 (12); 8824 (Ib); 8942 Chavelas-P6lito, J. & L. A. Perez. ES-899 (18): ES-900
(43al); 8950 (Ib); 9011. 9077 (43al). (20a).
Bye. R. A. & W. A. Weber, 8214 (Ib); 8215 (la); 8298 Chavelas-P61ito,J. & G. R. Soria, ES-4327 (20a).
(IOb). Chavelas-P6lito, J. & C. S. Zamora. ES-4786 (38): ES-
Bye, R. A. et al.. 12858 (19); 12872 (43al). 4787, ES-4790 (20a).
Caballero.J. & C. Mapes, 991 (26). Chazaro. M., 545 (34a); 643 (22a): 704 (16): 705 (15a):
Caborpet al.. 3 (28); 4 (14al). 736 (21): 751 (20a); 868 (28); 881, 885 (13).
Calder6n,S.. 1426 (20a); 2481 (24). Chazaro,M. & P. H. Chazaro,5438 (43al).
Caldwell & Baker, 7074 (2). Chazaro,M. & A. J. Dominguez. 1466 (22a).
Calzada, J. I., 1014 (20a); 2179 (28); 2460 (43c): 4439 Chazaro,M. et al.. 5449 (43al).
(22a); 5732, 5931 (34a); 7400 (38). Chez. G. J.. s.n. (13).
Calzada, J. 1. & J. Elizondo. 8549 (16). Chiang. F. & F. G. Medrano.3497 (41).
Calzada, J. I. & Fabian,2487 (43c). Chitty.s.n. (2).
Calzada, J. 1. & Horvitz, 2643 (22a). Christy,G. H., s.i. (2).
Calzada, J. 1. & E. Lozano. 4640 (43al). Christy,H.. s.n. (1a); 06/71 (34a).
Calzada, J. . & F. Lozano, 2460. 2482 (43c); 4665 (14b). Cibrian,D. & T. Mendez, s.n. (20b).
Camp, J. T., 83 (la). Clausen, R. T.. 7417 (22a): s.n. (la, 13. 15a).
Campos V., A., 3478 (20a); 3516 (22b). Clausen. R. T. et al., 80-2 (20a).
Candelario,M., 48 (34a); 51 (26). Clemente, Brother,5150 (7).
Capo, M. A.. 5 (37); 12, 13, 14. 15 (28); 19 (14b); 20 Clewell, A. F., 4511 (5c).
(14al); 22 (28); 23, 35 (14b); 36 (28); 38, 39 (14b); 41 Clewell, A. F. & G. Cruz, 4202 (5c).
(28); 42, 44, 45, 46, 47, 77 (14b); 80, 83, 85, 91 (44); Clewell. A. F. & D. Hazlett, 3949 (34a).

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270 Flora Neotropica

Clinton-Baker,H., s.n. (2). De la Torre,J., s.n. (37).

Clokey, 1. W., 4853 (29). De Laubenfels, D. J., 663, 665 (14al).
Cochrane,T. S., 8492 (la). De Le6n. D., 679 (15a); 691 (43al).
Cochrane,T S. et al., 8491 (la). De Puga, L. M., 8762 (37).
Coker, W. C., s.n. (6). De Soto, A. R. & A F Soto, 6543 (14al).
Collins, G. N. & C. B. Doyle, 57 (20a). Diaz, H. B., 2237 (28).
Collins, G. N. & J. H. Kempton,51 (16). Diaz Luna, C. L., 638 (la); 915 (19); 1723 (14al); 1724
Conrad,J. & R. Conrad,21, 25 (37); 3089, 3099 (28). (18); 1724 (21); 2054 (20a); 4542 (43al); 6714 (14al);
Contreras,E., 543 (5c). 9505 (la); 9581 (20b); 9582 (21); 13501 (43al); 18615
Contreras,M., s.n. (14b). (28); s.n. (41, 42).
ContrerasS. H., s.n. (16). Diaz Luna, C. L. et al., 1620 (20a).
Converse, O., 3 (15a). Dice, J. et al., 320 (30).
Conzatti, C., 2225 (15a); 3945 (20a). Diggs, G. & M. Nee, 2470, 2814 (14b); 2935 (22a); 2997
Cook, O. F., 49 (8). (Ib); 3022 (28).
Cook, O. E & C. B. Doyle, 61 (5c); 265 (20a). Diggs, G. et al., 2611 (13); 2651 (28); 3759 (41).
Cook, O. F. & R. F. Griggs, 317 (5c). Diguet, M. L., s.n. (43b).
Cook, O. F. et al., 25 (8). Dirzo, R. et al., 3, 4 (13).
Cooley, G. R., 2485 (6). Dominguez, A., s.n. (22a).
Cooper, E. H. & C. M. Rowell, 2535 (20a). Dominguez, G., s.n. (19).
Cordoba, 1036 (20a). Dominguez, P., s.n. (3, 11, 19, 35).
Cornelius, J., 34 (20a); 36, 38, 42, (5c). Don, D. B. & P. Don, 18800 (13).
Correll, D. S., 43153, 49463 (Sb). Donahue,G., I (24); 29, 46, 70 (34a); s.n. (18, 24, 34a).
Correll, D. S. & H. S. Gentry,22917 (19). Donahue, J. K., 20, 40, 80 (38); 103, 104 (34a); 136, 140,
Correll, D. S. & I. M. Johnston, 19857 (28); 19914 (lOc); 161, 190, 220, 320, 321, 322, 411 (38).
20101 (Ib); 20120 (12); 21492, 21619, 21622 (lb). Donnell-Smith, J., 2182 (13); 2632 (16); 2633 p.p. (5c);
Correll, D. S. & F. G. Meyer, 44623 (5b). 2633 p.p. (18); 2633B, 3622 (13).
Cota, F.M., 10(11). Dorado, O. et al., 1611 (43al).
Cota, H. & H. Le6n, 7439 (29). Dorantes,J. L., 350, 351 (13).
Cota, O., 88 (14al). Dorr, L. J., 2352 (41).
Coulter,T., 712 (29); s.n. (33). Dorr, L. J. & T. L. Atkins, 2321 (43al).
Courbasson& Mahieux, s.n. (4, lOb, 19, 25). Dorr, L. J. et al., 2085 (43al); 2099 (32); 2608 (28).
Covarrubias,L., s.n. (22a). Douglas, D., s.n. (35).
Cox, D. K., HCGI170, 1460 (18); 3141 (5c); s.n. (20a). Douglas, D. & B. A. Krukoff, 10177 (24).
Cox, D. K. & Guzman, 1326 (24); 1382, 1384 (5c). Dressier, R. L., 1663 (18); 2094 (28).
Coyne & W. B. Critchfield,s.n. (18). Duffield, J. W., 1 (47); 2 (3); 3 (35).
Coyne, W. B. Critchfield& Jurrialba,24 (18). Duig, H., 4495 (32).
Crane, C. J., 315 (5c). Duke, J. A. A., M3545 (22a).
Crespo, J. H., s.n. (38). Duncan,T., 2372 (14b); 2391 (13); 2401 (41); 2401A (22a);
Critchfield,W. B., 1 (20a); 2 (22a); 5 (20a); 7 (18); 8 (20a); 2417 (28); 2451 (22a).
79 (18); s.n. (18). Duncan, W. H., 22453 (22a).
Critchfield,W. B. & B. Hallberg, 1 (18); 2 (14b). Dunn, Dziekanowski & Bollibroke, 20254 (13).
Croat, T. B., 24005 (5c); 39356 (43al). Dunn, LeDoux & Wallace, 21800 (14al).
Cruz, Cruz & Espinosa, s.n. (16). Dunn, Walker& Enright, 17208 (la).
Cruz, M., s.n. (14al). Dvorak, W. S., s.n. (24).
Cruz-Cisneros,R., 45 (la); 192 (28); 193 (15a); s.n. (26). Dwyer, J. D. et al., 234, 321 (5c).
Cruz S., L. & M. Ducote, 8336 (20a). Dziekanowski, C. et al., 3474 (20a).
Cuanalo-Rafael,J., I (14b). Earle, F. S., 724 (2).
Cuellar, E., 8 (14b). Earle Smith, C. & S. Kitchen, 5034 (14b).
Cuevas & Rosales, 2403A (4). Ebel, F & J. Bisse, s.n. (7).
Danida, s.n. (5c). Echenique M., R., 46 (11); 47 (35); 49 (11); 52 (34a); 53
Dario, G. F., s.n. (14b). (22b); 60 (14b).
Davalo & M. Avila, s.n. (5c). Eckenwalder,J. E., 51 (13); 95 (20a).
Davidse, G. & A. E. Brant, 31992, 32267, 33017 (5c). Edwards,E. P., 11, 13 (20a).
Davidse, G. & J. Davidse, 9520 (16); 9602A (20a); 9875 Edwards,J. B., 342 (Sc); P47 (18); P163 (5c); P166 (18);
(la). P341 (5c); s.. (32).
Davidse, G. & G. E. Pilz, 31568 (20a). Edwards,M. T., 192 (28).
Davidse, G. & R. W. Pohl, 2076 (20a); 2076A (14b). Eggers, H. F. A. von, 2033 (8); 4172 (5b).
Davidson, C., 7551 (4). Eggler, W. A., 176 (27); 436, 462 (13); 505 (20a).
Dawson, E. Y., 26616, 26754 (5b). Eguiluz, T., 2 (24); 11 (ISa); 15, 16 (34b); 24, 28 (14al):
De la Cerda G., R., 54 (28); 479 (16). 34 (la); 35 (16); 36 (la); 38 (26); 44, 47 (20a);
De la Cerda, M., 1354 (19). 50 (14al); 61 (32); 67, 68 (14b); 69 (10c); 72 (44); 75
De la Cruz. J. A., s.n. (42). (37); 77 (44); 79 (37); 80 (32); 81 (14b); 82 (37); 84
De la Cruz, N., 14 (15a). (14b).
De la Fuente, J., s.n. (15a). Ehrenberg,C. A., 4 (28); 627 (22a); 629 (43al); 633, 636,

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List of Exsiccatae271

640, 643 (28); 645, 646, 647 (34a); 648 (13); 717 (20a); Fosberg, W., 11, 67 (la).
913 (34a); 915 (13); 1024, 1080 (28); 1083 (13); s.n. Franceschi,F., 2 (30).
(la, 20a, 42). Franco,M. & P. Adlard,s.n. (13).
Eilers, L. J., 866 (24). Frankis, M. P., 101 (43al); 114 (43a2); 115 (43al); 117
Ekman, E. L., 105 (8); 1533 (7); 2886 (8); 3126 (7); 3665 (37); 123, 137 (43a2); 142 (14al); 155 (44); 159(14al);
(8); 3829, 4815, 5058, 6379, 6805, 7026 (7); 7302, 7646 179 (43a2).
(8); 10137 (7); 10440 (5a); 10839 (2); 12279 (5a); 12385, Franklin.M. A., 5843 (29).
12801 (2); 12802 (5a); 13036 (8); 15980 (7); 42886 Franklin,M. A. & J. Chandler,5673 (3); 5682 (47).
(8). Freytag,G. F. & J. Baxter,44 (43al).
Elizondo, E., s.n. (14al, 14b, 41, 42). Fryxell, P., 1341 (43al); 1334a (45).
Ellis, 30 (la). Fremont,J. C., 367, s.n. (46).
Ellis, Dunn & Torke, 1081 (12). Fr6derstrom,F. & E. Hulten, 260 (la); 575 (28).
Endlich, R., 1272 (19). Fuentes, C.. s.n. (13).
Engard,R. G., L. A. McGill et al.. 155 (37). Fuentes, M., I (19).
Engelmann,G., 2293 (42). Fuentes, 0., 480 (16); 657 (13); 659 (14al); 675 (20a);
Enriquez,7320 (38). 710 (17); 711 (16); 719 (4); 720 (20a).
Ern, H., s.n. (18). Galeotti, H. G., 3 (13); 4 (22a).
Ernst,W. R., 2220 (16); 2221 (26); 2396 (17); 2397 (28). Galicia X., R.. 136 (34b).
Escudero, M., s.n. (42). Galindo, L. D., s.n. (38).
Espinosa, J., 811 (22a); 812 (15a); 813 (28); 814, 867 Gallegos, J. M., s.n. (3, 11, 33, 35).
(20a); 906 (15a); 908 (14b); 694b (28). Galeotti, H. G., s.n. (22a).
Estavillo, W., s.n. (29). Gallian. P. & M. Leake, 861 (13).
Estrada, E., 2 (Ib); 804 (24); 855 (26); 972 (28); 1239 Galvam, A., s.n. (14b).
(43a ). Gamon, J. A., 13 (34a).
Estrada,E. et al., 807 (34a); 809 (14al). Gandera,M., s.n. (19).
Evans, H., E/73/16 (18). Garcia,4 (28); 1973 (19); 2184 (25); 2187 (43al).
ex herb. J. H. Baltour, s.n. (16). Garciade la Cerda, R., 1354 (19).
ex herb. G. Gordon, s.n. (14b, 16). Garcia,A., 95 (44); 239, 891 (14b); 1162 (43a2).
ex herb. A. B. Lambert,XXVI 55 (15a). Garcia,A. & F. Martinez,2137 (20a).
ex herb.J. Lindley,3150 (20a); 3154 (37); 3164 (13); 3178 Garcia, B., 3, 7 (14b).
(14al); 3179 (16); (Deppe s.n.) (22a). Garcia,C., 102, s.n. (20a).
ex Herb. J. G. Zuccarini,376, 377 (16). Garcia, D. S., 66 (14al); s.n. (14b).
Fagoaga, C., 115 (16); s.n. (la). Garcia. E., III (14b); s.n. (43a2).
Fairchild,D., 3026 (8). Garcia, F., 62 (13).
Farjon, A., 244 (33); 260 (29); 261 (47); 263 (35); 274 Garcia, H., 3 (22a); 5 (la); 6 (43c).
(46); 275 (47); s.n. (31). Garcia, M., 137 (14al).
Farjon, A. & D. Mejia. 295 (14al); 297 (18); 300 (13); Garcia, S.. 35, 276 (la).
301 (20a); 302, 303, 304, 305 (14al); 306 (13); 307 Gardner,M. F. & S. G. Knees, 5122 (42); 53(X)(13); 5339
(14b); 308, 309 (16); 310 (22b); 311 (14b); 312 (38); (34b); 5377 (38); 5259A (16).
313 (20b); 314 (16); 315 (la); 316 (22b); 317, 321 (13); GarzaOcanas, F., s.. (lOb, 12).
326 (34b); 327, 328 (43al); 329 (14al); 330 (32); 331 Garza,A. S., 403 (la).
(22a); 332 (15a); 333, 334, 335 (41); 337 (42); 338 (40); Gentle, P. H., 1130, 1575, 3109, 3690. 8319, 8417, 9433
340, 341 (23); 342 (20a); 344, 345 (17); 349 (la); 350 (5c).
(17); 351 (4); 352 (17); 353, 354, 355 (39); 356 (5c). Gentry,A. & M. Lavin, 50962 (7).
Fassett, N. C., 27688 (28); 28798 (20a). Gentry,H. S., 1998 (lb); 2873 (19); 4394 (43b); 5881 (37);
Faull, J. H., 12944 (24). 5901 (20a); 5902 (19); 5903 (20a); 5904 (12); 6401 (37);
Favela L., S., 3 (44). 6402 (19); 6591 (20a); 7242 (10a); 8565 (43al); 11948
Feddema,C., 174 (20a); 664 (16). (37); 16753 (29); 5901a (20a).
Fernandez,Vales & Herrera,29156 (5a). Gentry,H. S. & F. Cech. 8978 (46).
Fernon, B. E., 159 (8). Gentry, H. S. & R. G. Engard,23110 (42); 23112 (43al);
Ferris, R. S., 5983 (16). 23113 (12).
Figueroa,A. M.. s.n. (16, 18). Gentry,H. S. & C. L. Gilly. 10536 (37); 10790 (16).
Fishbein, M. et al., 779 (37). Gentry,H. S. et al., 20021 (45); 20388 (26); 20388 (28).
Fisher, G. L., 37211 (la); 44235 (25); 44236 (12); 45101 Gereau, R. E., 728 (13).
(28); 45102, 554808 (20a); s.n. (la, 20a, 34a). Gereau,R. E. & G. J. Martin, 1927 (14b); 1932 (la); 2000
Fisher-Meerow,L. L., 687 (8). (16).
Flameca, R., s.n. (20a). Gereau, R. E. & A. J. Moyer, 2144 (14al); 2145 (22b);
Fley, s.n. (20a). 2181 (16); 2183 (14b).
Flores, A., 335 (27); 398, 1200 (28). Gerez, P., s.n. (43al).
Flores, R. O., s.n. (41). Ghiesbreght,A. B., 54 (42).
Flores-Franco,G. & R. Ramirez,2245 (la); 2572 (17). Gibson, A. C. & L. C. Gibson, 2396 (26).
Fogg, G. G. & M. T. Hall, 9, 10, 29, 31 (43al). Gifford, J. C. et al., s.n. (6).
Fonseca, R., 32 (4); 193 (15a); 202 (20a); 275, 276, 480 Gilbert, L. E., 43 (14b).
(27): 527 (18). Gilerz. L., 866 (20a).

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272 Flora Neotropica

Gillis, W. T., 11801 (5b). Gregory.P. & G. Eiten, 112 (20a).

Gillis, W. T. & G. R. Proctor, 12301, 34021 (5b). Grijalva,A., 5097, 5098 (24).
Gilly, C. L. et al., 6 (43al). Grijalva,A. & M. Araquistain,668 (20a).
Gilmartin,A. J., 1515 (43b). Grijalva. A. & E. Sediles, 5297, 5298, 5299, 5300 (24);
Gimate, J., 501 (22a). 5302, 5303, 5325, 5326 (24); 5329, 5330 (24).
Glassman, S. F., 1757 (24); 1843 (18). Guerrero,G., s.n. (13).
Godina, A. G., s.n. (40). Guevara,M., 13 (20a).
Goldman, E. A., 1 (28); 5 (la); 11 (15a); 16 (22a); 19 Guizar,E., 87 (la); 1906 (43al).
(20a); 20 (14al); 163 (12); 268 (20a); 290 (12); 341, Guizar, E. et al., BC8 (47).
798 (20a); 845 (16); 846 (20a); 903, 950 (15a); 960, 987 Gutierrez,C., 52 (14b).
(28); 988 (34a); 996 (20a); 997 (18); 1044 (15a); 1055 Gutierrez,E. A., 8, 9 (18).
(20a); 1132 (47); 1219 (35); 1220 (3); 1245 (11); 1246 Gutierrez.M. & J. Olayo, 837 (19); 855 (4).
(47); 1408 (43al). Gutierrez,M. G., 53 (4); 56 (22a); 513 (la).
Golfari, L., 77571 (5b). Gutierrez,P., s.n. (20a).
G6mez, A. & L. Nevling, 1300 (22a). Gutierrez,R. I., s.n. (38).
G6mez P., A., 69 (28); 610(12); 611 (37); 612 (lOb); 614 GuzmainH., C., H46 (43b).
(la); 643, 646 (12); 648 p.p. (la); 648 p.p. (19); 1420 Guzmin M., R., 497 (19).
(28); 4884 (14b); 5312 (28); 5313 (13). Guzmin. C., 61 (la); s.n. (20a).
G6mez, V. & R. Ruiz, 2 (14al). Guzman,G.. GH1509, 10591 (20a); s.n. (lOb, 16, 19).
Gonzalez, Castellanos & Zanalet, 9835 (14b). Guzmain,H. G., s.n. (13).
Gonzilez, B., 4 (32). Guzman, M. & D. Castro, 1646 (24).
Gonzalez, F. & F. Guevara, 10183 (41). Guzmain,M., A. Montiel & D. Castro, 1261 (24).
Gonzalez, F et al., 8490, 8658 (41). Guzman, R., 715 (47); s.n. (15a).
Gonzailez,G., 77 (43a2). Guzmain,R., I. Puga & A. Valenzuala,6559 (25).
Gonzilez. G. & S. Carvajal,27 (19). Hagen von, C. & V. W. F. H. von Hagen, 1218 (18).
Gonzailez,J. & J. Casteneda,s.n. (18). Hagen von, V. W. F. H., 1065 (5c).
Gonzalez, J. A., 248 (27). Hahn, W., s.n. (14b, 38).
Gonzalez, J. 0., 175 (34a); 307 (18); 308 (19). Haines, A. L. & G. Hale, s.n. (30).
Gonzilez, L. M. & S. Martinez,638 (20a); 639 (16); 768 Hallberg, B., CHIQI, YAHI, YOTI (38); 748A, 901 (13);
(la); 769 (39); 770 (4). 932a (14b); 932b (la); 932c (14b); 1017 (16); s.n. (18,
Gonzalez, L. M., 160 (20a); 504 (16); 531 (14al). 38).
Gonzilez, Q. L., 310, 363, 507 (22a); 508 (Ia); 1608 (20a); Haller,J. R., 3413 (35); 3427 (3); 3431 (47).
1976 (28); 1977 (13); 2070 (43al); 2099 (13); 2146, Hamilton,L. H. & C. M. Rowell, 3227 (28); 3229 (34a).
2435 (14b); 2536 (43al); 2540 (14b); 2545 (42); 2561 Hamilton,S. H., 138 (7).
(13); 2561 (28); 3821 (41); 3837, 28219 (43al). Hammerley,B. J., 383 (43b).
Gonzalez, R. T, 610 (37). Hanon, G., 6055 (47).
Gonzilez, S. & M. Gonzalez, 1708 (19). Hansen, B. F & M. Nee. 7664 (28); 7688 (14al); 7728
Gonzailez,X. N., 99 (la); 99-a (20b). (13).
Gonzalez Flores, 101, 102 (14b). Hansen, B. F. et al., 1582 (20a); 3788 (41); 3856 (28).
Gonzalez G., M., 77 (43a2); 334 (43al). Harbison,C. F., s.n. (47).
Gonzalez M., F., 1464 (41); 2375 (22a); 3058 (14al); Harmon,W. E., 2258 (20a).
17002 (15a); s.n. (22a). Harmon, W. E. & J. A. Fuentes, 2141 (20a); 4828 (13);
Gonzalez M., F. & F. Chiang, 3497 (41). 5848 (5c).
Gonzalez M., F. & C. Delgadillo, 1755 (44). Harmon,W. E. & J. D. Dwyer, 3702 (20a).
Gonzdlez M., F. & P. Hiriart, 12783 (43al). Harmon,W. E. & S. E. Fuentes, 6457 (20a).
Gonzalez M., F. & Protomast,9104 (41); 9213 (43al). Harper,C. C., 45 (43al); 46 (lb); 50 (12).
Gonzilez M., F. & M. Rosas, 1595 (14al). Harris,J. H., 20 (Ia).
Gonzalez M., F. & Sanchez, 2375 (28); 2390 (41). Hartman,C. V., 367, 475 (43a2); 538 (la); 541, 1004 (19);
Gonzalez M., F. et al., 3636 (22a); 8662 (41); 8730 (14b); s.n. (37).
9497 (42). Hartweg,C. T., 31(?) (15a); 92 (18); 440 (43al); 441, 442
Gonzalez O., J., 230 (19); 265 (20a). (28); 443 (22a); 620 (18); 723 (15a); s.n. (13, 14al, 14b,
Goodding, L. N.. 2108, 2109 (12); 2114 (37); 2116 (la); 15a, 15b, 16, 22a, 29).
s.n. (lb). Hastings, J. R. & R. M. Turner,66-57 (47).
Goodman, G. J., 3509 (34a). Hatch, E. D. & C. L. Wilson, 242, 326 (20a).
Graber.J., 4 (19); 113 (43al): 137 (28); 158 (22a); 162 Havrard,V., s.n. (43al).
(28); 190 (42); 260 (43al); 261 (28). Hawkes, J. et al., 1274 (12); 1481 (lb); 1650 (22a); 1651
Graber,R., 174 (15a). (14b); 1707 (26); 2047 (18); 2073 (5c); 2126 (20a).
Gray, C. E. & Rowell, 3000 (20a). Hawkins, T., 69 (34a); 98 (18); 99, 323 (24); 324 (34a);
Greaves, A., 298, 300 (5c); 484 (20a); (serial numbers] 10- 325, 461 (18); 462 (34a); 630 (24); 631 (18).
28, 29-54, 55-75, 76-102, 103-133 (20a); 190-217. Hawkins, T. & B. Allan, 324 (34a).
219-228, 240-250. 281-294 (20a): s.m.(5c). Hayden, F. V., s.n. (12).
Greenman,J. M. & M. T. Greenman,5905 (20a). Hazlett, D., 5, 6 (20a); 86, 109 (18); 546 (20a); 738 (5c);
Gregg, J., 52, 294, 321 (43al); 402 (32); 628 (28); 821 2263 (34a); 2277 (14al).
(la); 975, 628a (16); 689c (22a). Heller, A. A., s.n. (3).

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List of Exsiccatae 273

Heller, A. A. & Barber, 16 (13): s.n. (34a). Hoffman.J. M., 296 (6).
Helmecke, F., s.n. (7). Holdridge, L. R.. 1330 (8); 2353 (34a); 2354 (13); 2357
Hendricks,B. A., 489 (43a1); 513 (lOb); 713 (19). (16); 2358 (20a); 2365 (14al).
Hendrickson,J., 6141, 12146 (42); 14482(30); 14961 (45); Holmes, W. C., 4446(18).
19134 (41). Home. W. J., 1197 (5a).
Hermannvon, H. A., 2653, 3287 (2). Hort. soc. ?, 35 (14b).
Hernandez& Cedillo, 1142 (13). Howard,R. A., 5929 (7); 12251 (8).
Hernindez & Garcia, s.n. (41, 42. 43al). Howard,R. A. & E. S. Howard,8201, 8577 (8).
Hernandezet al., 981 (34a). Howard,R. A. et al., 675 (20a).
Hernandez,A., 201 (13); 201 (15a). Howell, J. T., 8183, 8184, 8268, 8269 (30).
Hernandez, D. 0. & L. Cornejo, s.n. (Sc, 18, 20a, 24, Huerta,J., 722 (28).
34a). Huerta,L. M., 14 (22a); s.n. (34b).
Hernandez,E. & P. Senegalen, 319 (34a); 322 (13). Huey, L. M., s.n. (47).
Hernindez, L., 14 (ISa); 229 (20a). Hughes. C. E., 2 (24); 3 (20a); 4 (34a); 8 (5c); 15 (20a);
Hernandez,N., s.n. (4). 18 (14al); 20 (34a); 21, 22 (13); 39, 43 (24); 44 (20a);
Hernindez, R. & Cedillo, 762, 1145 (28). 45 (24); 48 (24); 88 (34a); 89 (13); 165, 191 (5c); 203
Hernandez,R. & R. Rodriguez,5038 (43al); 5086 (28). (24); 224 (34a); 302 (24); 381 (13); 384 (20b); 387 (38);
Hernandez,R. & R. C. Trigos, 763a (22a); 1341 (13); 1142 470 (24); 480. 481 (8); 548 (34a); 619 (18); 620 (20a);
(13); 1177 (22a). 693, 694 (32); 966, 967 (43b); 970 (4); 971. 973 (19);
Hernindez E. et al., 1, s.n. (22a). 975 (37); 976 (25); 977 (10b); 978 (la); 980 (25); 981
HernandezM., R., 453 (41);.2072 (46); 3251 (37); 3343 (43al); 983, 984 (40); 1000 (27); 1261 (20a); 1262
(28); 3428 (34a); 3776 (42); 3863 (15a); 4938 (28); 7258 (14al); 1263 (24); 1264(18); 1265 (24); 1288(38); 1322
(34a). (14b); 1348 (20a); 1349, 1357 (16); 1425 (24); 1428,
HernandezM., R. & 1. Hernandez,6536 (43al). 1699 (18); 1751 (20a); 1773 (16); 1798 (14al); 1852
HernaindezM., R. & H. Susan, 5227 (27). (42); 1853 (43al); s.n. (34a).
Hernindez M., R. et al., 5176 (24); 5177 (26); 5195 (18); Hughes, C. E. & B. T. Styles, 108 (20a); 136 (18); 137
5198 (20a); 5200 (28); 5225 (20a); 5228 (24); 5230 (18); (4): 138 (25); 140 (18); 168, 169 (47); 170 (11); 172,
5231 (27); 5239 (20a); 5249 (22a); 5251 (20a); 5257 173 (35); 174 (3); 175 (11); 178 (29); 180 (47); 181
(27). (11); 183 (33); 184, 185 (46); 139a (25).
Hernindez X., E. & A. J. Sharp. 1278, 46237 (38). Humbert,H., 27625. 27699 (8).
HerreraC., I., s.n. (13, 32, 37). Hummel,C., 126. s.n. (5c).
Hess, W. & L. Byrne, 4731A (43al). Humphreys.C.. 5 (la).
Hess, W. & M. T. Hall, 534 (12); 546 (la); 554 (lOb); 597 Hunt, D. R., 1. 2 (24); 62. 62a, 62c (5c); 480, 481. 617
(15a); 597 (16); 591A (la); 598A, 598B (15a). (24); 9330 (43al); 2366154, 2366155 (24).
Heyde, E. T. & E. Lux, 3156 (34a). Huston, S. S., 6553 (5c).
Higgins, E. B., s.n. (47). Ibarra,G. M., 139 (15a); 175 (la); 176 (16); 912 (la);
Higgins, L. C., 12885 (37). 1150 (20a); 1163 (la); 1166 (16); 1194, 2362 (20a).
Higman, S. et al., 1 (34a); 2 (14al); 4 (13); 5, 6, 7 (14al); Iltis, H. H. & R. Guzmin, 3255 (19).
8(24); 9(14al); 10, 11(16); 12 (20a); 13(18); 14(14b); lltis, H. H. & R. Koeppen. 221 (22a); 306E. 368A (la);
15, 16 (14al); 17 (24); 18 (34a); 19 (14al); 20 (24); 21 541A (20a); 853A (la): 978, 979, 1031 (13); 1287A(28);
(20a); 22 (18); 23 (16); 24 (38); 25 (24); 26 (14b); 27 3069 (16).
(14al); 28 (14b); 29, 30, 31 (14al); 32 (20a); 33 (16); Jack, J. G., 6874 (2).
34, 35 (28); 36 (14b); 37, 38 (la); 39 (28); 40 (22b); 41 Jaeger. B.. 155 (8).
(la); 42 (22a); 43, 44 (17); 45 (34a); 46, 47 (28); 48 Janzen, D. & Ives, 1213 (5c).
(14b); 49 (28); 50 (la); 51 (17); 52 (18); 53 (20a); 54 Jasso. J., 1 ( 13).
(16); 55, 56 (27); 57 (15a); 58 (27); 59 (14b); 60 (la); Jeffrey,J., 731 (11).
61 (14al); 62 (16); 63 (14al); 64 (34a); 65 (18); 66 Jemmow,242 (20a).
(14b). Jennings,0., 82 (5a).
Hill, R. J. & K. Saltus, 2515 (47). Jimenez, C. R., s.n. (22a).
Hill, S. R., 1672 (18); 1673 (15a); 1767 (38); 1799 (28); Jim6nez,J. J.. 235 (13).
1805 (14al); 1815 (24); 1824 (16); 1825 (20a); 1846 Johannessen,C. L., 470 (5c); 590, 633. 897. 941 (20a);
(13); 3005 (5b); 10974 (6). 942 (18); 969 (5c).
Hinds. H. V. & E. Larsen,65, s.n. (25). Johansen,R., s.n. (14al, 15a).
Hinton, G. B., 3720 (la); 3721 (16); 3722 (14al); 3723 Johansen,R. & Mojica. s.n. (28).
(la); 3724 (20a); 5773 (14al); 5774 (15a); 5775 (18); Johnston,I. M. & C. H. Mueller,523 (45).
5776 (14al); 5777, 5784 (la); 5900 (28); 5929 (27); Johnston,1. M.. 8735 (42); 8935 (12); 8998 (37).
6222 (15a); 7483 (20a); 7486, 7487 (la); 8802 (20a); Johnston,M. C.. 8735 (42).
10357 (18); 10358 (20a); 17275 (44); 17768 (43al); Johnston,M. C. et al., 11575 (lOa); 12135 (42).
18753 (28); s.n. (16, 26). Jones, G.. 1040, 1053 (43al).
Hinton,G. B. et al., 5775 (14al); 8802 (34a); 8989 (14a1); Jones, M. E.. 492 (19); 27080 (43b); s.n. (la, 43a2).
17920 (44). Judd, W. S.. 1166. 1487 (8).
Hinze, V. L. & C. M. Rowell, 2573 (20a). Judd,W. S. & B. Judd, 2964 (8).
Hjerting& Odum, s.n. (12, 25). Judd, W. S. et al., 3022 (8).
Hoard, L. R & H. von Schrenk,s.,. (10a). Judziewicz & Cochrane,4996 (4).

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274 Flora Neotropica

Kaplan, L., 172 (38); 356 (16); 357 (15a); 380 (14al); Lanner.R. M., 2016, 2025 (47).
31ch57 (16); 32ch57 (20a); 59ch57 (18); 61ch57 (24); Lape F.. s.n. (37).
62ch57 (13); 63ch57 (15a); 128ch57 (16); 143ch57 Larsen,E., s.n. (25).
(20a); 148ch57 (24); 150ch57, 165ch57 (16); 172ch57 Lascurain,3 (43c).
(34a); 173ch57 (20a); 220ch56 (16). Lathrop,L. M., 5865 (24); 7488 (20a).
Kaplan, L. & E. L. Davis, 183 (13); 368, 369, 370 (24). Laubenfels,D. J. de, 663, 665 (14al).
Karwinski.F.. 519 (41); s.n. (13, 43al). Laughlin, R. L., 976 (24); 1200 (20a); 2917 (24).
Keil, Meyer, Lewis & Pinkava,5994 (45). Laughlin, R. M., 417 (14b); 562 (15a); 587 (16); 1196
Keller, C. S., 707 (47). (34a); 2911 (18); 2912 (16); 2975 (20a).
Kellerman, W. A., 4516, 4520, 4521, 4522, 4523 (20a); Leavenworth,W. C., 272 (la); 544 (14al); 600 (16); 601
4524 (5c); 4580 (13); 5026 (18); 5840, 6072, 6073 (13); (la); 706 (13); 707 (la); 719 (16); 886 (43al).
6081 (24); 6082 (18); s.n. (13). Leavenworth,W. C. & H. Hoogstraal, 1168 (13).
Kemp, R. H., 3 (20a); 5 (24); 12 (18); 16, 17 (5c); 20, 21 Ledoux, D., 1967 (Ib).
(5c); 36 (18); 40, 41, 42, 43, 44, 45, 47, 48, 55. 57 (5c); Lehmann,F. C., 1588, 1677 (34a).
58 (24); 61, 64, 70, 76 (20a); 87, 92, (20a); 95, 96 (5c); Lehto. E., 24344 (4).
99, 100, 101, 103, 104, 105, 106 (5c); K104 (18); 110, Lehto, E. et al., L21714 (43al).
114, 115(Sc); 116 (20a); 117(5c); 118, 119, 120, 121, Lemmon,J. G., 239 (la); 241 (14al); 242 (20a); 243 (la);
122 (20a); K119, K122 (18); 130, 131, 132 (26); K132 244 (16); 259 (13); s.n. (la, 12. 13, 16, 18, 28, 31, 37.
(18); 134, 135, 136, 137 (20a); 138 (14al); 140, (20a); 47).
K149 (18); 1/69 (20a); Y, Z (5c). Lemus, R., 49 (37); 400 (la).
Kenoyer, L. A., 1041 (la); s.n. (43al). Lemus, R. & G. Guzman, s.n. (la).
Kenoyer, L. A. & E. K. Crum, 2735 (43al); 3177 (42). Lemus, R. & R. Torres, 181 (37).
Killip, E. P., 43267 (6); 44027, 44135 (Sa); 44137, 44562 Lent, R. W., 611 (20a).
(2). Le6n, H., 4250, 4622, 17056 (Sa).
Killip, E. P. & F. Swetland, 41611 (5a). Le6n, H. & Charles,4902, 4935 (5a).
Kilterman& Raveret, 29 (22a). Le6n, H. & M. Roca. 10326 (7).
Kimnach, M. & Brandt,861 (Ib); 866, 1039 (43al). Leonard,E. C.. 3728, 3767, 7523 (8).
Kimnach, M. & H. Sanchez, 1812 (34a). Leonard,E. C. & G. M. Leonard, 14979 (8).
Kimnach, M. & Sanchez-Mejorada,1890 (19). Leopold, A. S., 23 (43al); 24 (Ib); 25 (28); 26 (37); 150
King, R. M. & T. R. Soderstrom,4794 (18); 4891, 4895 (la); 227 (12).
(20a); 5116 (26): 5143 (15a); 5144 (la); 5168 (14al); LeSueur.H., 207 (45); 550 (la); 551 (12); 557, 560 (19);
5170 (15a). 1274 (12); 1276 (Ib); 551a, 551b (la); s.n. (Ib).
Kirkby.V. T., 2695 (22a); 2778 (28). Leticia G., M. R., 184 (la).
Kiru, F.D., s... (8). Leuenherger,B. & C. Schiers, 2679 (18).
Klaus, W., 276, 277 (39); s.n. (12, 14b, 16, 34a. 37, 39, Levin, G. A., 1507 (Ib).
41. 42, 43a2). Lewis, W. H.. 7160 (5b).
Klaus, W. & J. Andrade,s.n. (40). Liebmann,2 (16); 3 (22a); 4 (20a); 9 (22a); 10 (20a); 1/7
Kluge, H. C., s.v. (5c). (28); s.n. (la, 34a).
Knobloch, D., 5637 (Ib). Liebmann,F. M., 14777 (34a); 14779 (20a); 14789, 14790
Knobloch, 1. W., 76 (37); 82 (19); 131 (Ib); 338 (43al); (34a); 14791 (la); 14792, 14794 (24); 14795, 14796
340, 410 (Ib); 1164 (10a); 1290 (Ia); 1321 (Ib); 1843 (20a); 14803 (22a); 14804 (4); 14805 (22a); 14810.
(19); 5025 (37). 14815, 14818 (28); s.r. (22a).
Knobloch, J., 5028 (la); 5637 (Ib); 5700 (16). Lind. W. L. Mittak& J. P. Perry,8445 (Sc).
Knutsen, N., x.n. (18). Linden, J. J.. I (22a); 2 (34a); 4 (28).
Koch, S. D., 7570 (13). Lindsay,G., 1947 (43b); s.n. (3, 30).
Koch, S. D. et al., 79253 (20a); 79331 (38). Liogier, H. (vide Alain, Brother). 18593. 21104 (8).
Kral, R., 27614 (19); 69204 (20a). Lipp, F. J., 72 (20a).
Kruger& Gillespie, 28 (28). Litchtey,P.. s.. (46).
Kruse, H., 1778 (16); 1810, 2406, 2408, 2409 (18). Little, E. L., 7845 (4); 11094 (28); 14046 (2); 14047 (5a);
Kuijt, J. & A. Miller, 1018 (30). 14048 (2); 14051 (5a); 14052 (2); 17831 (15a); 17832
Laguna,A. R., s.n. (la). (la); 17833 (14al); 17834 (25); 17835 (la); 17836
Lamas et al., 72 (4). (14al); 17837(16); 17838(18); 17839(26); 17840 (la);
Lamas G., A., s.n. (Ib). 17841 (16); 17842 (26); 17843 (20a); 17844 (16); 17845
Lamas, R., 12 (17); 26 (20a); 27 (17); 31 (16); 42 (18); 45 (4); 17846 (15a); 17847 (14al); 17849, 17850 (20a):
(40); 47 (16); 49 (37); 65 (20a); 66 (19); 67 (20a); 181 17851 (17); 17852 (16); 17853 (20a); 17854 (17); 17857
(37); 400 (la). (43al); 17858 (Ia); 17859(12); 17860(37); 17861 (19);
Lamas, R. & R. Torres,25 (26); 28 (18); 29 (17); 56 (23); 17862, 17863 (43al); 17864 (25); 17865 (37); 17867
60 (18); 178 (23); 235 (20a). (25); 17868 (lOb); 17870 (28); 17874 (12); 17875 (28);
Lamas, R. et al., 50 (20a). 17876, 17877 (IOb); 17878(19); 17880(25); 17881 (la);
Lamb, A. F. A., 10 (8); 11 (5c); 31, 32, 33. 34 (5c); s.n. 17882 (37); 17883, 17885, 17886, 17887 (19); 17890,
(2, 5a. 7, 8). 17891, 17892, 17893 (28); 17897 (la); 17899 (10a);
Landa, D. S., 29 (25). 17901 (25); 17903 (Ib); 17908, 17909, 17910 (37);
Langman. 1., 3722 (20a); 3722 (24). 17911 (12); 17912(b): 17913 (28); 17914 (10a); 17915

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List of Exsiccatae 275

(37); 17919 (10b); 17920(37); 17921 (lOb); 17922(19); 11608(5a); 11775, FH11778, 11779(20a); 11924. 11926,
17923 (la); 17924 (12); 17925 (Ia); 17936 (10a); 17938 11927, 11928, 11929, 11930, 11931, 11932 (5b); 11934,
(43al); 17940 (26); 17942 (14b); 17943 (34a); 17944 11935. 11936, 11937. 11938. 11939, 11940, 11941,
(13); 17946 (24); 17949 (14al); 17950(24); 17951 (13); 11942, 11943, 11944, 11945, 11946, 11947, 11948.
17952 (38); 17952 (16); 17953 (18); 17954 (34a); 17955 11949, 11950. 11952, 11953, 11956 (5c); 12085 (5a):
(13); 17956 (la); 17957 (24); 17958 (34a); 17961 (22a); 12086, 12087 (5b); s.n. (24).
17962 (15a); 17963 (14b); 17964 (15a); 17965 (38); Lugo, A.. 24 (43al).
17966 (18); 17969 (20a); 17971 (16); 17972 (18); 17973 Lumholtz.C., 3, s. . (19).
(20a); 17974 (16); 17976 (20a); 17978 (28); 17979 (41); Luna, D., 41, 1725 (2)a); s.n. (41. 42).
17980 (5c); 17981, 17982. 17983 (24); 17984, 17985, Luna Ledesma, V., 11 (40).
17986, 17987, 17988 (Sc); 18581 (46); 18901 (43ai); Lundell,C. L., 677. 678. 2846, 16764 (5c).
18902 (Ib); 18903, 18904 (43al); 18905 (12); 18906 Luteyn,J. L. & M. Luteyn-Lebr6n.11662 (13).
(Ib); 18907, 18908 (43al); 18910A(37); 18915, 18916 Lyonnet.E.. 2071 (34a).
(43al); 18917 (Ib); 18918 (12); 18919 (43a2); 18922 L6pez-G6mez. R.. 285 (14b); 301 (14al).
(lb); 18924, 18925 (37); 18927 (19); 18928 (37); 18929 L6pez-Sinchez, 297 (34a); 352 (14b); 354 (18).
(lb); 18930, 18931 (43al); 18932 (lb); 18933 (43al); Maas, P. J. M. et al.. 7750, 7751 (2).
18934 (12); 18935 (28); 18935A (43al); 18936 (19); MacDougal, F., s.n. (38).
18937 (10b); 18938, 18938A, 18939 (12); 18940 (la); MacDougall,T B., 261A (la).
18941 (lb); 18942 (10b); 18943 (25); 18944(37); 18945 Machuca. 933 (20a); 1282 (14al); 1287 (16); 6127 (Ib):
(lOb); 18946 (25); 18948. 18949 (12); 18950 (43al); 6128 (20a); 6129 (16): 6132 (19).
18952, 18953 (14al); 18956 (28); 18957 (15a); 18958 Madrano,11, 270. 271. 272, 273 (42).
(14al); 18960 (15a); 18961 (13); 18962 (34a); 18963, Madrigal S.. X., 24 (27); 91, 92 (14al); 251 (20a); 283
18964 (15a); 18965 (13); 18966 (14b); 18967 (14al); (15a); 303 (14a1); 325 (41); 653. 654. 655 (38); 1460
18968 (28); 18969 (26); 18970. 18971 (13); 18972 (26); (13); 1461 (38); 1464 (la); 1465 p.p. (17); 1465 p.p.
18974 (13); 18975, 18975A (28); 18976 (22a); 18977 (26); 1467 (17); 1471 (11); 1472 (47); 1473 (33); 1474,
(43al); 18979(14b); 18980(15a); 18980, 18981 (14al); 1475 (30); 1572 (13); 1634-1, 1634-2. 1634-3. 1634-5.
18982 (43al); 18984 (37); 18987 (44): 18988 (13); 1634-7 (14al); 2002 p.p. (la); 2002 p.p.. 2020 (19):
18989 (37); 18990 (43al): 23010, 23011 (43a2); 25002 2021 (20a); 2053. 2075 (16); 2076 (la); 2077 (17); 2078
(5c); s.n. (13). (26); 2079 (16); 2080 (la); 2081 (16); 2083. 2084 (17):
Little, E. L. & D. S. Correll. 18991 (45). 2085 (26): 2086 (la); 2087 (26); 2177, 2202, 2218 (39);
Little, E. L. & K. W. Dorman. 14033 (6). 2240, 2241 (20a); 2243, 2244. 2245 (17); 2246 (20a);
Littlepage,s.n. (30, 47). 2247 (17); 2248, 2249 (20a); 2260. 2290 (17); 2294.
Littmann,E. & T. D. Pennington,9043 (43al); 9052 (la); 2315 (19); 2341 (20a); 2527 (15a); 2548 (Ib); 2549 (12);
9053 (26); 9055 (14al). 2561 (37); 2638 (27): 2651 (17); 2757 (4); 2761 (20a);
Lloyd, F. E., 30 (42); 64 (43a2); 225 (42). 2762 (17); 2781 (16); 2800 (26); 2806 (20a); 2807 (17);
Lock, E. E. M., 113a (22b); 10280 (5c); 10572 (24); 2825. 2831 (20a); 2840 (la); 2844 (26); 2867 (17); 2890
10618 (14al); 11443 (Sc); s.n. (20a, 24. 27, 32). (13); 2930. 2931, 2932, 2933, 2934 (17); 3117 (14al);
L6pez et al., 144 (34a); 248 (14al); 651 (20a). 3139 (27); 3197 (14al); 3198 (39); 3211 (14al); 3212
L6pez, E. S., 19 (20a). (34b); 3213 (la); 3214 (15a); 3215, 3219 (16); 3220
L6pez, G. G., 249 (22a). (la); 3221 (4); 3222 (17); 3223 (20a); 3226 (14al); 3235
Lopez, I. R., 675 (28). (39); 3245 (20a); 3246 (26): 3255 (15a); 3256 (27); 3257
L6pez, M.. 107, 1239 (7); 2111 (8). (26); 3259 (la); 3267 (15a); 3269 (13); 3283 (18); 3324
L6pez, R., 284 (14al); 298 (27); s.n. (16). (14al); 3326 (20a); 3346 (28); 3423 (26); 3449 (15a):
L6pez. R. & A. Saynes, 659 (18). 3451, 3452 (16); 3460 (27); 3463 (18); 3466 (17); 3471
L6pez, R. et al., 254 (Ia). (14a1); 3474 (ISa); 3476 (26); 3478 (16); 3489 (14al);
L6pez, S., s.n. (14a ). 3490 (28); 3491 (14al); 3492 (27); 3500 (20a); 3501
Lorea, F. G., 1969 (Ia); 2229 (13); 2705 (27); 2878 (18); (18); 3537 (13); 3540, 3541 (16); 3544, 3545 (25); 3546
2879 (20a); 2973 (18); 2975 (4): 2986 (16); 3124 (17); (15a): 3550 (20a); 3552. 3553a (16); 3554 (14a1); 3556.
3138 (38); 3271 (16); 3483 (4). 35536 (15a); s.n. (14al, 14b, 15a, 16, 18. 20a. 22a. 24.
Lorea, F. H., 21()0 (34a). 26, 32. 38. 41, 42).
Lorence, D. H., 3761 (43al); 4073 (22a). MadrigalS.. X. & S. C. Boyas, 3511 (19).
Lorence, D. H. & R. Cedillo, 4069 (22a). Madrigal S.. X. & Bello Gonz;ilez. XMS3541 (20a):
Lorence, D. H. & A. Garcia, 3538 (13); 3597 (22a). MS3543 (17); XMS3547 (15a).
Lorence, D. H. et al., 3637 (22a); 4651 (13); 4691 (14b); MadrigalS.. X. & R. E. Perez. s.n. (14b. 15a. 17. 18. 20a.
4692 (16); 4758 (22a); 4759 (14b). 22a, 27).
Lorence, D. H. & R. Hernindez, 4910 (22a). Madrigal S.. X. & L. Vcla, s.n. (14al, 14b. 17. 27. 38.
Los Angeles Univ. (zool.), 260 (43al). 43al ).
Lot, A., 365, 1073 (22a). MadrigalS.. X. & Vela Galvez. s.n. (14al. 28).
Lott, E. J. & T. Wendt, P-93 (43al); P-133 (14b); P-134 MadrigalS.. X. & J. Villa. s.n. (15a).
(43c). MadrigalS.. X. et al.. 2291 (20b); 2293, 2312 (20a); s.n.
Lozano, G. V., 654 (16); 655 (15a). (13, 28).
Lucera, R., L66 (20a). Magana,J. L., 900, 990 (14b); 1002 (22b); 1030 (la); 1202
LiUckhoff,H. A., 11128, 11220, 11295 (5b); 11444 (5c); (15a); 1315 (14a1): 1324 (la); 1325 (14al).

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276 Flora Neotropica

Magana, M. A. & A. Zamudio,s.n. (20a). 3467 (30); 3468 (33); 3470 (31); 3477 (25); 3480 (20a);
Magana, M. A. et al., 290 (20a). 3486 (34b); 3505 (14a2); 3557 (14al); 3558 (21); 3559
Maj, M.. 2633, 3156 (la). (37); 3564 (24); 3565 (17); 3570 (20b); 3580 (20a); 3582
Mancera 0., A., AM217 (47); AM245 (35); AM249, (18); 3583 (25); 3585 (16); 3586 (lOb); 3588 (14al);
AM250, AM251, AM266 (47); AM268, AM269 (35); 3599 (4); 3601 (20b); 3610 (43a1); 3619 (11); 3624 (24);
M0371, M0374 (18); M0380. M0381 (20a); M0511 3625 (15a); 3628 (14b); 3629 (15a); 3634 (13); 3636
(lb). (12); 3637 (15a); 3640 (16); 3646 (15a); 3648 (20a);
Mancera0., A. et al., s.n. (14al). 3651 (34b); 3654 (38); 3659 (10b); 3661 (25); 3664
Manning, W. E. & M. S. Manning, 53275 (43al); 53328 (14al); 3665 (13); 3671 (16); 4000 (37); 4003 (14al);
(14b); 53597 (28). 4(009(28); 4029 (13); 4041 (32): 4045. 4046,4049. 4050
Manos, A. S., .s.n. (34a). (17); 4067 (25); 4078 (12); 4085, 4095, 4096, 4097 (21);
ManzaneroM., G. 1., 508 (15a). 4101 (42); 4102, 4104, 4105 (14b); 4108 (20b); 4110
Manzera,Viveros Z. & Takaki.s.n. (14al). (10b); 4125. 4125B (37); 4126 (43al); 5411, 5412 (17);
Marcks, B. & C. Marcks,s.n. (34a). 5413 (18); s.. (14b, 16, 19, 20a, 24, 28. 38, 42).
Marie-Victorin,Frere, 60010 (7). Martinez,M. & Trujillo, MATM13 (la).
Marie-Victorin,Frere& FrereClement, 21428 (7). Martinez.M. A. & F. Sim6n, 16 (12).
Marin, R., M65-23 (47). Martinez,S.. 495 (43al); 653 (la); 1092 (16); 1184 (20a).
Marineros,37 (24). Marlinez,T. V., s.n. (20a).
Marquez,W., 479 (22a); 692, 714, 721 (38). Mason, H. L., 2030 (30); 4096 (29).
Marsden.H. W., s.n. (30). Matuda, E.. 292 (24): 293 (34a); 870 (28); 1348 (22a);
Marsh. E. G.. 821 (43a2); 821 (37); 1358 (43al). 1530M (16); 2361, 2366 (34a); 2368, 2853, 2874 (13);
Marshall.J. T., 44 (Ib); 45 (12); 47 (37); 49 (10a); 52 (37); 2919 (34a); 4116, 4123, 4321, 4595 (20a); 4641 (13);
54 (10a); 72 (Ib); 73 (12); 104 (43a2); 114. 121 (25); 4657 (34a); 4658 (16); 4792 (20a); 5072, 5231 (24);
122 (10a); 123(12); 124, 126 (lb); 134 (10a); 141 (37); 5317 (18); 5524, 5530 (24); 5534 (16); 5549 (24); 5557
142 (43a2); 148 (25); 83-26 (15a); 83-27 (42); 83-28 (20a); 5561. 5562 (14b); 5583 (18); 5785 (24); 5913.
(lOb); 85-120 (41); 84F-104 (16). 5932 (20a); 5973, 6012 (24); 15524 (24); 15583 (18);
Marshall,S. A. & D. A. Neill, 6559 (Sc). 15771 (16); 18191 (13); 18602, 18848 (la); 18849
Martin.M48 (14b). (14al); 18862 (Ia); 18910, 19184, 19185, 19186 (13);
Martin, G. J., 249 (28); 320 (14b); 377 (34a); 378, 387 19292, 19495 (la); 19497, 19600 (28); 19602 (13);
(24); 503 (28); 508 (14b). 19690 ( a); 19693 (28); 20869 (la); 20871 (14al); 20872
Martin, P. S., H33 (37); 046 (13); H46, H47 (28); H48 (15a); 20873 (18); 20878 (13); 20879 (22a); 20905 (15a);
(15a); 50 (15a); 126 (28); 56021 (37); 56028 (10a); s.n. 20956 (la); 21950, 26213 (15a); 27299 (13); 27303
(14b). (15a); 27416 (la); 27543 (27); 28174 (14al); 28753
Martin,P. S. et al., 280 (Ib). (15a); 29610 (13); 30802 (20a); 30803 (18); 30863 (27);
Martinez,Soto & Ramamoorthy,3896 (27). 30968 (28); 30969 (16); 30971 (15a); 30973 (20a); 30982
Martinez,Tellez & Hernandez, 12514 (20a). (14al); 30985 (la); 31016 (14al); 31021 (20a); 31022,
Martinez,E., 523, 650 (la); 686 (lb). 31023, 31024 (14al); 31028 (20a); 31032, 31033 (17);
Martinez,E. M. & O. Tellez, 12681, 12683 (Sc). 31036 (15a); 31040 (20a); 32143, 32144 (17); 32145
Martinez,E. O. & F. Lozano, 781 (43c). (27); 32147, 32156 (20a); 32157 (27); 32611 (14al);
Martinez,E. O. et al., 773 (14b). 32612 (la); 32667, 32670, 32671 (10a); 32677 (la);
Martinez,F., 300 (13); s.n. (4, 11. 13, 19, 46, 47). 32701 (12); 32723 (10a); 38549 (43a2); 38573 (16); s..
Martinez,G., 223 (22a). (14al, 15a. 20a, 22a, 41).
Martinez,L. P., LP80 (26); LP221 (14al); LP325 (27). Matuda,E. et al., 31020 (18); 31038, 31041 (27).
Martinez,M., I (20a); 12 (14b); 19 (21); MATM-24(13); Maxon, W. R., 3428 (20a); 4341 (7).
67 (24); 88 (15a); 102 (la); 803 (38); 1109 (14b); 1620 Maxon. W. R. & R. Hay, 3141 (5c); 3370, 3375 (20a);
(14al); 2002 (15b); 2145 (24); 2148 (11); 2162, 2164 3428 (24); 3688, 3693 (13).
(19); 2168 (24); 2169 (Ib); 2184, 2185 (41); 2188 (34a); May Nah, A., AM42 (14al); 60 (4); 62 (17); AM63 (26);
2189 (34b); 2212 (15b); 2224 (14al); 2225 (34a); 2225 AM71 (32); AM79 (14al); AM81 (19); AM199, AM200
(15a); 2226 (28); 2229 (37); 2336 (34a); 2340 (37); 2341 (26); AM237 (14al); AM238 (27): AM239 (38); 245.
(34a); 2346 (12); 2358 (14al); 2363 (12); 2365 (15a); 246, 247, 248, 249, 250, 251 (14al); AM256 (38);
2367 (14al); 2375 (42); 2377 (14b); 2378 (34a); 2382 AM257 (20a); AM306 (17); 307, 308 (16); 309. 310,
(38); 2384 (26); 2392, 2393 (18); 2413 (16); 2427 (14b); 311 (la); AM312 (26); 313 (14al); AM314 (17); 318
2429, 2433, 2438, 2444 (16); 2453 (32); 2455 (41); 2460 (34a); AM321 (20a); AM322 (17); AM326, AM327
(lOb); 2487 (4); 2509 (12); 2526 (4); 3407 (43al); 3410 (22a); AM332. AM333 (11); AM335 (33); AM338 (II);
(46); 3411 (43al); 3412 (41); 3413 (37); 3415. 3417 AM341 (3); AM342 (35); AM347, AM348. AM349,
(34a); 3418 (35); 3419 (38); 3420 (la); 3421 (Ib); 3422 AM350 (22a); AM353 (13); AM361 (20a); 655, 712
(19); 3423 (26); 3424, 3425 (28); 3426 (26); 3427 (4); (44); AM1133, AM1135 (26); AM1136 (4); AM1148
3428 (18); 3429 (17); 3430 (14al); 3431 (14a2); 3432, (18); AM1150 (26); 3165, 3166 (17); 3167 (16); 3186
3433, 3434 (14al); 3435 (14b); 3436, 3437 (15a); 3438 (34b); 3278 (17); 3285 (27); 3291 (14al); 3532, 3648.
(15b); 3440 (13); 3441, 3442 (lOb); 3443, 3444, 3445, 3650, 3651 (20a); .Sn. (18. 19. 32, 43a2).
3446, 3447, 3448 (16); 3449, 3450 (12); 3451, 3452 Mayo, E., 261 (15a).
(11); 3453 (iOa); 3454 (25); 3455 (lOc); 3456, 3456a Mayr, H.. s.n. (12).
(27); 3457 (20a); 3458 (20b); 3459 (21); 3460 (20a); Maysilles, J. H., 7259 (4); 7262, 7357. 7366, 7375 (10b);
3461 (24); 3462 (22a); 3463 (32); 3464 (3); 3465, 3466, 7376 (12); 7399 (43al); 7580 (IOa); 7697 (10b); 7712

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List of Exsiccatae 277

(12); 7719 (lb); 7796, 7808 (37); 7924 (12); 7976 (19); 313 (13); 321 (24); 343 (13); 358 (24); 359 (18): 372
7981 (10a); 8030 (lb); 8064 (28); 8079 (10b); 8080 (37); (13); 376 (20a); 377 (5c).
8130 (lb); 8152 (10b); 8153 (37); 8154 (12); 8200 (37); Mejia, M. et al., 608, 612, 8834 (8).
8296 (Ia); 8297 (12); 8298 (37); 8307 (12); 8309, 8310 Mell, C. D., I (la); 13 (20a); 2288 (28); 710850 (16).
(lOb); 8311 (10a); 8312 (lOb); 8313, 8314, 8315, 8316 Mendez G., T., s.n. (15a).
(lb); 8317, 8318 (12); 8335 (10b); 8337, 8341 (10a); Mendez, T. M., s.n. (12).
8342 (Ib); 8380, 8435 (la); 8465 (19); 8490 (37). Messer, E., 228 (28).
McAdams, R. F., 73 (la). Mexican exped. E19 Univ. of Kansas,s.n. (22a).
McCarter,P. S., 23, 63, 94. 135. (24); 113 (34a); 114a, 116 Mexia, Y., 496 (10a); 724, 801 (15a); 1421 (18); 1463
(18); 118 (24); 123 (18); 130 (24); 146 (18); 174, 196, (14al); 1627 (16); 1759 (19); 1900 (20a); 2513. 2514
226, 245, 274, 275, 296, 306, 307, 310, 346 (24); 405, (Ib); 2515 (12); 2516(43al); 2607 (la); 2608 (12); 2609
406, 432, 437, 442, 482, 489, 490, 492, 519, 523, 527, (37); 2620 (Ib); 2727 (la); 9015 (27); 9083 (16); 9088
528, 538, 540, 549, 556 (24); 559, 560, 561 (20a); 563 (4).
(34a); 567 (34a); 568 (13); 570 (24); 571 (20a); 572 Meyer, F. G., 9914 (20a).
(14al); 574 (24). Meyer, F. G. & D. J. Rogers, 2740 (28); 2802 (22a); 3002
McCarter,P. S. & C. E. Hughes, I (5c); 13 (5c); 39 (34a); (37); 3087 (14al); 3002A (37).
41 (13); 43 (34a); 45 (13); 53, 55 (20a); 57 (16); 58 Meyer, W. C.. 87 (5c).
(24); 60 (14al); 61 (34a); 62 (14b); 79 (22a); 85 (43al); Meza, H. M., s.n. (13).
87 (14al); 88 (la); 94 (14b); 97 (22a); 98 (14b); 103 Meza, H. M. & G6mez, 98 (17); 100 (15a); 101 (20a).
(14al); 105 (10c); 223, 225, 226, 272, 273, 323, 324 Midence, J. D., s.n. (34a).
(24). Millan Espin, M. E., 369 (27); 582, 590 (4).
McCarter,P. S. & B. T. Styles, 35, 50, 72, 73 (24); 110 Miller, J. S. & G. Campos. 2844 (4).
(18); 122 (34a); 130 (24); 145, 163 (24); 184, 185 (16); Miller, J. S. & J. K. Meyers, 2761 (24).
195 (24); 198 (14al); 221, 237 (24); 253 (18); 255 (22a); Miller, J. S. et al., 305 (20a); 326 (17); 344 (lb); 7388
266, 267, 271, 272, 273 (22b); 279 (14al); 279a (17); (47).
281 (24); 288 (17); 291 (24); 305 (20b); 313 (22a); 315, Miller, W. et al.. 19 (18).
317, 322 (24); 336 (28); 340, 341, 342 (27); 343 (14b); Millspaugh,C. F. & C. M. Millspaugh,9187, 9214 (5b).
344 (la); 345, 346, 347 (27); 348 (20a); 349 (38); 387 Minoz, C. B. & A. Rebolledo, 17 (22a).
(13); 389 (34a); 394 (18); 398 (18); 410 (18); 412 (34a). Miranda,F., 64 (13); 73 (la); 151 (14al); 196 (ia); 1102
McCarter,P. S. et al., 3, 4, 5, 6, 7, 8, 9, 10 (13). (38); 3164 (22a); 3331 (38); 4643 (15a); 5080 (14al);
McDaniel, S. & W. J. Clark, 14318 (5c). 5770 (20a); 6085 (38); 6534 (18); 7094 (14al); 8885
McDonald & G6mez, 1170 (44). (46); s.n. (38).
McElwee, R. L., 508 (20b); 516, 518, 519, 520, 534, 535 Miranda,F. & F. A. Barkley. 17M134 (13).
(14al); 551, 552. 553 (20a); 517A, 531, 532, 533 (14al). Mirov, N. T., 2, 4, 5. 8, 10, 11 (20a); 1950 (14b); s.n.
McGill, L., D. & D. J. Pinkava,P12325 (35). (14al, 18, 34a, 42).
McPherson,G. D., 712 (20a). Mittak,W. L., 18, 3060, 3110 (13); 3140 (34a); 8147 (38);
McVaugh,R., 10072(13); 10121 (14al); 10135(la); 10347 8148 (24); 8153, 8155 (34a); 8159, 8162 (20a); 8164
(42); 10358 (43al); 10400 (34a); 10400 (34b); 11503 (24); 8167 (34a); 8168, 8169 (24); 8172 (20a); 8173
(37); 11505 (la); 11515 (25); 11517 (37); 11593 (19); (24); 8174 (16); 8176, 8176A (24); 8177. 8182, 8183
11786 (14al); 11796 (16); 11974 (20a); 12007 (Ib); (20a); 8184 (14al); 8185 (24); 8186 (34a); 8187 (5c);
12012 (15a); 12013 (19); 12063 (16); 12182 (20a); 8190 (20a); 8192 (24); 8232, 8237 (20a); 8238 (18);
12184, 12185(16); 12186(19); 12213 (20a); 12214(16); 8251 (24); 8254, 8256 (14al); 8257A (24); 8258, 8261
12917 (13); 13152 (16); 13162 (20a); 13443 (14al); (20a); 8265 (18); 8270-8272 (20a); 8283 (5c); 8294.
13733 (19); 13864 (14al); 14166 (15a); 14363 (17); 8295 (20a); 8298-8300 (14al); 8303, 8304 (24); 8308
14478 (37); 14920 (15a); 14927 (la); 16269, 17855 (18); 8309 (14b); 8313-8316 (24); 8317 (18); 8318-
(20a); 17877 (la); 18412 (28); 18467 (19); 19030 (21); 8320 (24); 8321-8325, 8333-8339 (20a); 8350-8353
20066 (20a); 20512 (Ia); 20651 (15a); 22370 (18); 22388 (24); 8354 (20a); 8356, 8357 (24); 8360 (13); 8362.
(38); 22823 (17); 22824 (20a); 22825 (17); 22833, 22927 8363 (20a); 8368 (18); 8373A, 8373B (34a); 8376-8378
(la); 22929 (15a); 23150 (25); 23199 (4); 23485 (la); (13); 8382 (34a); 8388 (24); 8389-8393, 8395-8404.
24330, 24339 (19); 25507 (18); 25599 (19); 25600 (25); 8410-8414 (Sc); 8427 (16); 8429 (34a); 8437 (24); 8440
25787 (lb); 25899 (25); 26128, 26128A, 26346 (4). (5c); 8441, 8442 (24); 8443 (13); 8444 (Sc); 8445 (13);
McVaugh, R. & W. N. Koelz. 175 (19); 404 (20a); 837 8454 (14b); 8463 (14al); 8464 (20a); 8465 (16); 8466
(16). (20a); 8468 (16); 8471 (24); 8473 (38); 8475 (24); 8478,
Mearns, E. A., 211, 390 (lb); 424 (12); 1623 (43al); s.n. 8481, 8482 (14al); 8483, 8484, 8486 (13); 8488 (18);
(37). 8489 (20a); 8490 (14al); 8491 (43al); 8492, 8493 (10c);
Mears, J. A., 337E (43al). 8494, 8495 (13); 8496 (44); 8497 (13); 8498, 8499 (28);
Medellin, F. L., 4 (la); 1346 (15a); 1348 (14b); 1361, s.n. 8500 (22a); 8501 (13); 8503 (1a); 8504 (13); 8505 (28);
(42). 8507 (la); 8513 (16); 8514 (26); 8519 (24); 8524 (13);
Medina, M. C., 265, 270 (22a); 704 (15a); 1673 (22b); 8525, 8526 (20a); 8527 (18); 8528 (20a); 8533 (14al);
2373 (10a); 2374 (Ib); 2374 (la); 2375 (37). 8534, 8535 (24); 8536 (14al); 8537 (34a); 8539, 8540
Medrano& Delgadillo, 1755 (44). (14al); 8542-8547 (5c); 8548 (20a); 8549-8552 (16);
Meigs, P., s.n. (31). 8553 (15a); 8555 (13); 8557 (15a); 8558 (14al); 8570.
Mejia, D., 40 (18); 93 (34a); 106 (14al); 107 (13); 108 8571 (24); 8574-8576 (13); 8578, 8579, 8582 (15a);
(24); 113(18); 114 (20a); 115(24); 154 (13); 160(24); 8585-8590 (13); 8591 (14al); 8593 (18); 8594 (20a);

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278 Flora Neotropica

Mittak, W. L. (continued) Molina R., A. & E. Montalvo, 21554 (20a).

8600 (14al); 8602, 8604 (20a); 8614 (18); 8618 (14al); Molina R., A. et al., 16132 (15a); 16285 (20a); 16401
8621-8623, 8628-8630 (20a); 8632 (24); 8634-8636, (34a); 16467 (13).
8638-8640 (20a); 8642 (24); 8644, 8646, 8647, 8651, Moloney, A., s.n. (5c).
8653 (20a); 8654, 8655 (24); 8657 (20a); 8658 (18); Montanez,A., 18 (4); 2425 (19).
8659 (14al); 8664 (20a); 8665 (38); 8668 (24); 8671- Monies, N., 1182 (20a).
8675 (24); 8681 (38); 8684 (14b); 8685 (13); 8691, 8693 Montes, S. G., s.n. (20a).
(14al); 8704 (20a); 8705-8711 (13); 8712 (15a); 8714 Montes, S. G. et al., 67 (22a).
(14al); 8717 (13); 8719 (14al); 8720 (34a); 8740, 8741 Montoya, A. E., 82 (14al); s.n. (22a).
(20a); 8760 (34a); 8761 (14al); 8762 (34a); 8769-8772 Moore, H. E., 1591 (la); 3499 (28).
(13); 8773, 8774 (14al); 8780, 8781 (24); 8783 (20a); Moore, H. E. & C. E. Wood, 3772 (15a); 3773 (32); 4048
8791 (14al); 8795 (18); 8797 (24); 8798, 8799 (18); (la); 4070 (32); 4145 (16); 4145 (15a); 4269 (43al);
8800, 8801 (20a); 8804 (14al); 8805 (13); 8806, 8807 4355 p.p. (14al); 4355 p.p. (14b): 4356 (43al); 4479
(14al); 8810, 8813 (20a); 8817 (16); 8818 (24); 8824 (28); 4480 (15a).
(20a); 8829 (Sc); 8830-8832 (18); 8833, 8834 (24); 8835 Moore, H. E. et al., 4251 (22a); 5612 (16).
(18); 8836 (20a); 8837, 8841 (24); 8856 (14ai); 8859 Moore, M. D., s.n. (20a).
(14b); 8863, 8864, 8867 (14al); 8868 (18); 8875 (14al); Morales, A., 4 (la).
8879-8881 (16); 8882 (18); 8883-8889 (20a); 8890 Morales, C. & J. M. Alcocer, s.n. (28).
(14al); 8892 (13); 8893 (14al); 8894, 8895 (15a); 8896 Morales, Ochoa & Sdnchez, s.n. (14b).
(13); 8897 (15a); 8898 (34a); 8899 (13); 8900, 8901 Moran, R.. 1334 (37); 3512 (30); 6220 (28); 6221 (12);
(15a); 8903 (14al); 8910-8916 (13); 8918 (34a); 8922 6432 (30); 8411 (33); 10133 (14b); 10905 (46); 10948
(37); 8923 (13); 8924, 8925 (44); 8929 (32); 8930 (43a 1); (47); 10967 (11); 11114 (47); 11161 (11); 11278 (35);
8931 (10c); 8932, 8933 (28); 8936 (22a); 8937 (15a); 11349 (47); 11360 (11); 11391 (47); 13150 (46); 13247
8939, 8941 (22a); 8942 (28); 8943-8945 (15a); 8948 (29); 13334 (37); 13402 (43al); 13496(46); 13497 (47);
(34b); 8949 (15a); 8950 (13); 8951 (la); 8952, 8953 13586 (46); 13587 (47); 13654 (46); 13701 (47); 13822
(15a); 8959 (28); 8960 (15a); 8962 (38); 8963 (18); 8964 (30); 14532 (31); 14552 (47); 14598 (35); 14834 (46);
(24); 8965 (20a); 8966 (18); 8967, 8968 (24); 8971- 14835 (47); 14855 (31); 15071 (47); 15076 (3); 15442
8973 (20a); 8975 (14al); 8976 (34a); 8977 (24); 8978 (47); 15457 (3); 15561 (46); 15562, 15594 (47); 15647
(14b); 8980 (16); 8981 (24); 8982 (18); 8983 (20a); 8985 (35); 16164 (33); 16390, 16538, 16591 (47); 16605 (11);
(13); 8987, 8988 (5c); 8989 (38); 8995 (18); 8999-9008 17655, 17742, 18024 (47); 18105 (46); 18110 (47);
(20a); 9009 (18); 9010-9012 (20a); 9014, 9016-9018 18243 (46); 18434 (30); 20540, 20762, 20954, 21445
(14al); 9019 (24); 9020 (18); 9041, 9044 (14al); 9047- (46); 23004 (47); 23053 (46); 23062 (47); 23583, 23630,
9050, 9053, 9054 (24); 9063 (20a); 9067 (14al); 9068 23707 (33); 24063 (46); 24501, 24664 (47); 24666 (46);
(20a); 9069-9076 (18); 9077 (20a); 9082 (18); 9083 24855 (11); 25581 (47); 25608 (35); 25624, 26933 (46);
(15a); 9090-9092 (24); 9093, 9094 (20a); 9096 (16); 27871, 27891 (29); 29151 (11); 29467, 29468, 29472
9098, 9099 (20a); 9104, 9106, 9108, 9109 (14a1); 9112- (29); 29839, 29846 (31); 30828 (46).
9127 (13); 9130 (20a); 9131 (18); 9134, 9135, 9139 Moran, R. & J. Henrickson, 10319 (46).
(24); 9144 (14al); 9146 (13); 9147, 9148 (14a1); 9149 Moran, R. & R. F. Thorne, 14266 (35); 14423 (46); 14424
(13); 9150-9152 (15a); 9158, 9162, 9163 (14al); 9168- (47); 14483 (11); 14500 (35).
9170. 9177, 9178 (13); 9179 (14al); 9182-9198, 9200- Moran,R. et al., 24244 (47).
9202 (13); 9204 (13); 9206, 9207 (18); 9208-9210 (15a); Moreno, J., 89 (43al).
9211-9213 (14al); 9215 (28); 9216, 9219, 9220 (14al); Moreno, P. P., 5831 (5c); 8058, 11388, 14253, 14447,
9221 (la); 9222-9224 (20a); 9229 (la); 9233 (26); 9234 24271 (20a).
(17); 9245 (27); 9258 (20a); 9259 (27); 9260 (26); 9261 Moreno, S. G., 150 (20a).
(28); 9262-9265, 9274-9276, 9289-9291 (14al); 9293 Morton,C. V., 9562 (8).
(14b); 9294 (14a1); 9295 (20a); 9296 (ISa); 8542/2 (5c); Motte, M. E., 178 (la).
8742/3, 8844-9 (13); 8952-5 (15a); 9090/2 (24); 9110-2 Moulis, B. & L. McGill, 556 (47).
(13). Mueller, C. H., 438 (14b); 439, 2075, 2124 (28); 2125
Mittak,W. L. & E. Espinosa, 8465, 8468 (16); 8469 (20a); (15a); 2156 (13); 2261 (44); 2281, 2282 (13); 2283 (37);
8473 (38); 8474 (24); 8483, 8485 (13). 2287 (28); 2653 (14b); 2714 (28); 2916 (43a2); 2929
Mittak, W. L. & J. P. Perry,8370 (24); 8373B (34a); 8383 (13); 3064 (45); 3208 (10a); 3210 (37); 3229 (43a2);
(13); 8413-7 (24); 8439 (14al); 8441 (24); 8460, 8461 3435 (Ib); 3436 (12); 3477 (45); 3523 (12); 3532 (la);
(14al); 8775-8779 (18); 8839 (14al). 3533 (37).
Molina R., A., 152, 1678, 2182, 3679 (5c); 6256, 6330 Mueller, C. H. & M. T. Mueller, 33 (14b); 438 (28); 439
(20a); 6368 (18); 6875 (20a); 6878 (5c); 7132 (20a): (1Sa); 510 (28); 537 (43al); 538 (28); 539 (14b); 822
7335a (5c); 7411 (18); 7551, 7575 (20a); 8004 (34a); (13); 1241 (44); 1244 (37); 1261, 1264 (13); 1278 (32);
8465 (5c); 8492 (20a); 8500, 8503-8506, 8509-8513 1339 (28).
(5c); 8514, 8685 (18); 8744 (20a); 10535 (5c); 11600. Muenscher,W. C., 12593 (34a).
11685, 12089 p.p. (18); 12089 p.p. (20a); 12579, 12929 Muller, F. 515 (13); 517 (22a).
(18); 13070, 13125 (20a); 13208, 13393, 14935, 15025, Muller, W., s.n. (13).
15607 (5c); 18506 (18); 18693 (20a); 18704 (Sc); 20423 Musselman,L. J., 4183 (6).
(24); 20608 (20a); 20786, 23373 (5c); 24161 (20a); Mufioz, C. & A. Rebollado, 17 (22a).
24741 p.p. (Sc); 24741 p.p. (20a); 25509, 27888 (18); Naranjo,N., 157 (la).
30265 (14al); 30706 (18); 30794 (20a); 31342 (34a). Narave, H.. 159 (34a): 160, 162, 163 (13); 164-166 (28);

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List of Exsiccatae 279

168 (22a); 795 (28); 797 (14b); 854 (28); 855 (13); 1502 Newman, R. J., 38 (28); 44 (22a); 54 (32).
(16); 1542, 1559, 1599 (38). Nickrent,D. L., 2020 (28).
Narave, H. & Vazquez,798 (13). Nickrent,D. L. & G. Keller, 2028, 2042 (4).
Narave, H. et al., 171, 182, 194 (13); 195 (22a); 196, 256 Nicolas, G. [Frere], 51 (la).
(34a). Nicotas de la Cruz, 14 (15a).
Narvaez, N. & A. Salazar,230 (19). Nixon, K. C., 4054 (41).
Nash, G. V., 825 (8). Nixon, K. C. et al., 3795 (28); 4047 (41).
Nash, G. V. & N. Taylor, 1325 (8). Norris, Peterson& Dowty, 20524 (19).
Nash, G. V. et al., L19563 (20a). Norris, D. H. & D. J. Taranto,14239, 14240 (20a); 14334
Nee, M., 22887 (22a); 22901 (13); 22939 (22a); 23122 (28); 14578 (la); 14583 (19); 14584 (15a); 14707 (20a).
(14b); 23165 (34a); 23174 (22a); 23191 (14al); 23530 Northrop,J. I. & A. R. Northrop,84A, 440 (Sb).
(1Sa); 23531, 23532 (28); 23535 (22a); 27719 (20a); Nunez, C. et al., 8144 (28).
32941 (43c); 32987 (13); 33006 (28); 33008 (14b). Ochoa, S. G., 440 (20a); 475 (14al); 607 (42); 737 (14b);
Nee, M. & G. Diggs, 24813 (13); 24885 (38); 24924 (28); s.n. (20a).
24930 (14b); 24931 (22a); 25226 (28); 25229 (22a). Oersted,A. S., 14797 (20a).
Nee, M. & G. Martin,32206 (14b); 32216 (la). Oliva, O. & S. Herrera,13904 (7).
Nee, M. & K. Taylor, 25867 (15a); 25870, 25871 (28); Olivares, A., 3 (22a).
25872 (15a); 25873 (28); 25874, 25875 (15a); 25877- Oliver, W. W. & M. D. Moore, s.n. (24).
25879 (14al); 25881 (13); 25891, 25897 (22a); 25925, Orcutt,C. R., 255 (47); s.n. (29, 43al).
25936 (14al); 25974 (13); 25976 (14al); 25977, 25979- Ordonez, W., 8595, 8596, 8597, 8598 (20a); 8984, 8985
25981 (13); 25982 (15a); 26061 (34a); 26230 (13); 26231 (13); 9021-9027 (34a).
(14b); 26235 (14al); 26247 (16); 26755 (28); 26760 Orr,R. T, s.n. (20a).
(15a); 26896 (22a); 26924 (28); 26927 (15a); 26965 Ortega,G., 173, 265, 325 (20a).
(13); 26971 (22a); 27004, 27006 (18); 27022 (22a); Ortega, 1. P., s.n. (28).
27024 (13); 27034 (14al); 27054 (34a); 27061 (22a); Ortega,J. G., 173, 4089 (20a); 4090 (19); 5069 (37); 6887
27062, 27064, 27071 (14al); 28834, 28835 (14b); 28849 (20a); 6888 (4); 6902 (19); 6905 (16).
(43c); 28851 (14al); 28855 (43c); 29070 (15a); 29074 Ortega, R. V. & M. Ortiz, 1664 (13).
(28); 29084 (34a); 29100 (15a); 29421, 29478 (14al). OrtegaC., T., s.n. (16).
Nee, M. et al., 19584 (la); 19616 (13); 26049 (13); 26050 OrtegaO., R., 209, 219 (13); 581 (22a); 1885 (38).
(15a); 26061 (13); 26078 (38); 27730 (20a). OrtegaO., R. et al., 127 (28); 161 (22a); 1035 (20a).
Neill, D. A., N192 (20a); 3899 (5c); 5314 (12); 6361 (20a). Ortiz, E., 134, 137 (15a).
Nelson, C., 8451, 10075 (5c). Ortiz, G. & F. G. Medrano,5 (32).
Nelson, C. & E. Vargas,2371 (20a); 2649 (5c). Ortiz, H. T., I (14al).
Nelson, C. & M. Hernandez,1004 (5c). Ortiz, M., 432 (la).
Nelson, C. et al., 3951 (18); 7920 (24). Othon Yanez, M., s.. (20a).
Nelson, E. W., 3 (28); 3? (14al); 4 (13); 5 (16); 7 (43a2); Ownbey, G. B. & F. Ownbey, 1883 (19).
8 (28); 177 (13); 276 (15b); 278 (13); 289 (22a); 941, Padilla, M., 97 (13); 126 (la).
978, 979 (20a); 980 (16); 981 (28); 982 (34a); 983 (13); Palacios, M., 65 (la).
984 (16); 985 (14b); 986 (la); 987 (28); 1486 (15b); Palacios, R. C., s.n. (la, 13, 14al, 15a, 16, 22a, 27, 43al).
1487 (la); 1760 (26); 1761 (20a); 1762 (16); 2040 (26); Palmar,W. & J. H. Riley, 273 (2).
2152, 2158 (20a); 2180 (15b); 2181 (26); 2182 (27); Palmer,E., 15 (20a); 80 (12); 82 (43al); 83 (lb); 90 (30);
2187 (28); 2188 (34a); 2189 (15b); 2406 (20a); 2520 301 (Ib); 303 (43al); 328 (la); 330 (12); 374 (20a); 449
(34a); 2521 (14al); 2522 (16); 2534 (13); 2535 (14al); (lOa); 763 (30); 768, 773 (43al); 774, 775 (Ib); 776
2536 (28); 2537, 2538 (26); 2539 (14b); 2540 (26); 2558, (12); 1299 (42); 1998 (21); 769A (43al); s.n. (10c, 30,
2678, 2854 (20a); 3128 (18); 3130, 3130A (20a); 3185 37, 43a 1).
(14al); 3186 (34a); 3218 (28); 3285 (15a); 3493 (20a); Panti, M. A., 35, 414 (28); 478, 480 (15a); 496 (13).
3650 (34a); 3653 (13); 3673 (20a); 3680 (16); 3681 Parkinson,J., s.n. (15a).
(20a); 3729 (13); 3778 (28); 3894 (43al); 3933 (28); Parra,A., 103, 160 (43al); 103163 (la).
3935 (15b); 3936, 3938 (22a); 3939 (13); 4003 (16); Parry,C. C., 1390 (47); s.n. (9, 11, 43al, 47).
4044 (20a); 4112 (19); 4117 (15a); 4118 (17); 4118 (18); Parry,C. C. & E. Palmer,844 (28); 845 (la); 846 (43al).
4119 (20a); 4498 (43al); 4500 (28); 4501 (41); 4502 Passini, J. & M.-F. Robert, 3093 (lb).
(10a); 4504 (13); 4555 (37); 4556 (43al); 4558 (Ib); Passini,M.-F. [n6e Robert],20105 (43a2); 20109 (42); 642-
4567 (15a); 4568 (12); 4907 (la); 4914 (19); 4915 (37); 20161 (19); s.n. (42, 43a2).
4916 (13); 5013 (12); 6013 (lb); 6043 (36); 6055 (12); Patino, s.n. (13).
6079 (43al); 6137 (13); 6140 (42); 6143 (43al); 6558 Patino,A. S., 421 (22a).
(la); 6563 (13); 6573 (16); 6588 (14al); 6599 (15b); Patino, F., s.n. (22a).
6736 (36); 6845 (20a); 6888, 6888a (14al); 6899 (13); Patoni, C., 22 (12); 138 (lb); s.n. (43al).
6958 (16); 7039 (20a); 7060 (34a); s.n. (14al). Patterson,T. F., 32, 33, 36, 1102, s.n. (44).
Nelson, E. W. & E. A. Goldman, 16 (13); 7457 (43b); s.n. Paxson, Barkley & Westlund,37, 48 (13).
(la). Pearce, R., 2351 (12).
Nelson Smith, J. H., 21, s.n. (5c). Peebles, R. H., 586 (43a2).
Nemesio R., C., 2 (28). Pena, J. M. & Socorro Gonzalez, 695 (28).
Nepamuceno,F. & N. Ubalde, 51-A7, 52-A4 (32). Pennell, F. W., 17389 (43al); 17435 (43a2); 19683 (12);
Nereyda,A. & M. Heinrich,48 (20a). 20125 (19); 20133 (25); 20134 (4); s.n. (16).

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280 Flora Neotropica

Pennell, F W., Dunn & Dziekanowski, 157 (13); 390 Proctor,G. R., 25316 (14al); 25402 (38); 38877, 38897
(28). (5c).
Perez, s.n. (20a). Puente, J. M. de la, s.n. (11, 33).
Perez & Gonzalez, 1218 (la); 1219 (28). Puga, L. M., 2541 (13); 5531 (43a1); 5801 (19); 5806 (16);
Perez, A., 1861 (18). s.n. (17).
Perez, E., 126 (34a); B-178 (13). Puig, H., 791, 6315 (la); 6316 (16); 6804 (28).
Perez, G. S., 277B (24). Purpus, C. A., 256 (la); 273 (28); 662 (13); 1803 (la);
Perez, R. E., s.n. (13, 14b, 32). 1805 (28); 3012 (13); 3514 (26); 3514 (28); 4377 (27);
Perez C., G., 182, 183 (14b). 4378 (28); 4379 (13); 4380 (28); 4381 (34a); 4885
Perez E., R., s.n. (19, 46, 47). (43al); 6924, 6984 (28); s.n. (14b).
Perez de la Rosa, J. A., 10 (25); 94 (16); 159 (18); 200 Quero, H., 2227 (20a); 2231 (34a); 2268 (14al); 2291
(21); 223 (15a); 263 (43al); 273, 285 (lb); 300 (25); (20a).
314 (23); 323 (37); 348, 354, 369, 370, 383, 386, 413 Quero, H. et al., 2186 (16); 2227 (20a); 2228 (26); 2264
(23); 496 (Ib); 528 (25); 610 (23); 654 (25); 767, 771 (16); 2269 (la).
(43al); 875, 877, 887, 931 (Ib); 991 (4); 1024 (19); Quiroz G., P. A., s.n. (13).
1089, 1231, 1305, 1352 (23); 1499 (21); 1898, 1899 Rabb, G. B., s.n. (Sb).
(34b); s.n. (4, 16). Ramirez& Alcocer, 451 (17).
Perez de la Rosa, J. A. & Gonzalez, 259 (16). Ramirez & P. Enrique,s.n. (14al).
Perez de la Rosa, J. A. et al., 1522 (21); s.n. (40). Ramirez,A., 22 (la); 548 (34a).
Perino, C. H., 3146 (15a); 3147 (la); 3334 (22a). Ramirez, E. A., s.n. (44).
Perino, C. H. & J. V. Perino, 3053 (14al). Ramirez,J., 6 (34b); s.n. (16).
Perry,J. P., SALI (24); 3-78 (14al); SAL3-79 (18); SAL7- Ramirez,J. A., s.n. (13, 20a, 28, 40).
77, SAL8-77, GUA 17-78, GUA 19-78 (14al); 20-78 (13); Ramirez,J. R., s.n. (43al).
GUA24-78, MEX24-79, MEX25-79, MEX26-79, Ramirez, M. D., s.n. (13).
GUA28-79 (14al); GUA32-79 (14al); 50-54 (43al); Ramirez, R. & G. Flores, 950 (21).
MEX74-83, MEX84-84 (14al); M96-81, M96-81A Ramirez,R. D., 231 (15a).
(14b); GUA112-78, GUA113-78, MEX151-83 (14al). Ramirez L., A., 22 (12); 480 (15a).
Peters, R. E. K., 253 (43b). Ramos, 156 (43al).
Phillips, A. R., s.n. (21). Ramos, C. H., 133 (43al).
Phillips, E. A., 659 (lb). Ramos, H. J., 138 (20a).
Pilsbry, H. A. & C. H. Harvey,s.n. (12). Ratter.J. A., 5181V (Sc).
Pina, 1., 136 (14b). Raven, P. H., 16808 (47); 16809 (46).
Pineda, A. R., 215 (la); 1037 (27); 1051 (20a). Raven, P. H. & D. E. Breedlove, 19893 (24); 19894 (24).
Pineda-Ochoa,1278 (27). Ray, H. E. et al., 358 (43al).
Pinkava,D. J., 5994 (45). Rebolledo, A., P-6 (43al); P12 (42).
Pipoly, J. J., 4073 (5c). Rebolledo, A. R., 278 (la); s.n. (42).
Piteka, F. A., 275 (20a); 294 (4). Record, S. J. & H. Kuylen, s.n. (5c).
Pittier,H. F., 1822 (24); 8486 (5c). Reeder,J. R. & C. G. Reeder, 3164 (19); 6827 (47).
Plancarte,A., s.n. (32). Rees, J., s.n. (14al).
Ponce, E. G., H-8, H-10 (14al). Reko, B. P., 4009 (28).
Popenoe, H., 1, s.n. (5c). Relame, O. E., s.n. (43al).
Powell, A. M., 1007 (20a). Rendon, A. V., s.n. (14al, 26, 27).
Prado, V. L., 58 (20a). Reyes, M., 10 (14al).
Prats,s.n. (5c). Reyes, P., 3900, s.n. (28).
Preciado,J. T. de la, s.n. (17). Reyna, M. L., s.n. (24).
Prettyman,F. P., s.. (5c). Reyna, O. R., 622 (19).
Prigge, B. A. et al., 4804 (47). Reyna B., O., 81 (16); 225 (23).
Pringle, C. G., 12 (la); 48 (20a); 959 (Ib); 1021 (28); Reyna B., 0. et al., 239 (23).
1448, 1673 (12); 1888(2109) (14al); 1899 (14al); 1964 Reznicek,A. A. & D. R. Gregory,305 (26); 315 (22a); 329
p.p. (15a); 1964 p.p. (28); 1965 (14al); 2109 p.p. (20a); (13); 338 (38); 355 (13); 364 (43al).
2109 p.p. (14al); 2293 (42); 2455 (20a); 2657 (14b); Reznicek, A. A. & S. A. Reznicek, 8120 (34b).
2659 (43al); 2660 (28); 3377 (la); 4018 (43al); 4171, Richards,D. K., 3228 (20a).
4358, 6180 (la); 6243, 6800 (28); 7835 (22a); 8054, Richards,D. K. & C. M. Rowell, 3226 (26).
8090 (14al); 8182 (la); 8785 (22a); 8786 (13); 8787, Ricketson,J. & H. Schmidt, 4819 (29).
8788 (14al); 8789 (13); 9402 (20a); 9403 (la); 9484, Riemer,W. J., 1977A (28); 2018C (37); 2018E (la).
10012 (13); 10013 (28); 10014 (19); 10015 (18); 10016 Rinc6n, E., s.n. (la).
(41); 10017 (26); 10018 (28); 10019 (27); 10139 (10a); Rivas, M. A., X-3 (15a).
10140 (25); 10141 (26); 10142 (32); 10170(15a); 10201 Rivera, G. R., 4 (14b); s.n. (15a).
(28); 10340 (27); 10399 (26); 11727 (la); 11728 (15a); Rivera, M., MRI (15a).
11729, 11730(28); 13207a(42); 13207b, 13207c (43a ); Robbins,T. G., 1434 (24).
13208 (la); 13209, 13262 (14al); 13614 (12); 13615 Robert, M.-F, 5936, 5936B, R59-75 (43a2); s.n. (18, 20a.
(37); 13616 (la); 13618 (37); 13710 (10b); 13711 (26); 43a1, 43c).
13714 (27); 13733 (14al); 13782, 13801 (13); 13803 Robert, M.-F & J. A. de la Cruz, s.n. (42).
(22a); 13804 (28); 13925 (26); 15042 (13); s.n. (la, 13, Robert, M.-F. & J. Passini, 5780 (42); 6416 (44); 6837
14a1, 14b, 15a, 22a, 26, 27, 28, 34a, 34b, 37, 47). (42); 8004 (14b); 9001 (41); 5936(B) (43a2).

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List of Exsiccatae 281

Robert-Passini,M.-F., 10011 (45); 10021 (43b). Rzedowski, J., 329 (43al); 906, 1750 (28); 2750 (13);
Roberts, M. L. & D. Keil, 10291 (37). 5959, 6061 (28); 6062 (43al); 6087 (37); 6642 (16);
Robertson,C. C., s.n. (20a, 22a, 27). 7149 (32); 14084 (19); 14281 (16); 14539 (20a); 14542
Robinson,A. N., 107 (15a); 108 (28); 111 (43al). (18); 15230 (20a); 15265a (20a); 15265b (16); 15279
Robinson, A. N. & R. J. Newman, 106 (37). (la); 15490a (20a); 15890 (34a); 15955 (4); 16490a
Robinson, R. G., s.n. (5c). (20a); 16507, 16508 (13); 16629 (20a); 17164 (13);
Rodriguez& J. Valerio,2131 (18). 17339, 17394, 17399 (20a); 17458 (16); 17611 (43al);
Rodriguez,A., s.n. (42). 17671 (19); 17714 (lb); 17714 (la); 18013 (34a); 18041
Rodriguez,A. & M., s.n. (42). (4); 18049, 18082 (26); 18083 (14al); 18172 (38); 18258
Rodriguez,A. & M. A. Carranza,2370 (43al). (40); 18287 (13); 18535 (38); 18601 (26); 18601 (18);
Rodriguez,A. & A. Vega, s.n. (15a). 19216 (28); 19274a (14b); 19388 (18); 19440 (34b);
Rodriguez,A. et al., 866 (42). 19465 (18); 19467 (26); 19593 (18); 19599 (38); 19620
Rodriguez,C., 5 (16); 16, 29 (Ia). (26); 19621 (17); 19625 (20a); 19644 (24); 19646 (20a);
Rodriguez,D. T., 126 (14b). 19650 (24); 19652, 19656 (17); 19659, 19675 (26);
Rodriguez,L. E. & R. A. Vasquez.s.n. (42). 19676 (14b); 19677 (la); 19683 (14b); 20827 (20a);
Rodriguez, M., s.n. (15a, 28). 21105 (34a); 21689 (14al); 22199 (22a); 22244a (13);
Rodriguez,P. M., 39 (22a). 23171 (14b); 24925 (19); 25170 (la); 25193a (16);
Rodriguez, R. M., s.n. (14b. 15a). 25193b (28); 25545 (14b); 25546 (16); 25712 (14b);
Rodriguez,S., 125 (28). 26235 (25); 27146 (28); 27153 (44); 27154 (37); 27166
Rodz, A. & M. A. Carranza,1444 (42). (32); 30196 (14al); 30305 (20a); 30306 (27); 30396
Roe, K. & S. Mori, 411 (14b). (26); 30397 (27); 30398 (14al); 30630 (24); 31624 (42);
Roe, K. & E. Roe, 1992 (14b); 1999, 2053 (la); 2095 31802 (43al); 31847 (38); 32666 (28); 32667 (ISa);
(20a). 32785 (14al); 33120 (24); 33204 (38); 33995 (14al);
Roe, K. et al., 334, 335 (13); 363 (28); 677, 848, 978, 34683 (42); 37542 (21); 45467 (28); 45789 (43al).
1099, 1105 (20a); 1153 (28); 1754 (20a). Rzedowski, J. & G. B. Hinton, 17275, s.,. (44).
Roezl, B., s.n. (la, 13, 14al, 15a, 16, 18, 22a, 28, 34a, Rzedowski,J. & R. McVaugh,83 (38); 804 (la); 805 (28);
43a 1). 996 (16); 997 (lb); 1045 (25); 1193 (43al).
Roig, T., 6221 (7). Sagra, R. de la, 277 (7).
Rojas, C., 321 (14al). Salas, J., 665, 666 (20a).
Roman, E. H., 198-9 (34a). Salas, J. & Taylor,2228 (20a).
Rosas R., M., 552, 555 (22a). Salazar,J. H., 2 (14b).
Rose, J. N., 1671 (17); 1755, 1997 (21); 2003 (20a); 2122 SalazarJ., F., 1. s.n. (16).
(15a); 2122b (15a); 2123 (la); 2123B (Ib); 2130 (28); Samek, V., s.n. (5a, 7).
2193 (13); 2193 (28); 2194 (19); 2195 (20b); 2262 (20a); Sanchez, F. M., s.n. (14al).
2263 (17); 2264 (20a); 2376 (25); 2377, 2378 (Ib); 2379 Sanchez, R., 68 (43al); 79 (44).
(25); 2405 (Ib); 2741, 2741B (28); 2742, 2742A (Ib); Sanchez, V. L., 1075 (20a).
2777 (15a); 2989 (19); 2990 (la); 3004, 3004A (25); Sanchez A., P. & E. Neri, s.n. (22b).
3005 (15a); 3025, 3025A (17); 3026 (21); 3027 (la); Sanchez M., H., 2682 (15a).
3068 (20a); 3083, 3586 (19); 3587, 3588 (Ib); 3726 Sanders,A. C. et al., 2664 (25); 2697 (Ib); 4449 (19).
(20a); 3727 p.p. (13); 3727 p.p. (25); 3736, 3737 (20a); Sandoval,A. J., 3 (13); 2037 (la).
11661 (Ib); 16001, 16186 (30). Santamaria,A., s.n. (14b).
Rose, J. N. & R. Hay, 5336 (la); 5403 (15a); 5537, 5780 Santana,M., 892 (37); 2948 (la).
(13); 5781 (16); 5968 (la); 6076 (13); 6234 (la); 6325 Santos, J. V., 3090 (22a); 3580 (20a); 3623 (22a).
(13); 6326 (la). Saylor, K., 22, 25 (22a); 67, 68 (20a).
Rose, J. N. & W. Hough, 4291 (22a); 4292 (15b); 4293 Saylor, L. C., 21 (22a); 41, 42, 52, 53 (14al); 41-45 (14b).
(28); 4294 (la); 4639 (26). Saynes, A., 89 (14b); 123 (16); 200 (28); 234 (14b); 240,
Rose, J. N. & J. H. Painter,7014 (28); 7015 (15b); 7153 249 (28); 265 (14al); 270 (28).
(la); 7216 (28); 7508 (20a); 7952 (15b); 7953, 7971 Scheinvar,L., 539 (43al).
(13). Schiede, C. J. W., 378 (la); 915 (15a); 944 (28); 945 (la);
Rose, J. N. & J. S. Rose, 11524 (43al). 946 (28); 955 (43al); 1109, 1117 (la); s.n. (15a, 16, 28,
Rose, J. N. et al., 8481 (22a); 8482 (la); 9092 (42); 9093 22a, 28, 43al).
(43al); 13087 (20a). Schiede, C. J. W. & F. Deppe, 13, 407 (28); 1108 (22a);
Rothrock,J. T., 652 (lOa); 654 (36); 1001 (36). s.n. (la, 15a, 22a, 28).
Rowe, K. & E. Rowe, 1968 (34a). Schipp, W. A., 386. 709 (Sc).
Rowell, C. M., 3032 (20a); 3049B (la); 3227 (15a); 3229 Schmutzenhofer,H., 1, 3 (24); s.n. (18).
(34a); 3898 (16). Schneider,R. A., 929 (13); 957 (44).
Rowell, C. M. & Class, 3897 (27). Schnooberger,I., 7942, 7943 (43al).
Rowlee, W. W., 232, 233 (Sa). Schoenwetter,J., JSOX-156 (28).
Rubin, N., 2 (19). Schomburgk,R. H., 43 (8).
Rushforth, K. D., 415A (37); 431 (43al); 440 (44); 441 Schott, A., s.n. (43a2).
(13); 465 (37); KR495 (12); 497 (15a); KR498 (14b); Schrenkvon, H., I (14al); 2 (18); 3 (34a); 6 (16); 7 (24);
557 (37); KR563 (41); 564 (43a ); 581 (17); 586 (14al); 50 (8).
KR588 (la); 591 (15a); 609 (16); 612 (20a); KR625 Schroeder,A. H., 95 (45).
(14al). Schuhrt& B. M. Massie, 9 (18); 2265 (43al).
Rutten-Pekelharing,C., 431 (2). Schultes, R. E., 586 (20a); 765 (38).

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282 Flora Neotropica

Schultes, R. E. & Reko, 910 (20a). (28); 7613 (26); 7868 (38); 8154 (14al); 8260 (28); 8261
Schultes, R. E. et al., 4 (2). (14b); 8281 (16); 8304 (14b); 8322 (15b); 8403, 8404
Schultze, L., 901 (20a). (26); 8425 (13).
Schumann,W., 1333 (43al). Soto N., J. C. & S. Silva, 1927 (la).
Schwerdtfeger,F., 2 (18); 3 (24); 4 (14al); 6 (24); 9, 11, Soto N., J. C. & A. R. Soto, 7968 (26).
13, 19 (20a); 23, 24 (18); 25 (20a); 30, 32 (24); 34, 37 Soto N., J. C. & F. Soto R., 6543 (14al).
(20a); 38, 40, 42 (24); 43 p.p. (18); 43 p.p. (14al); 44 Soto N., J. C. et al., 5526 (la); 5857 (14al); 6293 (16);
(16); 46 (13); 47 (14al); 48 (24); 49 (20a); 50 (18); 51 6502 (16); 7915, 7965, 8121 (15a); 8140 (16); 8144 (4).
(34a); 52 (24); 53-55 (13); 56 (14al); 57 (16); 59, 61 Soule, O. H., 453, 456 (37); 483, 485 (19); 489, 493 (37);
(34a); 62 (14al); 64, 65 (13). 513 (12); 2153, 2154 (la); 2160 (12); 2164 (19); 2210
Seaton, H. E., 216 (13). (28); 2211 (25); 2218 (la).
Seemann, B. C., 1060 (Ib); 1959 (43al); 1960, 1961 (19); Sousa, M., 2522 (22a); 2602 (17); 2603, 3246 (20a); s.n.
1962 (la); s.n. (19). (la).
Seidler, P., s.n. (38). Souviron, M. J. & C. 0. Erlanson,60 (20a).
Seymour, F. C., 3650, 4833 (5c); 7635 (24). Spellenberg.R. et al., 10063 (25); 10064 (Ib); 10065 (25);
Shafer, J. A., AD (7); 425 (2); 1688, 3156, 3206, 3216, 10069(12); 10076(Ib); 10079(25); 10080(37); 10081,
3799, 3834, 4087, 4112, 4112A, 4154, 4158, 4174, 4189, 10082 (Ib); 10083 (10a); 10086 (4); 10090 (Ib).
4194, 4254, 8128, 8453, 8842 (7); 10687, 10905, 11017, Spellman, D. L., 1530 (Sc).
11207 (2); 11209, 11255, 11697 (5a); 11715 (2); 11867 Spellman, D. L. et al., 978 (14b).
(5a); 11944 (2); 11968 (5a). Spjut, R. & R. Marin,5299 (29).
Shank, P. J., 122 (5c); 286, 287, 14755 (20a). Spross, B., 45 (24); 109 (38).
Shannon, W. C., 3616, 3622 (13). Stahl, E., s.n. (22a).
Sharp,A. J., 461 (38); 4521 (20a); 4634 (18); 4634 (20a); Standley, P. C., 1702, 4123 (20a); 8088 (18); 9543 (24);
4634 (14al); 4649, 4694 (38); 4696 (14al); 41322 10080, 10116 (20a); 14963, 16886 (Sc); 20399 (24);
(43al); 44166 (la); 45779 (44); 44873 (19); s.n. (13, 24227 (5c); 25177 (18); 25386, 55921 (20a); 56144 (18);
38). 58510, 58695 (20a); 60994 (14al); 62456 (16); 62552
Sharp.A. J. & E. Hernandez,44502 (la). (20a); 62689 (34a); 64397 (20a); 67370 (34a); 67618,
Sharp,A. J. & E. H. Xolocotzi, 46237 (38). 67618, 67704 (13); 69330 (20a); 69700 (24); 75523
Shaw, G. R., 1170 (19); s.n. (la, lb, 2, 5a, 12, 13, 14al, (20a); 76252 (24); 77109 (20a); 77595 (24); 80828 (16);
15a, 15b, 16, 18, 19, 20a, 22a, 26, 27, 28. 37. 41. 42, 80987 (18); 81156 (13); 81508 (16); 81663, 81692,
43a ). 81713 (13); 82066, 82083, 82091 (20a); 82111, 82336
Shilom Ton, A., 650, 717 (14b); 1829 (15a); 1879. 1940 (16); 82339, 82343 (20a); 83977 (13); 84014 (34a);
(20a); 2539 (38). 84106 (13); 84115 (14al); 84174 (34a); 84406 (14al);
Silva, G., S. Aureoles & C. Pizarros,8649 (28). 84477, 84519 (34a); 89696 (18).
Sim6n, F., 122 (38); 123 (22a). Standley, P. C. & P. Chac6n, 5531 (5c); 6523 (20a).
Sim6n, F. & M. A. Martinez, I (19); 29 (lOa). Standley, P. C. & L. 0. Williams, 1288 (5c).
Siplivinsky, Beck & Davis, 3910 (Ib); 3911 (37). Standley, P. C. et al., 1288 (5c).
Skutch, A. F, 113 (18); 295, 753 (20a); 822 (14al); 852 Stanford,L. R. et al., 110 (43a2); 148 (42); 456 (37); 2397
(13); 868 (34a); 1270 (13); 1566 (18). (43al); 2451 (28); 2675 (41).
Small, J. K. & J. J. Carter, 1249 (6); 8497, 8881, 8973 Stead, J. W., 22 (5c); 190 (18); 192, 194 (18); 229 (20a);
(Sb). 232 (14b); 235 (18); 271, 272, 273 (24); 303, 304 (13);
Small, J. K., J. J. Carter& G. K. Small, 3575, 3635 (6). 305 (14al); 306, 307 (16); 308 (14al); 309, 310 (16);
Small, J. K. & G. V. Nash, s.n. (6). 332 (18); 337 (34a); 338 (14b); 233-255 (14al); 278-
Smith, C. E., 4438 (28). 302 (14b); 335 (18).
Smith, C. E. & S. Kitchen, 5034 (14b). Stead, J. W. & B. T. Styles, 39 (14al); 54, 62 (5c); 77, 78
Smith, C. E. & N. Tejada,4457 (15a). (5c); 95 (15a); 97-99(16); 100 pp. (18); 100 p.p. (14a1);
Smith, C. E. et al., 3850, 3850-bis (14al). 105 (5c); 116 (5c); 125 (18); 126 (20a); 134-136, (18);
Smith. C. L., 1475 (22a). 141 (18); 142 (16); 143 (14al); 144 (14b); 145 (14al);
Smith, D. N. & D. Clancy, 557 (34a); 558 (14al); 569 148-150 (14al); 151 (14b); 154 (14al); 156 (16); 158
(13); 571 (34a). (20a); 163 (16); 168 (20a); 175 (38); 177 (20a); 178
Smith, J. G., 361 (22a); 492 (13). (38); 180 (14al); 181 (38); 190, 192 (34a); 193 (13);
Snajberk,K. & E. Zavarin.s.n. (45). 194 (14b); 195 (13); 201 (34a); 213 (20a); 215 (38); 233
Snook, s.n. (38). (20a); 234, 235 (5c); 244 (5c); 247 (20a); 248 (Sc); 249
Socconini, A., s.n. (28). (32); 250 (36); 251 (5c); 252, 253 (20a); 254 (18); 257
Solbrig, O. T. & R. Orduff, 4612 (12); 4637 (lOb). (20a); 259, 270 (24); 272, 273, 275, 277 (18); 278 (16);
Solis, R., 42 (24). 281 (18); 281a (16); 284-286 (14a1); 287 (14al, 14b);
Solorzano, L. B., 654 (la); 655 (15a); 656 (14al); 657 288-292 (14al); 293 (14al, 14b); 294-307 (14al); 309
(15a); 802, 803 (20a); s.n. (la). (34a); 336 (14b); 337, 338 (14al); 339 (16); 341 (34a);
Sosa, A., 4 (27). 342-345 (14a ); 346 (14a1, 14b);347 (14a1); 349 (14b);
Sosa, A. H., 2 (18); 10 (27). 350-368, 372 (14al); 375 (16); 376 (14al); 377 (15a);
Sosa M., M., s.n. (15a). 378 (18); 381 (16); 383, 385 (20b); 387 (14b); 388 (la);
Soto N., J. C., 3448, 3640 (20a); 4896 (la); 6110 (20a); 389 (17); 390 (14b); 392 (18); 393 (20b); 419, 420 (38);
6114, 6131, 6135 (14al). 421 (18); 423-447, 449 (14b); 450 (20b); 455, 456 (17);
Soto N., J. C. & S. Aureoles C., 7436 (28); 7463 (4); 7479 461 (14al); 462 (la); 465 (27); 466 (16); 470 (27); 473

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List of Exsiccatae 283

(13); 474 (19); 475 (21); 476, 477 (17); 478 (14al); 35 (18, 20a); 36-41 (18); 42 (Sc); 44-56 (20a); 57 (la,
479-492, 503, 504 (17); 506-522 (14al); 523-527 (17); 24); 58, 59 (24); 60 (17, 24); 61-64 (24); 65 (18); 72
528 (la); 529 (16); 530 (27); 531 (26); 532 (17); 533 (20a); 73, 77 (24); 78, 79 (5c); 80. 95-101 (24); 102
(18); 534 (14al); 535 (16); 536, 537 (15a); 538 (28); (20a); 104 (5c); 105-107 (24); 108, 109, 111, 112, 114
539-558 (17); 559 (14al, 17); 560-563 (17); 565-589 (20a); 115 (22a); 117 (34a); 118 (22a); 130 (20a); 135
(14al); 590, 592 (15a); 593 (43al); 595-601 (14al); 602 (22a); 143, 144 (18).
(28); 603-606 (14b): 607, 607b, 608-613 (14al); 614 Styles, B. T. & P. S. McCarter,s.n. (24).
(10c); 615 (43al); 616 (14al); 625 (13); 628 (10c); 629 Styles, B. T. & J. W. Stead, s.i. (24).
(32); 630, 631 (37); 632 (13); 633 (44); 634-636 (37); Styles, B. T., J. W. Stead & P. S. McCarter,17 (20a).
637 (15a); 638-662 (14al); 663. 664 (22a); 668-692 Styles, B. T. & K. Wilkinson, 1 (27); 3 (19).
(14al); 693 (28); 694 (13); 695 (22a); 696 (15a); 697 Sullivan, J. R., 518 (22a).
(28); 700 (5c); 725, 726 (Sc); 727 (20b); 734 (14al); Susan, H., 29 (41); 33 (28).
744 (14b). Swartz, O. P., s.n. (8).
Stead, J. W. et al., 730 (15a); 731 (4); 734 (14b); 735 (26). Taboya,S. R., 38 (34a).
Stephenson,R., s.n. (29). Takaki,F., S109, s.n. (la, 14al, 15a, 16, 18. 28).
Stevens, W. D., 1128 (5c); 3143 (20a); 7638 (5c); 7753 Taylor,A. A., 232 (2).
(5c); 10177 (20a); 10270 (24); 12787a (5c); 21679 (5c). Taylor,J. & C. Taylor, 1183 (26): 5310 (43a1); 6062 (20a);
Stevens, W. D. & M. Fairhurst,1944 (20a). 6203 (Ib); 6204 (19).
Stevens, W. D. & A. Grijalva, 15073 (18). Taylor. K. & H. Narave, 347, 365 (14al); 369 (15a); 370
Stevens, W. D. et al., 14917 (24). (28); 371, 1211 (34a).
Stevenson, J. A., 719 (8). Taylor, K. & M. Nee, 54, 64 (20a); 186 (22a); 209, 217
Stevenson, R., s.n. (29). (14al); 219 (28); 232 (14al); 236 (43al); 244 (14al);
Stewart, R. M.. 933 (45); 1243 (43a2). 250 (43al); 252 (22a); 253, 269, 287 (14al); 310 (22a);
Steyermark,J. A., 29699, 32162 (20a); 32703, 34083 (34a); 323 (la): 324 (22a); 325 (28); 329 (30).
34173, 35493 (13); 35571 (34a); 38416, 38548, 39665 Taylor,K. et al.. 186 (22a).
(5c); 42277 (20a); 42623 (34a); 42822 (24); 43122, Taylor,M.. 104 (12); 108 (43al); 157 (13); 192. 855 (28).
43272 (34a); 43878 (20a); 45155a (5c); 45269 (5c); Taylor,N., 373 (8).
47038 (16); 47473 (13); 47979 (18); 47979 (24); 49489 Tellez V., 0., 10121 (16).
(38); 49578 (18); 49603, 49729 (38); 50321 (13); 50449 Templeton,B. C., 123a (16); 8677 (43al): 9431 (17); 9443
(20a); 50497 (16); 51891 (38); 52264 (34a). (la).
Stinson, M. L.. s.n. (30). Tenorio L.. P. & R. Hernandez,018 (43al).
Stockwell, P., 2005, s.n. (30). Tenorio L., P. & C. RomeroT., 9034 (14al); 13296 (3).
Straw, R. M., 1968, 2003 (10a). Tenorio L., P. et al., 8721 (38); 10554 (43b); 12769 (38);
Straw, R. M. & D. P. Gregory,21, 1002 (13); 1069, 1092 13761 (14al); 14819 (24); 14881, 14882 (20a).
(14al); 1139 (14b); 1210(16); 1210(13); 1218 (la). Terrazas,S., s.n. (13).
Straw,R. M. & M. Forman. 1350, 1376 (43al); 1411 (42); Thieret, J. W.. 36980 (13); 36981 (44).
1438 (43a2); 1477 (28); 1478 (14al); 1625, 1691 (lb); Thomas, C. H., 65 (la).
1694 (12); 1756 (43al); 1757 (Ib); 1775 (28); 1968 Thomas, J. H., 7879 (43b).
(10a); 1970 (la); 2003 (10a); 2040 (19); 2041 (37); Thompson, Rawlins & Zimmerman,522 (43a2).
130251 (10a). Thorne, R. F., 31988, 62364 (47).
Streator,C. P., s.n. (28). Thorne, R. F. & Lathrop,40455 (20a).
Students Univ. Illinois, 957 (44). Thorne. R. F & W. Wisura,57435 (47); 61124 (33).
Styles, B. T., I (la); 2 (16); 3 (14al); 4 (la); 5, 6 (14al); Thorne, R. F. et al., 62364 (47).
7, 8 (la); 15 (26); 16 (la); 20 (16); 23 (20a); 24 (17); Tirado,J. L., s.n. (28).
25 (18); 27, 28 (27); 29, 30 (16); 34, 36, 37 (20a); 41 Toledo. C., 24 (la).
(4); 42 (17); 43 (27); 44 (22a); 45 (20b); 46 (26); 47 Tomlin, S., 175 (18); 189 (24).
(32); 49 (18); 50 (17); 51 (16); 54 (20a); 55 (17); 55 Torke, Dunn & Wallace. 182 (la).
(14al); 56 (la); 63 (16); 65 (20a); 66 (la); 67-69 (4); Torres& Vargas,s.n. (13, 22a).
74 (20a): 76 (17); 77 (20a); 79-81 (13); 89 (la); 93, 94 Torreset al.. 1031 (43al).
(43a1); 100 (20b); 101 (14b); 104 (28); 105 (34a); 107 TorresC., R.. 2 (15a); 22 (14al); 44 (16); 57 (17); 63 (19):
(28); 108 (14al); 109 (13); 110 (28); 112, 113 (22a); 163, 164 (16); 1081 (44); 6693 (14b).
114 (34a); 115 (24); 117 (18); 119 (24); 121 (16); 122 TorresC., R. & A. Campos, 10859 (18).
(20a); 126 (24); 149 (24); 54A (20a); s.n. (28, 32). TorresC., R. & L. Cort6s,9359 (18).
Styles, B. T, J. Burley,S. Favela et al., I (43al); 2, 3 (42); TorresC.. R. et al.. 7144 (14b); 7145 (la): 11850 (16).
4 (45). Torres,R. & D. Neil, 2296 (20a).
Styles, B. T. & C. Cozatl, 4 (la). Torres,R. & R. Cedillo. 2750 (4).
Styles, B. T, S. Favela et al., I (Ib); 2 (28); 3 (12); 4 (19); Townsend,C. H. T., s.i. (11).
5 (Ia); 6 (4); 7 (17); 8 (20b); 9 (1Oa);10(25); 11 (lOb); Townsend,C. H. T. & C. M. Barber,355, 356 (12); 359
12, 13(37); 14 (14al); 15 (20a); 16 (14al); 17, 18 (10b); (lb).
19 (43al). Traylor,J. D.. 44 (la).
Styles, B. T., S. de Hoogh & L. Favela, 3 (44); 4-6 (41); Trejo, R. A., s.n. (13).
16, 17 (32); 19,24 (13). Trelease, W., 179 (14b).
Styles, B. T & R. F. Kemp, 1-4 (20a); 6 (18); 7 (20a); 8- Trevino.C., s.n. (14b).
10, 14, 15 (Sc); 18, 20 (20a); 23-28 (20a); 29 (18); 30- Troll, C.. 118, 450 (28); 576 (la).

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284 Flora Neotropica

Trott, S. et al., 183 (20a). Villasefor, J. L. & W. W. Thomas, 785, 786, 787 (20a);
Tucker,J. M., 766, 809, 982 (20a); 1067 (34a); 1114 (14b); 829 (16).
1203 (34a); 1205 (20a); 2479 (12); 2505 (Ib); 2579 (37); Villegas, I., s.n. (27).
2580 (12); 2582 (10a); 2594 (37); 2596 (9); 2600 (43a2); Vincelli, P. C., 511, 625 (5c); 719 (24); 827 (20a).
4206 (47). Viveros, Zendejas & Mancera,s.n. (20a).
Tuerckheimvon, H., 1016, 2065 (18); 3088 (8); 8755 (18); Viveros, I. & A. Mancera,S114 (20a); SI15, s.n. (15a).
8756 (20a); s.n. (18). Viveros, M. A., 3 (35); 001 (3); 007 (11).
Ubaldo, N., s.n. (26). Viveros Z., F & Manzera,s.n. (27).
Ugent, D. & R. C. Flores, 1634 (14al). Wadsworth,F. & F Bazan, s.n. (38).
Uhde, C. A., 723 (16). Walker,L. W., s.n. (30).
Ullea S., M., s.r. (15a). Watkins,Wallace & Dunn, 620 (15a).
Underwood, L. M. & F. S. Earle, 1368 (7). Weber,W. A. & L. A. Chevette, 11801 (19).
Valdez, J. & M. A. Carranza,262 (43al). Webster,G. L., 11621 (la).
Valdez, J. et al., 1581 (43al). Webster,G. L. & G. J. Breckon, 15970 (21).
Vald6zT., V. et al., 15 (32). Webster,G. L. & S. L. Breckon, 15507 (43al).
Valenzuela,Z., 3 (17). Webster,G. L. et al., 11328 (28); 20205 (22a).
Valerio,J., 1627, 1699 (20a); 2131 (24). Webster,J. D., I (43al); 7 (la).
Valerio,J. & Rodriquez,202, 1655, 1699 (20a). Weed, C. M., s.n. (12).
Valeur,E. J., 252 (8). Weider,Bennett, Dunn & Torke, 255 (12).
Vargas,Ruiz, Mendez & Diaz, 132 (11); 133 (35). Weir,J. R., s.n. (8).
Vargas,F., 13 (la); 58 (15a). Welcer, D. & D. B. Dunn, 53 (la).
Vargas,F. M., 40, 41 (27); 42 (la); 58 (15a). Wendt, T., 556B (43al); 2481, 2494, 2498 (43c); 2499
VargasN., A., S100 (26); s.n. (la, 3, II, 14al, 15a, 26). (43a ); s.n. (43c).
Vasquez,J., KR561 (41); s.n. (14b, 18). Wendt.T. & D. K. Bailey, 2481, 2495 (43c).
Vasquez, R. A. & L. E. Rodriguez,645 (43al). Wendt,T. & E. J. Lott, 638 (43a2); P16 (45).
Vasquezde la Parra,R., s.n. (19, 20a). Wendt,T. et al., 116 (45); 1944 (45).
Vaughan,J. et al., 1043 (43al). Wendt,T. & D. Riskind, 1658 (45).
Vazquez, 166 (16); 3205 (la). West, R. C., Q-23, Q-18a (14al).
Vizquez, M., 1995 (22a). Whight, A. E., 75 (5b).
Vaizquez.N., 84 (14al). White, S. S., 528 (12); 565 (Ib); 959 (43a2); 2344 (12);
Vfizquez,R. B., 74 (14al). 2345 (Ib); 2346 (43al); 2732 (Ib); 2748 (10a); 2827
Vaizquez,V., 475, 513 (22a); 514 (15a). (Ib); 2828 (43a2); 2829 (12); 3174 (Ib); 3175 (12);
Vazquez S., J., s.n. (20a). 3176, 3360 (10a); 3409 (Ib); 3423, 3467 (37); 3509,
Vega A., R., 63, 576 (15a); 577 (13). 3941 (43a1); 3942 ( b); 3943 (12); 4202 ( b); 4266 (12);
Vela & Guzman, s.n. (12). 4321 (43a2); 4738 (43al); 4789 (Ib); 5246 (20a); 5406
Vela Galvez, L., 126 (22a); 1090 (43a2); 1149 (18); 1153 (34a); 7264 (12).
(la); 1258 (22a); 1302 (14al); 1326b (22a); 1333 (38); White, S. S. & R. M. Chatters,230 (14b).
1348(16); 1349 (20a); 1409(32); 1465 (22a); 1553 (38); Whitefoord,C., 2319 (Sc).
1564 (18); 1592 (20a); 1596 (17); 1599 (20a); 1600, Whitford,H. N. & L. R. Stadtmiller,54 (20a); 76 (5c).
1605, 1608 (26); 1609 (14b); 3005 (Ib); s.n. (13, 14al, Wiggins, I. L., 9055 (47); 9079, 9176 (11); 11239 (46);
19, 22a). 11261 (11); 11270 (47); 11274 (35); 11683 (Ib); 11684
Velasco, V., s.n. (18). (12); 11691 (Ib); 13196 (19); 13205 (43al); 16549,
Velazquez, B. B. & F. P. Valera,s.n. (14b). 20974, 21634 (47).
Ventura,10771 (13). Wiggins, I. L. & D. Demaree, 4920 (11); 4963 (3); 5013
VenturaA., F., 384, 486, 745 (13); 789, 885 (Ia); 900 (35); 5017 (11); 5033 (3); 5052 (47).
(15a); 907, 928, 977, 1031, 1086, 1166, 1242, 1437, Wiggins, 1. L. & W. R. Ernst, 51 (30).
2015, 2474, 2606, 2733, 4117, 4370 (la). Wiggins, I. L. & J. W. Gillespie, 4123 (11); 4161, 4169
Ventura,E. V., 617 (Ia). (47).
Ventura,F. A., 1548 (28); 3016 (14b). Wilbur,R. L. & C. R. Wilbur, 1699 (20a); 1902 (4); 1983,
VeraA., A., s.n. (16). 1986 (25).
Victoria,A. & R. Fernindez, s.n. (30). Wilkinson, K. & B. T. Styles, I (23); 2 (21); 3 (19).
VidauriD., E., s.n. (16, 17). Williams, L. 0., 13156 (34a); 17063 (20a).
Vila, A. K., I (22a). Williams, L. O. & A. R. Molina R.. 1475, 11211 (5c);
Villa, 183 (22a). 11281 (20a); 11315(5c); 11864 (20a); 12234(18); 12271
Villa, A., s.n. (11, 33). (20a); 14754 (5c); 19029 (20a).
Villa, J. V.. 22 (14b); sn. (20a, 22a, 41). Williams, L. O. & T. P. Molina, 22228 (13).
Villamar,A., s.n. (la, 16). Williams, L. O. & R. P. Williams, 18401 (20a).
Villarreal,J. A., 3082 (43al); 3983 (10a). Williams, L. O. et al., 22228 (13); 22490 (24); 22496 (16);
Villarreal,J. A. & M. A. Carranza,3404 (28); 3745, 4041 23142 (34a); 24767 (24); 25399 (34a); 27969 (24); 40003
(43al); 4242 (44); V3313 (43al). (20a); 41500, 41725 (34a); 42900 (Sc); 43400 (18).
Villarreal,J. A. et al., C219, 2690 (43al); 2976 (28); 3143 Williams, R. M. C., 5 (24); 9 (16).
(32); 3497 (44); 4945 (43al). Wilson, P., 975 (20a); 7694 (5b); 9236 (5a).
Villarreal,L. M., 5531 (43al); 5801 (19). Windier,B. K., 2953 (20a).
Villarreal,R. C., s.n. (44). Windler,D. R., 2964 (14al).

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List of Exsiccatae 285

Windier.D. R. & B. K. Wilson, 2953 (14al). ZamoraC., P. & V. Velasco, s.n. (38).
Wislizenus, F. A., 155, 231 (37); 232 (lb); 233 (12); 534 Zamudio, S., 2124, 2380 (43al); 3828 (16).
(9); s.n. (12). Zanoni, T. A., 2518 (12).
Witsberger,D. & M. L. Reyna, 684 (18). Zanoni, T. A. & M. Mejia, 12288. 16658, 17954. 35801
Wolffsohn, A., 5a (24). (8).
Wood, C. E. & E. Atchison, 4251 (22a). Zanoni, T. A. & J. Pimentel, 20700, 25908 (8).
Woronow,G., 2773 (20b). Zanoni, T. A. et al., 3897A, 10889, 12933. 29734, 32443.
Worthington,R. D., 7884 (43al). 36453, 40760, 40761, 40832, 40833. 41210, 41212 (8).
Worthington,R. D. et al., 9368 (20a). Zavaleta, P. & F. Guevara.8490 (41).
Wright, C., 598, 1462, 1462A, 1462B, 1462C (7); 3189 Zenteno, M., 1 (27); 4 (17).
(2); 3189A, 3190, s.n. (2). Zepeda, M., 5 (26).
Wright,C. et al., s.n. (8). Zepeda, S., 34 (22a).
Wynd, F. L. & C. H. Mueller, 166 (45); 630 (37); 650 (9). Zing, R. M., 45 (19).
Yanez, M., 175A (44). Zobel, B., 1 (14b); 2 (37); 3 (14b); 4 (28); 5, 6 (13); 7. 8
Yanez, O., s.n. (18, 26, 38). (28); 9 (13); 10 (15a); 12 (14b); 14 (15a); 16 (14b); 17
Youngpeter,J. & T. J. Cohn, 40 (43al). (28); 18, 20, 20X (10c); 21 (14b); 23 (15a); 24 (28);
Yuncker,T. G. et al., 5686 (20a); 6205 (18): 8182 (5c). 1555 (43a ).
ZamoraC., P., 1002 (18).

INDEX OF LOCAL NAMES

Species numbersare given in parenthesesafter each local name. The numbersrefer to the NumericalList
of Taxa, above.

acalote (34) pino chalmaite (14)

acahuite(37) pino chino (1, 4, 10, 20. 21, 28)
ayacahuite(34) pino colorado (20. 28)
chichonuda(31) pino gretado(34)
ocote (1, 4, 5c, 13, 15, 17, 18, 22, pino hayarin(17)
23, 24, 25, 27. 28, 35, 39) pino lacio (14. 16, 19)
ocote blanco (15) pino liso (14)
ocote chino (20) pino llor6n (19)
ocote dormido (19) piii6n (40, 41, 42, 43, 44, 45, 46, 47)
ocote gretado (34) pino nayar(36. 37)
pinabete(17, 34, 36. 37, 38) pino negro (11)
pino (1, 2, 3, 4, 5c. 7, 9, 11, 13, 22, piionero (43, 45)
23, 24, 27, 29, 30. 33, 39) pifi6n prieto (41)
pino amarillo(10, 19) pifi6n real (40)
pino barbacaida(19) pino oriundo (8)
pino blanco (10, 14, 16, 17, 25, 37, pino prieto (1, 16, 20, 21, 32)
38) pino real (10, 12, 15, 17, 25, 28)
pino de azucar (35) pino rojo (27)
pino del caribe (5a) pino rosillo (28)
pino de montezuma(15) pino tabla (34)
pino de pifia (31) pino triste (19, 22)
pino cans (18) South Floridaslash pine (6)

INDEXOF SCIENTIFICNAMESOF PLANTSAND ANIMALS

New names and combinationsare in boldface and synonyms are in italics. Page numbersin boldface indicate primary
page [Link] numberswith an asterisk (*) indicate pages with illustrationsor maps.

Abies, 11, 14, 17, 26, 49, 58, 144. 173, 198, 213, 242 Agave, 126, 219, 229. 232, 236, 242
concolor, 58, 82, 111, 205 lecheguilla, 245
guatemalensis,58, 82, 112, 207 Alnus, 84
religiosa, 58, 134, 146, 161 Andropogon,99
vejarii, 193 Apinus, 62
Acacia, 139, 153 Araucariaceae,48
Adenostoma, 186, 253 Arbutus,84, 102, 213, 232

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286 Index of Scientific Names of Plants and Animals

Arceuthobium,58 Mimosa. 139. 229

Arctostaphylos,75, 102, 186, 232, 236, 242, 253 Opuntia.75, 126, 229. 232, 236. 245
Artemisia, 186, 253 Persea. 84
Brahea,226 Picea, 11. 14, 31,42,49, 58, 144
Bromeliaceae, 119 Pinaceae, [11] 2, 4, 11, 14. 15, 24. 29, 30. 36. 42, 48, 49.
Brosimum,216 55
Buddleja, 126 Pinus, 4, 5, 6. 7, 8. 9, 11, 13, 14, 15, 16*, 17, 18*, 20*,
Byrsonima, 153 22, 23*, 24, 25*, 27*, 28*, 29, 30, 31. 35, 36,
Cactaceae, 226, 232 38, 41, 42. 43, 47, 48. 49, 50*, 51*. 52*, 55, 56,
Calliandra, 139, 153, 169. 238 57. 58, 61, 62, 63, 75, 84, 93, 99, 102. 105. 116,
Carpinus, 169 120, 135, 148, 184, 198, 224, 226, 238. 240, 254
caroliniana,216 subgen. Balfouria, 199
Carrvopitys.62 subgen. Cemnbrapinus, 199
monophylla,247 "div."Crassisqcuanmue 71
Cassia, 126 subgen. Diplopinus. 71
Ceanothus, 186, 232. 241. 253 subgen. Diploxvlon. 4. 13, 30, 31. 32. (36), (42). 71
Cedrus, 11, 14 subgen. Ducamlpopinus,199
Cercocarpus,229, 236, 245. 253 subgen. Eupinus.71
Clethra,84, 169, 216 subgen. Eupivs, 71
Clusia, 84 subgen. Gerardia, 199
salvinii, 166, 219 subgen. Hatplopinus.199
Comarostaphylis,226 subgen. Haploxylon,4. 30, 31, 33, (36), (42). 199
Comsostrobus,48 subgen. MalacopitYs.199
Cupressaceae,48 subgen. Parrceinhr(pinus, 199
Cupressusguadalupensisvar. forbesii, 184 subgen. Pelce, 198
Cupressus lusitanica,58, 75, 119, 134, 144, 169, 173, 193 subgen. Pintster. 12, 71
Danthoniadomingensis, 99 subgen. Pinea, 71
Dasylirion, 226 subgen. Pinus, 4, 11, 12, 13, 15. 16. 17, 20, 21, 22. 24,
Dendropanaxlempirianus,119 25, 26, 30. 32, 35, 36, 42, 43, 44, 45, 46, 47. 48,
Dodonaea, 139, 238 49. 57, 59. 63, 71, 221
Drymis granadensis, 119 subgen. Sabinianw,71
Duccmnpopinus, 4, 42. 62. 199 subgen. Scleropit\s, 71
Ericaceae, 119 subgen. Strobus. 4. 12. 15, 16, 17. 20, 24, 25, 26. 30.
Eriodictyon,253 33. 35, 36, 42, 43. 44, 46, 47, 48, 49, 63, 65,
Erytheaedulis. 187 198, 221
Fagus mexicana, 216 subgen. Tamilulipasa,199
Fouquieria,229 "div."Tenuisqurnae, 199
splendens, 245 sect. Diploxylon, 12, 22, 71
Fraxinus, 193 sect. Leiophyllae,43, 44, 47, 63
Garrya,236 sect. Lumholtziae,44
Gaultheria, 126, 139 sect. Pinaster, 11. 12. 45. 47
Gochnatia,229 subsect. Attenuatae.43, 45, 46, 47, 63, 64
Heteromeles, 186 subsect. Australes,43, 45, 47, 63. 64
Hybosema, 153 subsect. Contortae,45, 47, 48, 63, 64
Juglans, 84 subsect. Leiophvllae.35, 43
Juniperus,56. 75, 107, 114, 175, 226, 232, 236 subsect. Oocarpae,4, 20, 35, 43, 44, 45, 46, 47, 48, 64
ashei, 244 subsect. Ponderosae,22, 35, 43, 45, 47, 63, 65
californica, 184, 187, 248, 253 subsect. Pseudostrobi,43, 45, 47. 63, 65
deppeana, 105, 173, 238 subsect. SabLiniaae,45, 46
flaccida, 105, 126, 193, 229, 238 sect. Pinea, 11, 44
monosperma, 102, 244 sect. Pinus, 44, 45, 47
standleyi, 119 subsect. Sylvestres,35, 48, 80
Karwinskia,229 sect. Cembra, 11
Larix, 11, 14, 26 sect. Haploxvlon. 12, 22, 198
Lebachiaceae,48 sect. Parryanae,13, 15, 26, 30, (34), 46, 48. 221, 224
Leucaena, 139, 153, 169, 229 subsect. Cembroides, 17, 24, 36, 40, 46, 47, 48, 52, 65.
Leucopitvs,62 224, 226, 246
Leucothoe, 153 subsect. Nelsoniae, 47, 48, 65, 66
Liquidambar.123. 139, 161, 169. 183, 193 subsect. Rzedowskianae,221
styraciflua,57, 146, 216 sect. Pence, 62, 198
Lycopodiaceae, 119 sect. Strobus, 11, 12, 15, 17, 22, 25, 26, 30, (31), 46.
Magnolia, 169, 216 47, 199
Mahonia, 226 subsect. Cembrae,26, 46, 48, 49, 209
Meliaceae, 3 subsect. Strobi, 29, 30, 46, 47. 49. 65, 66, 209

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Index of Scientific Names of Plants and Animals 287

aculcensis, 118 231,232, 238, 239, 240, 241, 243. 244, 246, 247,
albicaulis, 46, 49, 209, 215 250, 254, 255
altamiranoi, 175, 177 subsp. cembroides var. bicolor, 8, 44. 48, 51. 53, 54.
alvordensis,45 232, 233, 234*, 236, 237*, 239, 240, 244, 268
amecaensis, I 18 [Link],44, 48, 51, 54, 232, 233*. 235*. 236,
angulata, 122 238, 239, 240, 241, 268
antoineunu, 131 [Link], 251
apacheca, 112,116 subsp. lagunae, 8, 44, 48, 54, 61, 207*, 232. 236,
apulcensis, 127, 128, 129 237*, 238, 240, 268
arizonica,8. 9, 45, 51, 54, 59, 65, 67, 75, 102, 103, 105, var. lagunae, 236
109, 150, 173, 174, 183, 213, 232 subsp. monophylha,247
var. arizonica,47, 54, 104*, 105, 106*, 107, 109, 267 var. monophylla,247
var. cooperi, 8, 47, 54. 102*, 105. 107, 108*, 109, subsp. orizabensis, 11. 44, 48, 54, 232. 234*. 237*.
267 236, 238, 240. 241, 268
var. stormiae, 8, 47, 54, 102*, 105, 107, 108*, 109, var. orizahbensis,238
193, 244, 267 subsp. parr-ana. 251
attenuata,9, 10, 16, 18, 25, 26, 27, 28, 45, 47, 54, 56, var. parryana, 251
64, 66, 188, 190, 191*, 192*, 195. 267 var. quadrifolia. 251
ayacahuite,6, 21, 28, 30, 31, 34, 35, 36, 44, 46, 54, 58, var. remote, 244, 246
66, 68, 70, 144, 146, 150, 161, 169, 173, 199, chalmaensis, 255
200, 204, 207, 209, 210, 213. 214, 215, 216 chiapensis. 215
var. ayacahuite,48, 54. 200*, 201*, 202*, 203, 267 chihuahuana,75, 76
var. brachlyptera204, 211, 215 coarctana, 133
subsp. loulotiana, 203 coloradensis,44
var. loudo,liana, 203 comonjortii,72
var. novogaliciana, 211, 215 var. gracilis. 72
var. oaxacana, 200),204 var. lerdoi, 73
var. reflexa. 207 var. monte-allegri,72
subsp. stroifornnis, 211 contorta,34, 45, 82
var. strobifornnis,207 var. contorta,255
var. veitchii, 8, 28, 44, 48. 54, 200*, 202*, 203, 204, var. latifolia, 82, 255
267 subsp. murrayana,80
aztecaensis, 133 var. murrayana,8, 18, 45, 47, 51. 54, 58, 64, 66, 80*,
bahlanmensis. 89, 91 81*. 82, 111, 205. 255, 267
balfouriana,12, 199 cooperi. 107, 109
banksiana,34, 45 var. ornelasii, 107
belgica, 48 cornea, 133
besserianca,180 coulteri. 9, 10, 17, 25, 28, 30, 39, 46, 47. 51, 54, 64, 66.
bicolor, 240 195, 196*, 197*, 198*, 267
bonapartea, 211 cubensis, 8, 18, 25, 34, 45. 47. 54, 55, 57. 61, 64. 71.
boothiana, 130 79*, 91, 94, 95*, 96, 97. 99, 267
boucheiani(, 130 var. anoinala, 86. 91
hrachvpterta,100 var. terthrocarpla77, 79
hullata, 133 culminicola, 9. 10, 14, 29, 40, 44, 48, 52, 54. 55, 56.
bungeana,46, 230 58, 61, 65, 68. 209, 213, 224. 240. 241, 242*.
californiarurn,247, 251 243*, 244, 247. 256, 268
subsp. californiarum,251 var. discolor. 234
[Link], 247, 251 [Link]. 234
califjrnica, 188, 190, 192 decaisneana. 133
canariensis,34 var. wilsonii, 116
caribaea, 8, 34, 35, 45, 51, 54, 57, 60, 64, 71, 77, 79, decandolleana, 72
85, 86, 89, 90, 91.93, 94, 96, 97 var. depenlens. 73
var. bahamensis, 8, 16, 47, 49, 54, 86, 87*, 88*, 89, var. ehrenbergii, 116
267 var. verrucosa,72
[Link], 16, 25, 47, 54, 77, 86, 87*, 88*, 91, 267 deflexa. 109
var. hondurensis,8, 9, 18, 40, 47, 49, 50, 51, 54, 55, densa. 91
57, 59, 60, 70, 86, 87*, 88*, 89, 90, 91, 267 var. austrokevsensis,91
carrierei, 133 densiflora,34
catarinae, 244, 247 depauperata,255
cedrus, 73 dependens. 73
cembra, 31, 34, 46, 49, 238, 240 devoniana, 4, 6, 9, 17, 23. 25, 45, 47. 53, 54, 57, 65.
cembroides,5, 6, 11, 12, 14, 26, 29, 33*, 34, 36, 40, 47, 67, 68, 69, 70, 133, 136, 137, 138*, 139, 140*.
53, 54, 56, 58, 59, 61, 65, 67, 68, 69, 75, 107, 141, 142, 144, 146, 153, 158, 179, 267
114, 126, 139, 150, 193,221, 224, 226, 229, 230, discolor. 40, 234, 239, 240, 244

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288 Index of Scientific Names of Plants and Animals

Pinus (continued) hendersonii, 131

doelleriana, 131 herrerae.6. 8, 11, 45, 47, 54, 59, 64, 67, 82, 83*. 85*,
don-pedrii, 200 144, 166, 175, 219, 267
c(onn)ell-smithii.118, 120 heteromorpha,255
douglasiana, 8, 9, 21, 45, 47, 54, 65, 68, 70, 75, 115*, heterophylla,93, 94. 96
142, 143*, 144, 146, 148, 150, 153, 166, 179, hondurensis,89, 91
267 horizontalis, 133
var. martinezii, 171 hoseriana, 144
var. maximlinoi,144 hugelii. 180
durangensis.8. 9, 16, 18, 27, 46, 47, 51, 54, 64, 68, 75, huisquilucaensis,72
107, 114, 120, 171, 172*, 173, 174*, 175, 183, inflexa, 133
213, 267 insignis. 7. 188
to. quinquefoliata, 171, 174 var. hinata, 7, 186, 188
echinata, 34 izltacihuatlii,116
edgariana,!184 jaliscana, 11, 18, 25, 46. 47, 54, 61, 64, 68. 164, 165*.
edulis, 36, 47, 49, 51, 56, 102, 230, 239, 246, 248, 250, 166, 167*, 267
255 jeffreyi. 8. 39, 45, 47, 51, 54, 58, 65, 66, 82, 103. 109,
[Link]., 247, 251 110*, 111*. 112, 198. 205, 248, 253, 256, 267
var. monophylla. 247 var. baja-californica, 109, 112
ehrenbergii, 116 var. deflexa. 109
elegcns., 130 johannis, 40, 234, 239. 240, 244
elliottii, 35, 89, 91, 94 jostii, 131
subsp. densl, 91 juarezensis. 251, 254, 255
var. densa, 14, 22, 35, 45, 47, 50, 51, 54, 57, 64, 71, kesiya, 34
87, 91, 92*, 94, 267 keteleerii, 131
endlicheriana I116 koraiensis.31, 34
var. longifolia. 255 krempfii.4. 11. 17.42. 199
var. richardiatac, 133 lhgunae, 236
engelmannii, 6, 17, 45, 47, 51. 54. 65, 67, 69, 75, 103, lambertiana,8, 16, 24, 25, 30, 31, 44, 46, 48, 51. 54.
105, 112, 113*, 114, 115*, 153, 173, 174, 183, 58, 66, 82, 111, 204, 206*. 207*, 209, 267
213, 232, 267 var. mclrtirensis,204
var. bliancoi, 112, 116 attifolia. 112, 116
escandonianca,144 Iatisquamna, 226, 230
eslavae, 183 lawsoniana, 190
estevezii, 40, 122, 129 lawsonii, 8, 35, 44. 46, 47, 53, 54, 64, 70, 75. 144, 167*,
exserta, 133 175, 176*, 177. 179, 180, 267
filifolia. 137, 141 var. gracilis. 255
flexilis, 35, 204, 207, 209, 210, 211, 215, 230, 243 leiophylla. 6, 12. 14. 17, 23. 26, 34. 35, 36. 37*, 44, 49.
var. flexilis, 209, 211, 256 54. 59, 63, 67, 68, 71, 72. 76, 77, 107, 114, 144,
var. nacrocarpa, 211 150, 153, 161, 173, 175, 183
subsp. reflexa, 207 subsp. chihuahuana,75
var. reflexa, 8, 44, 48, 51, 54, 66. 67, 68, 204, 207, var. chihuahuana,6, 22, 44. 47. 54. 72, 73*, 74*. 75,
208*, 209, 210*, 211. 214, 215, 256, 267 76, 77, 213, 222, 232, 267
var. serrulata, 211 var. leiophylla. 47. 54. 69. 72, 73*, 74*, 75, 76. 267
fremnontiaca,247 lerdoi, 73
gerardiana,34, 46, 199, 224, 230 leucodermis,34
gordoniana, 134, 136 lindgrenii.47
gracilis. 72 lindlevana. I18
grlndis, 130 linguiformis, 192
var. carrierei, 133 llaveana, 232, 238, 240. 251, 253
var. doelleriana, 131 loudoniana, 203
var. haageana. 131 lowii, 16
[Link]. 133 lumholtzii. 5, 6. 17, 35, 44, 45, 47. 49. 54, 65, 67. 76,
var. thelemucnnii, 133 114, 148, 149*, 150, 151*. 158. 213. 267
greggii. 6. 21, 26, 45, 46, 47, 54, 64, 69, 104*, 161, var. rnicrophvlla.73. 76
183, 192, 194*, 195, 267 lutea. 107, 109
grenvilleae. 137. 141 var. ornelasi, 107, 109
hacgeana. 131 mnacrocarpa, 255
halepensis, 15, 34 macrophvylla.112. 116. 137. 141
hamnata,200 var. blancoi. 112, 116
hartwegii, 6, 23, 41, 45, 47, 49, 54. 55, 58, 65, 68, 69, mccvaughii, 164
70, 109, 116, 117*. 118, 119*, 120, 136. 139, naestrensis,94. 97
141. 161, 209, 213, 241. 267 magnifica, 137
var. rudis, 116 magothensis,48

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Index of Scientific Names of Plants and Animals 289

martinezii, 171, 173, 174 nubicola, 123, 129

maximartinezii, 11, 14, 15, 22, 23, 24, 29, 40, 44, 46, oaxacana, 127, 129, 255
47, 48, 54, 56, 59, 61, 65, 67, 210*, 221, 221, var. diversiformis,255
223*, 224, 230, 267 ocampii, 137
maximinoi, 6, 23, 25, 45, 47, 54, 58, 60, 65, 68, 69, 70, occidentalis, 130, 135
139, 144, 145*, 146, 147*, 148, 153, 161, 166, occidentalis, 6, 35, 45, 47, 54, 55, 57, 63, 64, 71. 79*,
169, 171, 179, 216, 267 97, 98*, 99, 100, 267
inavriana, 112, 116 var. baorucoensis, 97, 100
merkusii,34, 44, 221 var. cubensis, 94, 97
nmichoacaensis,137 var. maestrensis,94, 97
michoacana, 9, 137, 141 oocarpa, 6, 8, 23, 26, 32*, 35, 40, 46, 54, 55, 57, 58,
var. cornuta, 137, 141 59, 60, 64, 67, 70, 71, 75, 90, 91, 139, 144. 146.
var. cornuta to. nayaritana, 137 150, 151, 152. 153, 156, 158, 159, 166, 169, 170,
fo. procera, 137 171, 173, 175, 179, 183, 190, 219
var. quevedoi, 137 var. macvaughii, 164
fo. tumida, 137 var. manlzanoi,152, 156
microcarpa, 180 var. microphylla.156, 158, 159
monophylla, 9, 10, 15, 17, 18, 21, 25, 29, 34, 36, 40, var. ochoterenae.91, 156, 164, 166, 167, 170, 171
44, 48, 51, 54, 56, 65, 66, 247, 248, 249*, 250*, var. oocarpa,47, 68, 152, 154*, 155*, 156, 267
251, 253, 254, 255. 268 var. oocarpoides, 152
var. californiarium,247 var. trifoliata, 8, 47, 54, 67, 152, 153*. 154, 155*,
var. iIllax, 247 156, 267
monstsrosa,130 to. trifoliata, 154
var. hendersonii, 131 orizabae, 122
var. nesselrodiacna.133 orizabensis, 238
monte-allegri,72 ortgiesiana, 133
monteziumae,255 osteosperna, 232, 239
montezumae,5, 6, 16, 19*, 35, 40, 45, 53, 54, 55, 57, palustris.34, 139
58, 59, 65, 68, 69, 70, 75, 76, 109, 119, 121, papeleuii, 118
128, 129, 130, 134, 135, 136, 139, 141, 142, 148, parr!ana, 251, 253
161, 173, 174, 175, 183, 193 patula, 148, 150
var. gordoniana,6, 27, 47, 54, 123*, 130, 132*, 134, patula. 6, 8, 21, 34, 40, 45. 46, 54, 58, 59, 60, 65, 69,
136, 137, 267 70, 75, 76, 85, 146, 150, 160, 163, 164. 166. 170,
var. hartwegii, 116, 141 175, 179, 183, 193, 195, 216
var. lindlevana, 118 [Link], 164
var. lindleyi, 130, 135, 136, 141 var. longipedunculata,9, 40, 47. 54, 160. 161*, 162*.
fo. macrocarpa, 133 163, 164, 171, 267
var. macrophylla, 137 var. macrocarpa, 160, 163
var. mezambrana,133,136 var. patula.47. 151*, 160, 161, 162*, 163, 171. 267
var. montezumae,47, 54, 130, 131*, 132*, 134, 141, var. stricta, 180
267 subsp. tecunumanii. 164, 166, 171
var. rudis, 116, 120, 141 ptiawlowkskiana, 137
monticola, 39. 207, 218 paxtonii, 131
mugo, 241 pescatorei. 133
mulleriana, 180 pinaster,34
muricata,9, 10, 26, 28, 45, 54, 55, 56, 186, 188, 192, pinceana, 6. 26, 29, 34, 40, 44, 46, 47, 48, 52, 53, 54,
195 55, 56, 61, 66. 68, 224, 226, 228*, 229, 230*.
subsp. anthonvi, 184 232, 247, 267
var. anthonvi, 184 pinea, 30, 34. 44, 49, 71, 76
var. borealis, 184 planchtonii,133
var. cedrosensis, 186. 188 ponderosa,5, 10, 34, 35. 45, 102, 103. 105. 109. 112,
var. muricata,47, 64, 66, 184, 185*, 188*, 267 116, 253, 256
subsp. remorata, 184 subsp. arizonica, 105
var. remorata, 184 var. arizonica, 105, 174
var. stantonii, 184 subsp. coulteri, 195
murravana,80, 82 [Link], 109
nec-plus-ultra, 137 [Link]!i,103, 109
nelsonii, 7, 8, 12, 14, 17, 18, 22, 23, 24, 26, 29, 34, 40, var. macrophvlla,112. 174
44, 46, 47, 48, 49, 52, 53, 54, 56, 60, 61, 66, 68, var. mariana, 112, 116
199, 224, 225*, 226, 227*, 232, 267 var. ponderosa, 102, 103, 256
nesselrodiana, 133 subsp. scopulortmn,100
nigra, 31, 34 var. scopulorum, 9. 10, 47, 51. 54, 65, 100, 101*.
northumterlandiana,116 102*, 103, 105, 109, 267
novogaliciana, 215 var. stormiae, 107

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290 Index of Scientific Names of Plants and Animals

Pinus (continued) sabiniana,71, 198

popocatepetlii, 203 san-rafaeliana, 131
praetermissa,8. 18, 26, 46. 47. 54, 64, 68, 156, 157*, scoparia, 16
158, 159*, 267 scopulorumn,100, 103
prasina, 255 sibirica, 46
pretuherculata, 190 skinneri, 255
pringlei, 7, 8. 44, 46, 47, 53, 54, 64, 70, 75, 175, 177, soulangeana, 130
178*, 179*, 190, 267 spinosa, 131
protuberaits. 122, 128 standishii, 116
var. angulata,. 122 strobiformis,6, 18, 21, 25, 31, 35, 36, 44, 46, 48, 51,
var. exserta, 133 54, 58, 66, 67, 68, 105, 107, 204, 207, 209, 210.
pseulostrobus, 130 211, 212*, 213, 214*, 215, 256, 267
pseudostrobus,5, 6, 23, 35. 40, 44, 45. 53, 54, 58. 59, var. carvajalii, 211
60, 65, 68, 69, 70, 75, 107, 114, 121, 127, 128, [Link], 211, 215
129, 130, 139, 141, 142, 144, 146, 148, 161, 169, strobus, 10, 21, 31, 34, 148, 199, 207, 209, 216, 218
175, 179, 193, 216, 219, 232, 238, 255, 256 subsp. chiapensis. 215
var. apulcensis. 128, 129 var. chiapensis, 8, 9, 16, 18, 27. 28. 34, 44, 48, 53.
var. apulcensis, 6, 25, 27, 28, 47. 54. 85*. 122. 124*, 54,57,61.66, 69, 70, 169, 214*, 215, 217*, 218,
125*, 127, 255, 267 267
subsp. apulcensis. 123. 128, 129 var. strobus,61
var. coatepecensis, 123, 128 sulhpatula,255
var. estevezii, 122, 128 suffruticosa. 118
var. laubenfelsii, 123 sylvestris, 31. 34, 35, 39, 62, 71, 82
fo. mnegacarpa.123 taeda, 34
var. o)axacana, 127 var. heterophylla.94, 96
fo. protuberans,8, 122, 123*, 125*, 126, 128, 267 tecunumanii,6, 8, 40, 46, 47, 52, 54, 57, 60. 64, 70, 71.
subsp. pseudostrobus, 128 90,91, 146, 152, 153, 164, 166,168*.169*,170,
var. pseudostrobus,47, 122*, 123. 124*, 125*, 127, 171, 216, 267
128, 267 tenangaensis, 130
var. tenuifolia. 144 var. longifolia. 133
pumila, 241 var. ortgiesiana, 133
quadrifolia,6, 7. 27, 29, 34, 40, 44, 48, 51, 54, 56, 66, tenuifolia, 5, 6, 60, 144, 148
230, 247, 248, 250, 251, 252*, 253. 254*, 255, teocote, 6, 35, 44, 46, 47, 54, 58, 59, 65, 67, 69, 70, 75,
268 76, 85, 107, 150. 161, 173, 175, 180, 181*, 182*,
quichensis, 256 183, 193, 267
radiata,26, 34, 45, 60, 61, 188, 190, 192, 194, 195 var. herrerae, 82
fo. binata, 186, 188 fo. mtacrocarpa,180
subsp. binata, 186 var. macrocarpa, 180, 183
var. binata, 7, 8, 47, 50, 54, 61, 64. 66, 184, 186, fo. quinquefoliate, 183
188*, 189*, 190, 256, 267 terthrocarpa,77
var. cedrosensis, 186, 190 thelemannii, 133
fo. guadalupensis, 186, 188 thibaudiana, 131
var. radiata,50, 187, 188, 256 tropicalis.8, 18, 44, 45, 47, 54, 57. 63, 71. 77, 78*, 79*,
recurvata,86, 91 80, 91, 267
reflexa. 204, 207. 215 troubezkoifana,130
regeliana. 122 tuberculata, 190, 192
remorata. 184. 188 var. acutca,190, 192
remota, 9, 44, 48, 51, 54, 66, 68, 226, 230, 244, 245*, turnida,255
246*, 268 tzompoliana. 144
resinosa. 116 valida, 133
resinosa, 34, 80 vangeertii, 131
richardiana, 133 vanhouttei, 131
rigida, 76 veitchii, 203, 204
rinzii, 133 verrucosa,72
robusta, 116 verschaffeltii,137
var. inflexa, 133 vilmoriniana, 180
roezlii. 116 var. besseriana, 180
rohanii, 133 wallichiana,34
roxburghii,34 wilsonii, 116
rubescens, 133 wincesteriana, 137, 141
rudis, 109, 116, 120 wrightii,94, 96, 97
ruimeliana,130 yecorensis, 255
russelliana, 130, 141 var. sinaloensis, 255
rzedowskii, 11, 17, 23, 24, 36, 37*, 44, 46, 47, 48, 54, zacatlanae, 133
61, 65, 67, 210*, 218, 220*, 221, 247, 267 zamoraensis. 255

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Index of Scientific Names of Plants and Animals 291

zitacuarensis, 137 chrysolepis, 197

var. nitida, 137 devia, 236
Pityostrobus,48 macrophylla,222
Platanus, 193 tomentella, 187
mexicana, 216 turbinella,248, 253
Podocarpaceae,48 Rhus, 253
Podocarpus,216 Rubus, 169
Populus tremuloides,82 Sabal palmetto, 89, 90, 93
Prosopis, 229 Salvia, 126, 186
Pseudolarix, 14, 35 Sophora,226, 229
Pseudotsuga,35, 84, 213, 242 Strobus,62
menziesii, 35, 150, 193 Styrax, 183
Pteridium,99 Symplocos, 169
aquilinum,90, 99, 126, 146, 153, 169 Tilia, 84
Quercus, 56, 57, 75, 102, 105, 107, 111, 114, 119, 126, Ulmus mexicana, 216
139, 144, 146, 150, 166, 169, 179, 182, 193,213, Voltziaceae,48
216, 219, 222, 226, 229, 232, 236, 238, 242, Voltziales,48
244 Yucca, 226, 229, 232, 236, 238, 253

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