Historic, archived document 


Do not assume content reflects current 
scientific knowledge, policies, or practices. 


GEOGRAPHIC DISTRIBUTION 


OF THE 


PINES OF THE WORLD 


Elbert L. Little, Jr. 


U.S. DEPARTMENT OF AGRICULTURE MISCELLANEOUS PUBLICATION 99] 
FOREST SERVICE February 1966 


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GEOGRAPHIC DISTRIBUTION OF THE 
PINES OF THE WORLD 


by 
WILLIAM. B. CRITCHFIELD 
Geneticist 
Pacific Southwest Forest and Range Experiment Station 
Forest Service, Berkeley, Calif. 
and 
ELBERT L. LITTLE, JR. 


Dendrologist 
Division of Forest Management Research 


Forest Service, Washington, D.C. 


Miscellaneous Publication 991 


U.S. DEPARTMENT OF AGRICULTURE FOREST SERVICE 
WASHINGTON, D.C. FEBRUARY 1966 


ACKNOWLEDGMENT 


The authors owe a special debt of gratitude to Dr. Nicholas T. Mirov, 
who began this project before his retirement from the U.S. Forest Service 
in 1961. His many specific contributions, particularly to the information 
about the eastern and southeastern Asian species, are acknowledged in the 
text. Dr. Robert Z. Callaham has been helpful and encouraging through- 
out the preparation of this publication. Many persons have provided 
valuable information about pine distribution, and their contributions are 
acknowledged in the text under the appropriate species. 

For their work in compiling and drafting the maps, the authors 
are greatly indebted to Audrey E. Kursinski and Barry K. Ford. 


Il 


Library of Congress Catalog Card Number: Agr 65-212 


For sale by the Superintendent of Documents, U.S. Government Printing Office 
Washington, D.C., 20402 - Price 75 cents 


CONTENTS 


Species distribution, genus Pinus ___ 


Subgen. Ducampopinus __ 
Cc 


Sect. Ducampopinus 
Subsect. Krempfiani 


Pinus krempfii 
Subgen. Strobus _ 


Sect. Strobus 


P. koraiensis 


P. pumila 


P. sibirica 


P. cembra 


IPs GUROIOWG 2 


iPAEmMOonticolay se oe 
. lambertiana 


. flexilis 


ayacahuite 
peuce 


armandii ___ 


griffithii 


. dalatensis 
parviflora 
. morrisonicola 


. fenzeliana _ 


saa a Ma. a a aaa a) 


. wangil 
Sect. Parrya 
Subsect. Cembroides 
P. cembroides 
. edulis 
. quadrifolia 
. monophylla 
. culminicola 


. pinceana 
. nelsonii 


my y 


P. gerardiana 
P. bungeana 

Subsect. Balfourianae 
P. balfouriana 


P. aristata 
Subgen. Pinus 
Sect. Ternatae 


Subsect. Leiophyllae 
P. leiophylla 
P. lumholtzii 


9 


r 


DAN DM OF Or Ot Ot Ot Ot St OF OT OT OT OT Ot 


a STS SS) 


C0 00 00 CO ~I ~I 


2a 


Map 


2b (11) 


3 


16, 


(4) 


CONTENTS 


Subsect. Canarienses __ een Bl Ne av ea 
IRANGamlansien suse seeaa iee eens Pople a rena aun tyes 
P. roxburghii fan Ala Se ese Aer che ara oe WIR MRE AES Telok tet 


Subsect. Pineae 


Ie 


Sect. Pinus 


Les 


Sub 


to gg) Ine) tae}ine} oe] 


Su 


1 
P 


. nigra 


. sylvestris 


. taiwanensis 
. luchuensis 
. hwangshanensi 


ae) Ing) Ine} tae} toe} Inch Ine) nsf ing} Inel lech Inch tne}! Ine} toe), las} tas) ine} 


. merkusii 


. palustris 
. taeda 


ay ae Madi 


. elliotti 
. caribaea 
. occidentalis 
. cubensis 


. ponderosa 
. washoensis 


. Montezumae 


. hartwegii 


. douglasiana 
. teocote 


a Iia~ la-la-la-la Iilg- la-la-la-la la~ia~ 


. coulteri 


Sy LITE aa aa 


Subsect. Sylvestres ____ 
TESUTNOS aly eet WM utara ale are en Ed Lae ee 


. tropicalis 


. heldreichii 


pinaster 
. halepensis 
bauititaeses 


PATTNUUG ©) iycoet teat Siac fe aegis es 4 elias preys 78 


densiflora 


. thunbergiana 
. massoniana 


. tabulaeformis 
. yunnanensis 
. Insularis 


Sr) Shae! 


sect. Australes 


. glabra 


rigida 


serotina 
. pungens 


sect. Ponderosae 


jeflreyi 


. engelmannii _ 


. durangensis 


cooperi 


. michoacana 


. pseudostrobus 


. sabiniana 


Dy we 


38 (37) 
39 


CONTENTS 


Page 

EPRLOULEN Ada teen, ete Meine Ls es! Soe ee eee 18 
Sribsecie GOntontac ues = same ne ee ek is 19 
Pet oe keeyneh veal} ee 2 ae ee 19 
ARC OUCOLUameeees See een. ge ea Bs ee ee it) 
Ee pva Oui ak eee tee eS en See ee 19 
JP QTC 2) ce SS ae Ss Pa ek get 19 
Subsectus Oocanpicpes = 1m. eee Se Sa Ae ee 19 
1e2, TeEMOUTE NE, at eee aes ee eee ere 19 
AALCe IAC mnie Mer ON St be ee ee S| 19 
PASTA Cat aweeeens eect ee WR el eS 20 
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AO OCALD Ames Stee on 2 oy pe ne A UE ee 20 
PA Gti cinerea ek Ah on BB oo EAR 20 
WiferatinecaGl(CUume a tae Se Se ee ee 21 


NIB ON: Seeks SS Sane melo Se SL en pe on oe le 30 


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GEOGRAPHIC DISTRIBUTION OF THE 
PINES OF THE WORLD 


INTRODUCTION 


The genus Pinus is one of the most widely distributed genera of trees in the 
Northern Hemisphere. In the North Temperate Zone it is rivalled only by 
Quercus, which has more species and may occupy a greater area. The pines 
extend from the polar region to the tropics, crossing the Equator at one point 
(Map 1). Within this enormous range, they dominate the natural vegetation 
in many regions. The genus includes some of the most valuable timber trees in 
the world, and is readily distinguished from other plants. For all of these 
reasons a great deal of information has accumulated about the natural dis- 
tribution of the pines. This publication summarizes this information in the 
form of maps, and indicates the sources of information wherever possible. 

A previous attempt to summarize information on the distribution of all 
the pines is that of Schmucker (1942), who included them in his survey of 
the distribution of the trees of the North Temperate Zone. His diagrammatic 
maps are small in scale (1:50,000,000 to 1:80,000,000) , and his principal sources 
of information are indicated only in a general way. 

The scope of this publication is limited to mappable information about 
where pines grow naturally at present. We have not tried to relate present 
distribution to the fossil record, nor have we tried to indicate the ecological and 
historical factors controlling species distribution. The elevations at which pines 
grow are mentioned only in the most general terms; this aspect of distribution 
is intimately related to latitude, topography, and other features of the local 
environment. 

In relation to the evolution of Pinus, the arrangement of species maps has 
been made as meaningful as possible by grouping the species according to their 
presumed relationships. The classification of the soft or white pines, subgenus 
Strobus (Haploxylon), follows Shaw (1914, 1924). The hard pines, subgenus 
Pinus (Diploxylon), are arranged according to the relationship scheme of Duffield 
(1952) , except that we have retained Sabinianae (group Macrocarpae of Shaw) 
as distinct. 

We have brought up to date the names of the subdivisions of the genus 
Pinus, as explained in a separate report.! As indicated under Contents, the genus 
Pinus is divided here into 3 subgenera, 5 sections, 15 subsections, and 94 species. 
The rank subsection corresponds to that designated by some authors as series, 
“group,” or section. One subgeneric name by another author is validly published 
here, and three names of subsections are proposed as new. 

The geographic distribution or natural range of 94 species of Pinus (pine) 
is summarized and presented on 64 maps. Several other names are mentioned 
or combired as synonyms following conservative usage. English common names 
have been added. 

Varieties and subspecies have not been mapped separately except in a few 
special cases. Many taxa of these ranks have been established on fragmentary 
evidence in Pinus, and cannot be considered mappable entities because of un- 
certainties or disagreement about their characterization or distribution. 


1 Little, Elbert L., Jr., and Critchfield, William B. Subdivisions of the genus Pinus. 1965 
(Unpublished report on file at U.S. Forest Service, Washington, D.C.) 


SOURCES OF INFORMATION 


Forest trees of economic importance differ from most 
other plants in the very diverse kinds of information that can 
be used in mapping them. Publications for tree identifica- 
tion, floras, vegetation maps, forest-type maps, and a sizable 
part of the literature of forestry and plant ecology are 
sources. ‘Travelers’ accounts can occasionally be used, and 
zoological studies are sometimes helpful—for example, a 
report by J. I. Marshall (1957) on the birds of the pine-oak 
woodland of northern Mexico is an important source of 
information about pine species distribution in that region. 
Dot maps based on herbarium specimens convey only a frac- 
tion of the available information, and we have used them 
only for the pines of Mexico. The main source of distribu- 
tion information about these Mexican species is Martinez 
(1948), who lists the localities where each species was col- 
lected. For the other pines, we have incorporated the 
available information into maps showing their total distri- 
bution, often with a special symbol (x) to indicate isolated 
or scattered occurrences. 

The amount and quality of published information about 
where pines grow varies greatly from species to species and 
from country to country. The distribution of such economi- 
cally important species as Pinus ponderosa, P. sylvestris, and 
P, pinaster is usually better known than that of species hay- 
ing little or no economic importance, such as P. albicaulis, 
P. mugo, and P. pumila. 

The countries of eastern Asia present the greatest ex- 
tremes in available information: The pines of Japan have 
all been mapped in detail at a large scale (Hayashi 1952, 
1954), whereas the natural distribution and even the identity 
of some of the pines of eastern and northeastern China are 
uncertain. More is known about the distribution of pines in 
some of the remotest parts of western China than in eastern 
and southern China because of the remarkably concentrated 
plant exploration of parts of Kansu, Szechuan, Sikang, and 
Yunnan provinces (Cox 1945). In western and southern 
Europe, the distribution of the pines of Switzerland and 
Yugoslavia is more fully documented than that of most other 
countries. In the Western Hemisphere, poorly known areas 
include northwestern Canada, parts of the Great Basin and 
adjacent regions in western United States, parts of Mexico, 
and much of Honduras. 

We have supplemented the published sources of informa- 
tion by having preliminary versions of all of the maps re- 
viewed by specialists. “The European and American species 
received a more thorough review than the Asian species. 
Information obtained from reviewers and cited here is filed 
at the Institute of Forest Genetics, Placerville, Calif. Addi- 
tional working maps of the pines of the United States are in 
the files of the Forest Service dendrology project at Wash- 
ington, D.C, 

With certain exceptions, all of the published and unpub- 
lished sources of information are cited with brief notes on 
each species. Where a source pertains to only one or two 
countries, states, or provinces, these are indicated in paren- 
theses. ‘The following abbreviations are used for countries, 
states, and provinces: 


2 


Countries: 


Union of Soviet Socialist Republics 


United States of America___ 


States (U.S.): 
Arizona 


California 


Colorado 
Florida 


Illinois 


Indiana 


Minnesota 


Missouri 


Montana 


Nebraska 
Nevada) Baebes Remit sine 


New Hampshire 
New Mexico_ 


New York_ 


North Carolina 


North Dakota 


Oregon 
Pennsylvania 
South Carolina 


Washington 


Wyoming 
Provinces (Canada): 
Alberta 


British Columbia__ 


Newfoundland 


Nova Scotia 
Ontario Zoe See eee 


Owebec Lee Ss zs 
Q) 


States (Mexico): 
Aguascalientes 
Baja California 


Chika a's) yp 3 cotillion ya 


Chihuahua 


Coahuila 


Durango 


Guerrero 


Hidalgo 


Jalisco 


México 


Nayarit Woe MAUS Soe OS 
Nuevo Leon 


Oaxaca 


Puebla 


@UEKE tao; sees eae ee 


San Luis Potosi_ 
Sinaloa 


Sonora 
Tamaulipas 


Veracruz __ 


Zacatecas 


Abbreviation 


Ind. 
Minn. 
Mo. 
Mont. 
Nebr. 
Nev. 
N.H. 
N. Mex. 
INSYS 
N.C. 

N. Dak. 
Oreg. 
yas 

S.C. 
Wash. 
Wyo. 


Alta. 
BAG 
N.F. 
N.S. 
Ont 


Que. 


Ags. 
Baja 
Calif. 
Chis. 
Chih. 
Coah. 
Dgo. 
Gro. 
Hgo. 
Jal. 
Mex. 
Mich. 
Nay. 
N.L. 
Oax. 
Pue. 
Qro. 
Sues 
Sin. 
Son. 
‘Tamps. 
Ver. 
Zac. 


4 


All of the sources used for the distribution maps of the 
European and Asian pines are listed with the species notes. 
Additional source material, not listed with species notes, was 
used for the pines of the United States, for those of Califor- 
nia in particular, and for the pines of Canada and Mexico. 

United States.—Sargent (1884) was perhaps the first to 
record in detail the distribution of the native trees of the 
United States, including the genus Pinus, and to publish 
distribution maps of a few native pines. Afterwards he pub- 
lished revised ranges without maps in his Silva (1891-1902) 
and Manual (1905, 1922, 1926). Mohr (1897) prepared 
maps of the species of Pinus in southeastern United States. 

Early work by the Forest Service on tree distribution 
maps has been reviewed by Little (1951). In charge of this 
work was George B. Sudworth, dendrologist with the former 
Bureau of Forestry from 1886 to 1905 and with the Forest 
Service thereafter until his death in 1927. Sudworth sum- 
marized ranges in his Check Lists (1898, 1927) and cited 
detailed distribution records in his Forest Trees of the Pacific 
Slope (1908). He and his assistants prepared distribution 
maps of most native tree species for publication under the 
title, “Forest Atlas Geographic Distribution of North Amer- 
ican Trees.” Only Part I—Pines (Sudworth 1913), contain- 
ing maps for 36 species of Pinus, appeared. Some smaller 
maps were issued in five bulletins on Rocky Mountain trees: 
one of them covered the pines (Sudworth 1917) . 

Munns (1938) published 170 maps of important forest 
trees of the United States, including 26 of pines, revised from 
Sudworth’s older maps. Many of these maps were redrafted 
on a smaller scale and often with slight revisions in the Forest 
Service leaflets known as the American Woods series, several 
on pines, by Betts (1945). New small maps of 165 species 
including 20 pines were made for the 1949 Agriculture Year- 
book, Trees (Little 1949). An atlas showing the distribution 
of the native tree species is in preparation (Little 1951). 

The revised maps of the 36 species of Pinus native in the 
United States published here have incorporated with  re- 
visions the Forest Service maps prepared for other publica- 
tions. These include maps of 18 species for the book, “‘Silvics 
of Forest Trees of the United States” (Fowells 1965), and 
maps for the forthcoming atlas. 

In compiling these maps, the earlier Forest Service maps 
and other references mentioned above have been consulted, 
and the sources for the small maps in the 1949 Yearbook 
have been rechecked (Little 1951). 

Current State floras (Blake and Atwood 1942; Gunn 
1956) and tree books (Dayton 1952), several with maps by 
species and detailed range notes, have been important sources 
for plotting distribution on a large base map of the United 
States showing county boundaries. Some local floras have 
been checked, especially to fill in any gaps in State publica- 
tions. Regional floras and tree publications have been helpful 
also. Maps of several native tree species, including a few 
pines, have appeared in monographic studies. Some general 
references for tree identification contain small maps. 

Several State herbaria possess unpublished maps for each 
species showing by dots the counties or localities of the speci- 
mens collected. Credit is due the curators for permission to 
use this unpublished information. Where other records 
seemed inadequate, some herbarium specimens have been 
examined, for example, in a few States without species maps 
and for uncommon species of imperfectly known range. 

The Forest Survey of the Forest Service has provided 
much detailed information on the distribution of commercial 
tree species. From measurements of trees on thousands of 
uniformly spaced sample plots within a State, highly accurate 
maps have been compiled, especially for the southeastern 
species (Janssen and Weiland 1960). States with distribution 
maps of commercial tree species published by the Forest Sur- 
vey include: Virginia (Evans 1942), North Carolina (Roberts 
and Cruikshank 1941a), South Carolina (Roberts and Cruik- 
shank 1941b), and Mississippi (Sternitzke and Duerr 1950). 


Similar unpublished records of pines and other conifers by 

counties for other Southeastern States have been consulted. 

Some very early western forest survey reports contained 
species maps not duplicated later. Forest type maps prepared 
mostly by the Forest Service have been published by States or 
regions. These have been helpful in mapping limits of 
dominant tree species, associated trees, and nonforested areas, 
particularly in western States of great altitudinal range and 
vegetation zonation. Vegetation maps have been prepared 
for some States, and one of the United States by Zon and 
Shantz (1924). 

Accuracy of the latest maps has been improved through 
review by many foresters, both within the Forest Service and 
outside, and by botanists. Their assistance is acknowledged. 

Principal ‘sources, published and unpublished, for the 
maps of the 36 species of Pinus native in the United States are 
summarized below by States. California, with additional spe- 
cial sources, is discussed separately. Pinus is not native in 
Hawaii, Kansas, or Puerto Rico. 

Alabama.—Harper (1928); Mohr (1901). ; 

Alaska—Hultén (1941); Sudworth (1908) ; Taylor and Little 
(1950). 

Arizona.—Kearney and Peebles (1960); Little (1950). Un- 
published maps by Robert A. Darrow at University of 
Arizona Herbarium. 

Arkansas.—Moore (1960). Unpublished maps by Dwight M. 
Moore at University of Arkansas Herbarium. 

California—Jepson (1910); Munz and Keck (1963); Sud- 
worth (1908). Special sources are cited on page 4. 

Colorado.—Cary (1911); Harrington (1954). Unpublished 
maps by H. D. Harrington at Colorado State University 
Herbarium. 

Connecticut.—Graves et al. (1910) ; Taylor (1915). 

Delaware.—Taber (1960); Tatnall (1946). 

District of Columbia.—Hermann_ (1946); 
Standley (1919). _ 

Florida—Kurz and Godfrey (1962); Ward (1963); West and 
Arnold (1956). 

Georgia.—Bishop (1948); Harper (1907-08); Thorne (1954) . 
Unpublished maps by Wilbur H. Duncan at University of 
Georgia Herbarium. 

Idaho.—Davis (1952); Johnson (1961) ; 
Merriam (1891); St. John (1963). 

Hlinois—Jones and Fuller (1955); Miller and Tehon (1929). 

Indiana.—Deam (1940); Deam and Shaw (1953). 

Towa.—Aikman and Hayden (1938); Campbell (1956). 

Kentucky.—Braun (1943): Garman (1913). Unpublished 
maps by Edward T. Browne, Jr., at University of Kentucky 
Herbarium. 

Louisiana.—Brown (1945); Cocks (1921). 

Maine.—Hyland and Steinmetz (1944); Ogden, Steinmetz, 
and Hyland (1948). 

Maryland.—Shreve et al. (1910); Tatnall (1946). Examina- 
tion by Russell G. Brown of specimens at University of 
Maryland Herbarium. 

Massachusetts—Emerson (1875); Illick (1927). 

Michigan —Otis (1931): N. Smith (1952). Unpublished 
maps by John W. Andresen from specimens at University 
of Michigan Herbarium, Michigan State University Her- 
barium, and other herbaria. 

Minnesota.—Rosendahl and Butters (1928). Specimens ex- 
amined at University of Minnesota Herbarium. 

Mississippi—Lowe (1921); Sternitzke and Duerr (1950). 


Missouri.—Settergren and McDermott (1962); Steyermark 
(1963). 


Hitchcock and 


Kirkwood (1930); 


Montana.—Booth (1950); Hutchison and Kemp (1952); Kirk- 
wood (1930); Rydberg (1900); Standley (1921) 

Nebraska.—Pool (1951). 

Nevada.—Billings (1954): Linsdale, Howell, and Linsdale 


(1952): Little (1956). 
New Hampshire.—Foster (1941) 


w 


me (1911); Taylor (1915). 
ttle (1950); Wooton and Standley (1915). 
‘own (1921); Taylor (1915). Unpublished 
. Ogden and Stanley Smith at New York State 
\Ibany. 
ina.—Coker and Totten (1945); Pinchot and 
897); Roberts and Cruikshank (1941a). Unpub- 
ed maps by Albert E. Radford, Harry E. Ahles, and C. 
itchie Beil at University of North Carolina Herbarium. 
or th Dakota .—Stevens (1950). 
i aun (1961); Transeau and Williams (1929). 
ims jee hie Gibbs, and Mattoon (1939). Specimens 
exa ayned at University of Oklahoma Herbarium. 
yregon:—Benson. (1930); Peck (1961); Randall (1957); Sud- 
worth (1908). 

Pennsylvania.—Illick (1925 Unpublished maps by John 
M. Fogg, Jr., Edgar T. The: and others at University of 
Pennsylvania Herbarium. 

Rhode Island.—Russell (1900) . 

South Carolina~—Coker and Totten (1945); Roberts and 
Cruikshank (1941b). Unpublished maps by Albert E. 
Radford, Harry E. Ahles, and C. Ritchie Bell at University 
of North Carolina Herbarium. 

South Dakota.—McIntosh (1949); Over (1932); Ware (1936). 

Tennessee.—Shanks (1952). Unpublished maps by Aaron J. 
Sharp and Royal E. Shanks at University of ‘Tennessee 
Herbarium. 

Texas.—Cory and Parks (1937) ; 
and Sperry (1951) ; 
Arendale (1953). 

Utah.—Graham (1937). Unpublished thesis: Erdman, Kim- 
ball S. Classification and distribution of the native trees of 
Utah. 221 pp., illus. Brigham Young Univ., M. S. thesis. 
1961. 

Vermont.—Burns and Otis (1916); Dole (1937). 


Gould (1962) ; McDougall 
‘Texas Forest Service (1943); Vines and 


Virginia.—Evans (1942); Massey (1961). 
Washington. palace (1936); Mosher and Lunnum (1951) ; 
Piper (1906); St. John (1963); Sudworth (1908). Speci- 


mens ee at University of Washington Herbarium. 

West Virginia—Brooks (1920); Strausbaugh and Core 
(1952). Unpublished maps by Earl L. Core and Elizabeth 
Ann Bartholomew at University of West Virginia Herbar- 
ium. 

Wisconsin.—Fassett (1930). Unpublished maps by John W. 
Andresen from specimens at University of Wisconsin Her- 
barium and other herbaria. 

Wyoming.—Cary (1917); McDougall and 
Porter (1959). 

California.—Our principal sources of information for all 
of the pines that occur in California and adjacent parts of 
Nevada are the Vegetation Type Maps* and Soil-Vegetation 
Maps* prepared by the Pacific Southwest Forest and Range 


3agegley (1956); 


*Vegetation type and forest condition maps of California and western 
Nevada. September 20, 1963. U.S. Forest Serv., Pacific Southwest Forest 
and Range Expt. Sta., Berkeley, Calif. [List of available maps.] 

*Timber stand and soil-vegetation maps of California. January 1963. 
U.S. Forest Serv., Pacific Southwest Forest and Range Expt. Sta., Rerkeley, 
Calif. [List of available maps.] 


Experiment Station in cooperation with other agencies. We 
have also used the more detailed, unpublished information on 
which the older Vegetation Type Maps were based, Soil- 
Vegetation Maps currently being prepared for publication, 
data from the Forest Survey, and other unpublished informa- 
tion of the Pacific Southwest Forest and Range Experiment 
Station. 

Canada.—Similar published references have been con- 
sulted for the maps of the eight species of Pinus native in 
Canada. Small distribution maps have appeared in the pub- 
lication, Native Trees of Canada (Canada Department of 
Forestry 1961). Halliday and Brown (1943) mapped the 
distribution of important forest trees from forest survey data, 
and Rowe (1959) mapped and described the forest types of 
Canada. Principal additional published sources, including 
floras and tree and forestry publications by Provinces, are: 

Alberta.—Moss (1959) 

British Columbia.—Garman (1963) ; 

(1918). 

Manitoba—Lowe (1943); Scoggan (1957). 

New Brunswick.—Morison (1938); Roland (1947). 

Newfoundland.—Rouleau (1956). 

Nova Scotia.—Roland (1947). 

Ontario.—White and Hosie (1946) . 

Quebec.—Hustich (1949); Marie-Victorin (1927, 1935). 

Saskatchewan.—Budd (1957) ; Fraser and Russell (1954). 

Yukon.—Hultén (1941); Porsild (1951) . 

Mexico.—The mapping of these pines differs from that 
of the other pines because of the scarcity of distribution in- 
formation other than the small-scale distribution maps and 
the lists of collection localities in Martinez (1948). His lists 
are the chief source of information for the Mexican distribu- 
tion of these species (the majority of the pines that occur in 
Mexico) : Pinus strobus, P. strobiformis (as P. ayacahuite var. 
brachyptera and P. reflexa), P. ayacahuite, P. cembroides, P. 
pinceana, P. nelsoni, P. leiophylla (including P. chihuahu- 
ana), P. lumholtzi, P. ponderosa (as P. arizonica), P. engel- 
mannii, P. durangensis, P. cooperi (as P. lutea), P. monte- 
zumae, P. hartwegii (including P. rudis), P. michoacana, P. 
psceudostrobus (including P. tenwifolia), P. douglasiana, P. 
teocote (including P. herrerai), P. lawsonii, P. patula, P. 
gregen, P. oocarpa, and P. pringlet. 

Many of the place names listed by Martinez could not be 
located on the available maps. Most of the others were lo- 
cated with the assistance of a recent gazetteer (U.S. Board 
on Geographic Names 1956). 

Another source of distribution information for many of 
the Mexican pines was the Mexican pine collection in the 
herbarium of the Institute of Forest Genetics at Placerville, 
Calif. Included in it are the 1960 and 1963 collections of 
E. L. Little, Jr., and the 1962 collections of North Carolina 
State College. We also used the species distribution map in 
the Guidebook to the Seminar and Study Tour of Latin 
American Conifers (Mexico Instituto Nacional de Investiga- 
ciones Forestales 1960). The approximate limits of the 
species were drawn with the help of topographic maps and 
several forest-type maps covering part or all of Mexico (Com- 
ision Forestal del Estado Michoacan 1958: Mexico Instituto 
Nacional de Investigaciones Forestales 1960; Leopold 1950). 


Whitford and Craig 


SPECIES DISTRIBUTION, GENUS PINUS L. 


SUBGEN. DUCAMPOPINUS (A. Cheval.) 


de Ferré 
SECT. DUCAMPOPINUS 
SUBSECT. KREMPFIANI Little & Critehfield’ 


Pinus krempfii Lecomte MAP 2a 
Pinus krempfii Lecomte, Paris Mus. Natl. d’Hist. 
Nat. Bull 27: 191, fig. 1921: 

This unique species—the only pine with flattened needles 
—is known only from the high mountains of southern Viet- 
nam. 

Sources: 
Published—Bui 1962; Rollet 1955. 
Unpublished—M. Schmid to N. T. Mirov 1961. 


SUBGEN. STROBUS Lemm. 
SECT. STROBUS 
SUBSECT. CEMBRAE Loud. 


Pinus koraiensis Sieb. & Zuce. MAP 2b 
Korean pine (see also Map 11) 


Pinus koraiensis Sieb. & Zucc., Fl. Jap. 2: 28, 
t. 116, fig. 5-6. 1844; exclud. fig. 1-4. 


Korean pine ranges through Korea and eastern Man- 
churia into southeastern Siberia, with outliers on the Japanese 
islands of Honshu and Shikoku. 

Sources: 

Published—Hayashi 1952 (Japan); Solovev 1958 

(U.S.S.R.); Uyeki 1926 (Korea); Wu 1956 (Manchuria). 


Pinus pumila Regel MAP 3 
Japanese stone pine (see also Map 4) 


Pinus pumila Regel, Index Sem. Hort. Petrop. 
1858: 23. 1859. 


This pine ranges widely through northwestern Asia, ex- 
tending north almost to the Arctic Ocean, east to the Bering 
Sea, west to northern Mongolia and Lake Baikal, and south 
to Korea and central Honshu, Japan. In the southern part 
of its range this shrubby, thicket-forming species is restricted 
to high elevations. Its presence on the Commander Islands 
east of Kamchatka is debatable; Tikhomirov (1949) shows it 
on one of the islands (Mednyy), but Tatewaki (1958) states 
that it is not present on either island. 


' Pinus subgen. Ducampopinus (A. Cheval. ) de Ferre, subgen. nov. 
Pinus sous-genre Ducampopinus (A. Cheval.) de Ferré, Paris Acad. Sci. 
Compt. Rend. 236: 228. 1953 (not validly published under [CBN Art. 33 
because reference to basionym lacked original publication with page) . 
Ducampopinus A. Cheval., Rev. Bot. Appl. d’Agr. Trop, 24: 30. 1944. 
Holotype species: Pinus krempfit Lecomte, Paris Mus. Natl, Hist. Bul. 27: 
LOE tiga O21 


5 Pinus subgen. Ducampopinus sec, Ducampopinus subsect. Kremp/fiani 
Little & Critchfield, subsect. nov. Characteribus subgeneris Pinus subgen. 
Ducampopinus (A, Cheval.) de Ferré. Folio cum 1 fasciculo vasculare, 2 


In the past this species has generally been considered a 
variant of the Pinus cembra—P. sibirica group of eastern Si- 
beria and Europe, but recent investigators consider it more 
closely allied to the Japanese P. parviflora (de Ferré 1960; 
Malyshev 1960) . 

Sources: 

Published—Hultén 1926; Imanishi 1950 (Manchuria) ; 
Kung 1934 (Manchuria); Malyshev 1960 (Siberia); Takahasi 
1944 (Manchuria); Tatewaki 1958 (Kurile Islands); Tik- 
homirov 1949; Utkin 1961 (Siberia); Uyeki 1926 (Korea) ; 
Wu 1956 (Manchuria) . 


Pinus sibirica Du Tour MAP 4 


Siberian stone pine 
Pinus sibirica Du Tour, Nouv. Dict. Hist. Nat. 18: 18, 1803. 


Siberian stone pine ranges from the Ural Mountains 
through western and central Siberia to northern Mongolia, 
with outliers on the Kola Peninsula of northern Russia. This 
species is the Siberian counterpart of the central European 
Pinus cembra. Its range overlaps that of P. pumila in the 
region of Lake Baikal, Siberia. P. stbirica sometimes assumes 
a shrubby growth form like that of P. pumila, and the two 
are often confused (Malyshev 1960). 

Sources: 

Published—Hultén 1950 (Kola Peninsula); Lavrenko and 

Soczava 1956; Nekrasova 1949 (Kola Peninsula) . 


Pinus cembra L. MAP 4 


Swiss stone pine 
Pinus cembra L., Sp. Pl. 1000. 1753. 


Swiss stone pine grows at high elevations in the Alps and 
the Carpathian Mountains. It is closely related to Primus st- 
birica, from which it is separated by a 1,500-mile gap. 
Sources: 

Published—Fekete and Blattny 1913-1914; Generali 1937 
(Italy); Marchesoni 1959 (Italy); Ozenda 1961 (France) ; 
Rubner 1953, p. 364; Scharfetter 1938; Schmid 1949 (Switzer- 
land) . 


Unpublished—Forestry Research Institute, Rumania, 
1964; M. Vidakovié 1964 (Yugoslavia). 
Pinus albicaulis Engelm. MAP 5 


whitebark pine 
Pinus albicaulis Engelm., Acad. Sci. St. Louis Trans. 
2: 209. 1863. 
Whitebark pine occurs at high elevations from central 
British Columbia south to the southern Sierra Nevada olf 


in fasciculo, anguste lanceolata, valde complanata (15-4 mm. late 
minute serrulata vel integra, stomatibus ventralibus atque raro dorsalibus 
ductis resiniferis subexternis, et vagina mox decidua. Bases bractearum 
non decurrentes. Strobilus junioris solitarius, sine aculeis. Strobili ovoid 


symmetricales, aperti postquam maturi. Squamae carinatae, apophysi 
crassa pyramidata et umbone dorsali. Semina cum ala longa articulata 
Lignum modice durum, leviter resinosum, annulis incrementi, sinc 
tracheidiis radiis, alburno albo luteolo, ligno interiore col monis. A 


subgeneribus aliis differt follis complanatis atque absen tracheidiarum 
radiis. Holotypus: Pinus kremp/it Lecomte, Paris Mus. Nz Hist. Nat 
Bul. 27: 191, fig 1921. 


or 


> Ruby Mountains of Nevada, and the moun- 

western Wyoming. Through much of its range 

ristic of timberline, where it often forms dense 

is distribution is still not fully known. Many 

it grows are relatively inaccessible, and in the 

id southeastern part of its range it is often confused 

rus flexilis. The cones of the two species differ, but 

yecause of similarities in most other features there is con- 

siderable uncertainty about their relative distribution in areas 

of overlap. 

\ ditional sources: 

Published—Ayres 1900a (Mont.), 1900b (Mont.) ; Bailey 
and Bailey 1941; Bedwell and Childs 1943 (Wash.) ; Leiberg 
1897 (Idaho), 1900b (Oreg.), 1900c (Idaho), 1904a (Mont.) 
1904b (Mont.) . 

File report—‘Blister rust occurrence survey in the Mar- 
ble Mountains, Klamath National Forest’ by Benton How- 
ard, October 4, 1956. Forest Service, Region 5, San Francisco, 
Calit. 

Unpublished—T. C. Brayshaw and A. E. Porsild 1964 
(B.C.); J. F. Franklin 1963 (Oreg. and Wash.); J. R. Grif- 
fin 1963 (Calif.); F. D. Johnson 1963, 1964 (Idaho); O. V. 
Matthews 1964 (Calif.); H. Weaver 1963 (Wash.) . 


’ 


SUBSECT. STROBI Loud. 


Pinus strobus L. MAP 6 


eastern white pine 
Pinus strobus L., Sp. Pl. 1001. 1753. 


Eastern white pine ranges from Newfoundland through 
southern Canada to southeastern Manitoba, through the Lake 
States south to Iowa and Illinois, and throughout much of 
northeastern United States, south in the Appalachian Moun- 
tains to northern Georgia. A discontinuity of more than 
1,200 miles separates it from a southern variant usually called 
Pinus strobus var. chiapensis Martinez. This Middle Ameri- 
can white pine grows at low and middle elevations in south- 
ern Mexico and Guatemala. 

Additional sources: 

Published—Aguilar 1961 (Guatemala) ; Crespo 1963 
(Pue.); Haddow 1948a and 1948b (Ont.); Miranda and 
Sharp 1950; Sharp 1946 (Guatemala); Standley and Steyer- 
mark 1958 (Guatemala); Wagner 1962 (Chis.) . 

Unpublished—T. C. Brayshaw and A. E. 
(Canada); B. Hallberg 1964 (Oax.). 


Porsild 1964 


Pinus monticola Dougl. MAP 6 


western white pine 
Pinus monticola Dougl. ex D. Don in Lamb., Descr. 
Genus Pinus. Ed. 3 (8°), v. 2, unnumbered page between 
paletand pe lao 832. 

Western white pine ranges from southern British Colum- 
bia to northern Idaho, northwestern Montana, and eastern 
Oregon, and south through the Cascade Mountains to the 
southern end of the Sierra Nevada, Calif. It is separated from 
the closely related Pinus strobus by more than 1,000 miles. In 
the northern part of its range P. monticola nearly reaches sea 
level in places, but in the Sierra Nevada it is a high-elevation 
tree. Early reports of this species in the mountains of south- 

o California (Sudworth 1908) apparently were the result 
mnfusion with either P. flexilis or P. lambertiana. 


Additional sources: 

Published—Ayres 1900a and 1900b (Mont.); Baker 1951 
(Oreg.); Leiberg 1900a (Mont.), 1900b (Oreg.). 

Unpublished—R. T. Bingham 1964 (Idaho); T. C. Bray- 
shaw and A. E. Porsild 1964 (Canada); J. F. Franklin 1964 
(Oreg.); J. L. Jenkinson 1963 (Oreg.); F. D. Johnson 1964 
(Idaho); O. V. Matthews 1964 (Oreg.); R. Schmidt 1964 
(B.C.) ; C. A. Wellner 1964. 


Pinus lambertiana Dougl. MAP 7 
sugar pine 
Pinus lambertiana Dougl., Linn. Soc. London Trans. 
15: 500. 1827. 


Sugar pine ranges from western Oregon through the 
North Coast Range of California, the Sierra Nevada, and the 
mountains of southern California, with outliers in the Santa 
Lucia Mountains of west-central California and the Sierra 
San Pedro Martir of northern Baja California. 

Additional sources: 

Published—Baker 1956 (Oreg.); Goldman 1916 (Baja 
Calif.); Langille et al. 1903 (Oreg.); Leiberg 1900b (Oreg.); 
Ornduff and French 1958 (Oreg.). 

Unpublished—B. K. Ford 1963 (Calif.); J. F. Franklin 
1964 (Oreste) Re Griffimel9o7 (Callie) ss )reks: Jenkinson 1963 
(Calif. and Oreg.); O. V. Matthews 1964 (Oreg.) ; W. I. Stein 
1964 (Oreg.). 


Pinus flexilis James MAP 8 


limber pine 
Pinus flexilis James, Exped. Rocky Mts. 
RPA OO AUSAR: 

Limber pine ranges through the Rocky Mountains from 
southern Alberta and British Columbia to northern New 
Mexico, extending west through the mountain ranges of the 
Great Basin to southern California, western and northern 
Nevada, and eastern Oregon. It grows at high elevations 
through much of its range, but in Canada and the northern 
United States it is often confined to the prairie margins at 
the lower edge of the forest zone. It may be more widespread 
than the map shows on the poorly-known mountain ranges of 
the Great Basin. In the northern and western parts of its 
range it is often confused with Pinus albicaulis. It overlaps 
and to some extent intergrades with P. strobiformis at the 
southern edge of its range. Its southern limits are based on 
information provided by J. W. Andresen and R. J. Steinhoff 
from an unpublished investigation of variation in these two 
species (see below). 

Additional sources: 

Published—Ayres 1900a and 1900b (Mont.); Bacigalupi 
1933 (Calif.); Bailey and Bailey 1941; Douglass and Douglass 
1955 (Colo.) ; Goodding 1923 (Nebr.) ; Howell 1951 (Calif.); 
Leiberg 1904b (Mont.); Peck 1947 (Oreg.); Potter and Green 
1964 (N. Dak.). 

Theses—S. J. Preece, Jr. 1950. Floristic and ecological 
features of the Raft River Mountains of northwestern Utah. 
M.S. thesis, Univ. Utah, 103 pp., illus.; Raphael J. Steinhoff 
1964. Taxonomy, nomenclature, and variation within the 
Pinus flexilis complex. Ph.D. thesis, Michigan State Univ., 81 
pp. illus. 

Unpublished—]. W. Andresen 1963; W. H. Baker 1961 
(Idaho); R. T. Bingham 1963 (Mont.); T. C. Brayshaw and 
A. E. Porsild 1964 (Canada); W. C. Bullard 1963 (Calif) ; 
D. B. Coombs 1963 (Alta.); W. C. Cumming 1963 (Nev.); 
Margaret M. Douglass 1962 (Colo.); L. W. Hoskins 1963 
(Nev.); F. D. Johnson 1964 (Idaho); O. V. Matthews 1964 
(Idaho); M. A. McColm 1963 (Nev.); R. A. Read 1964 
(Nebr.). 


Pinus strobiformis Engelm. MAP 8 
southwestern white pine (see also Map 9) 


Pinus strobiformis Engelm. in Wisliz., Mem. Tour 
North. Mex. 102. 1848. 


Southwestern white pine extends from the southern part 
of Colorado through the mountains of Arizona, New Mexico, 
western Texas, and northern Mexico as far south as San Luis 
Potosi. Common synonyms for Pinus strobiformis include 
P. flexilis var. reflexa Engelm., P. reflexa (Engelm.) Engelm., 
and P. ayacahuite var. brachyptera Shaw. This species forms 
a link, both geographically and morphologically, between its 
neighbors P. flexilis and P. ayacahuite. The northern limits 
of P. strobiformis are based on information supplied by J. W. 
Andresen and R. J. Steinhoff (see P. flexilis) and Margaret M. 
Douglass (as P. flexilis var. macrocarpa Engelm.). Southwest- 
ern white pine is apparently separated from its southern rela- 
tive, P. ayacahuite, by a 100-mile gap that does not seem to 
correspond to any comparable discontinuity in suitable 
habitats. 

Additional sources: 

Published—Bailey 1913 (N. Mex.); Gentry 1942, 1946 
(Sin.); Johnston 1943 (Coah.); J. T. Marshall 1957; Plum- 
mer 1904 (Ariz.); Plummer and Gowsell 1904 (N. Mex.) ; 
Rixon 1905 (N. Mex.) ; G. C. Rzedowski 1960 (S.L.P.) ; Zobel 
andeGech: 1957(N-L.):. 

Thesis—James H. Maysilles 1956. Floral relationships 
of the pine forests of western Durango. Ph.D. thesis, Univ. 
Mich., 165 pp., illus. 

Unpublished—J. W. Andresen 1964; M. B. Applequist 
1964 (Ariz.); Margaret M. and J. R. Douglass 1962 (Colo.); 
E. Larsen 1962 (México); R. J. Steinhoff 1963 (Colo.); O. 
Zarzosa L. 1964 (Dgo.). 


Pinus ayacahuite Ehrenb. MAP 9 
Mexican white pine 


Pinus ayacahuite Ehrenb. in Schlecht., Linnaea 12: 492. 1838. 


Mexican white pine grows at high elevations from Jalisco 
and Hidalgo through central and southern Mexico to Guate- 
mala, El Salvador, and western Honduras. In _ northern 
Mexico it is replaced by the closely related Pinus strobiformis. 
The single collection from western Jalisco is, according to 
Martinez (1948, p. 126), morphologically intermediate be- 
tween these two white pines. 

Additional sources: 

Published—Allen 1955 (Honduras); Castanos 1962 
(Oax.); Hinds and Larsen 1961 (Jal.); Schwerdtfeger 1953 
(Guatemala); Standley and Steyermark 1958 (Guatemala); 
Wagner 1962 (Chis.). 

Unpublished—P. H. Allen 1962 (El Salvador); B. Hall- 
berg 1964 (Oax.). 


Pinus peuce Griseb. MAP 10 


Balkan pine 


Pinus peuce Griseb., Spicil. Fl. Rumel. Byth. 
rapt ae 

This species is confined to the high mountains of south- 
eastern Europe—southern Yugoslavia, western Bulgaria, north- 
ern Greece, and Albania. It is morphologically similar to the 
blue pine of the Himalayas, Pinus griffithii, from which it is 
separated by more than 3,000 miles. 
Sources: 

Published—Buffault 1931 (Greece); Heske 1959a (AI- 
bania) ; Markgraf 1932; Rikli 1943-1946. 

Unpublished—B. Pejoski (n.d.) to N. T. Mirov. 


Pinus armandii Franch. MAPS 11 AND 12 


Armand pine 
Pinus armandii Franch., Paris Mus. Hist. Nat. 
Nouv. Arch., Sér. 2, 7: 95-96, t. 12. 1885; “armandt.” 

This white pine of moderate and high elevations has a 
notably disjunct distribution. It ranges through western 
and southwestern China from Shansi and Kansu south to 
Yunnan and Kweichow, and west to northern Burma, €x- 
treme northeastern India, and southeastern Tibet. It is also 
present on the islands of Hainan and Taiwan and on two 
islands just south of Kyushu, Japan—Yaku Shima and Tanega 
Shima. In 1924 it was discovered in the Tsangpo Valley of 
southeastern Tibet by the plant explorer F. Kingdon Ward 
(Marquand 1929, Ward 1941b). Not until 1940 was it re- 
ported on the higher mountains of southern Hainan (Mer- 
rill and Chun 1940). 

We have tentatively included here a five-needled pine 
recently discovered in the mountains of western Anhwei 
Province, China. The description of this pine, Pinus an- 
hweiensis, has apparently not been published, but the locality 
where it was found has been reported by Wei (1963) . 

Another little-known white pine, found near Hué in 
central Vietnam, has been variously classified as Pinus arman- 
dit (Chevalier 1944), P. excelsa (P. griffithit) (Chevalier 
1919), and part of the P. fenzeliana complex (de Ferré 1960; 
see P. fenzeliana and Map 14). 

Sources: 

Published—Cheng 1932 (Kweichow), 1939 (Szechuan) ; 
C. K. Chow 1947 (Kansu); Diels 1900 (Szechuan); Farrer 
1926 (Kansu, Szechuan); Handel-Mazzetti 1927 and 1929-1936 
(Szechuan, Yunnan); Hers 1922 (Honan); Hsu 1950 (Yun- 
nan); Kabanov 1962 (Kansu, Shensi); Ku and Cheo 1941 
(Szechuan); Limpricht 1922 (Szechuan); Ludlow 1951 
(Tibet); Marquand 1929 (Tibet); Merril! 1941 (Burma); 
Merrill and Chun 1940 (Hainan); Orr 1933 (Yunnan); Reh- 
der 1923 (Shensi); Rock 1947 (Yunnan) ; Schweinfurth 1957; 
H. Smith 1925 (Shansi); Teng 1940 (Szechuan, Yunnan), 
1947 (Kansu); Walker 1941 (Kansu); Wang 1961; Ward 
1924 (Yunnan), 1941c (Burma), 1949 (Burma), 1952b (As- 
sam); Wilson 1907 (Hupeh), 1913 (Hupeh, Szechuan), 1916 
(Japan), 1926 (Yunnan); Wu 1956. 

Thesis—Chung-Iwen Wu 1947. The phytogeographical 
distribution of pine in China. M.F. thesis, Yale Univ., 74 
pp., illus. 

Unpublished—Forest Research Institute, Taiwan, to N. 
T. Mirov 1961 (Taiwan) ; Shiu-ying Hu 1963; Wang Chi-Wu 
1964. 


Pinus griffithii McClelland MAP 12 
blue pine (see also Map 14) 
Pinus griffithii McClelland in Griffith, Notul. 

Pl. Asiat. 4: 17. 1854; Icon. Pl. As. 4. t. 365, excl. fig. 1-3. 1854. 

Blue pine has also been known as Pinus excelsa Wall., 
P. wallichiana A. B. Jacks., and P. chylla Lodd. It ranges 
throughout the Himalaya Mountains, extending beyond them 
to eastern Afghanistan, northeastern Baluchistan (West Paki- 
stan) , northern Burma, and Yunnan Province, China. It is an 
important component of the middle- and high-elevation Hima- 
layan forests, especially in the drier inner valleys. Its eastern 
outliers in southeastern Tibet and northern Burma have been 
found by the plant explorer F. Kingdon Ward within the past 
few decades. Only recently it has been observed in the cen- 
tral part of India’s Northeast Frontier Agency approximately 
28° N. lat., 90° E. long.) , where it may have been introducet 
(Fiirer-Haimendorf 1955, pp. 18, 62). 

Sources: 

Published—Aitchison 1879, 1881-82 (Afghanistan); Ban- 
erji 1958 (Nepal) ; Bor 1988 (Assam) ; Burkill 1909 (Baluchi- 
stan); Cooper 1983 (Bhutan); Duthie 1893 (Kashmir), 1898 
(Pakistan); Fiirer-Haimendorf 1955 (Assam); Gammie 1898 


933 (India) ; Kawakita 1956 (Nepal); Lin- 

zorovsky 1949 (Afghanistan); Malik 1962 

aaston 1922 (India); Pirson 1962 (India); 

Nepal); Schweinfurth 1957; Shebbeare 1934 

zh 1929 (Kashmir); Stebbing 1906 (Baluchi- 

[955-1956 (Nepal); Troup 1921 (Pakistan); Ward 

bet), 1936b (Tibet), 194la, 1944-45 (Burma), 1949 

jurma) ; L. H. J. Williams 1953 (Nepal). 

Unpublished—Wang Chi-Wu 1964 (Burma, China). 


; dalatensis de Ferré MAP 14 


Pinus dalatensis de Ferré, Toulouse Soc. d’Hist. 
Nat Bul) 952) 178) fig: 271351960: 


This recently described pine is confined to the mountains 
near the town of Dalat in southern Vietnam. De Ferré (1960) 
considers it most closely related to Pinus griffithit, from which 
it is separated by more than 1,000 miles, and to P. peuce of 
southeastern Europe. 

Source: 
Published—de Ferré 1960. 


Pinus parviflora Sieb. & Zuce. MAP 13 


Japanese white pine 
Pinus parviflora Sieb. & Zucc., Fl. Jap. 
221, tet lo. L844. 

Japanese white pine grows throughout Japan as far 
north as southern Hokkaido, and on the Korean island of 
Utsuryo (Ullung). It is the northernmost of a chain of 
related white pines which replace each other along the east- 
ern edge of Asia from Vietnam to Hokkaido, Japan (Pinus 
parviflora, P. morrisonicola, P. fenzeliana, and possibly P. 
wangi). P. parviflora consists of two geographical varieties 
which intergrade in central Honshu, The northern variety is 
sometimes called P. pentaphylla Mayr; the southern, P. hime- 
komatsu Miyabe & Kudo or a variety of the former. 

Sources: 
Published—Hayashi 1954 (Japan); Uyeki 1926 (Korea). 


Pinus morrisonicola Hayata MAP 14 
Taiwan white pine 
Pinus morrisonicola Hayata, Gard. Chron., Ser. 


Oe Oa O08! 


This pine is confined to the mountains of Taiwan. It 
is closely allied to both Pinus parviflora of Japan and P. 
fenzeliana of southern China. 
Source: 

Unpublished—Forest Research Institute, Taiwan, to N. 
T. Mirov 1961. 


Pinus fenzeliana Hand.-Mazz. MAP 14 
Pinus fenzeliana Hand.-Mazz., Oesterr. Bot. Ztschr. 
80: 337. 1931. 

This pine grows in southern mainland China (Kwang- 
tung, Kwangsi, and southern Hunan), on the island of 
Hainan, and possibly in central Vietnam. Neither southern 
China nor Vietnam has been thoroughly explored botanically, 
and the range of the species is still poorly known. 

We are including here Pinus kwangtungensis Chun ex 
Tsiang (Tsiang 1948), which appears to be synonymous 
with P. fenzeliana. Also included is a white pine of central 
Vietnam which de Ferré (1960) has tentatively classified as 
P. fenzeliana on the basis of a vegetative specimen. This 
Vietnam pine had earlier been variously referred to P. excelsa 
(P. griffithiz) (Chevalier 1919) and P. armandii (Chevalier 
1944). 


8 


Pinus fenzeliana is part of the P. parviflora group, and 
was first classified as P. morrisonicola after its discovery in 
the 1920’s (Merrill 1927). Recently Wu (1956) has re- 
named both P. fenzeliana and P. morrisonicola as varieties of 
P. parviflora. 

Sources: 

Published—de Ferré 1960 (Vietnam); Handel-Mazzetti 
1931 (Hainan); Merrill 1927 (Hainan); Tsiang 1948; Wang 
1961, p. 144 (Kwangsi) . 


Unpublished—Arnold Arboretum Herbarium: W. T. 
Tsang 28496 (N. Kwangsi) . 
Pinus wangii Hu & Cheng MAP 14 


Pinus wangit Hu & Cheng, Fan Mem. Inst. Biol. 
Bul., n.s., 1: 191. 1948. 


This pine is known only from the limestone hills of south- 
eastern Yunnan where it was discovered. In many features 
it resembles the Pinus parviflora group, but its leaf structure 
is notably different (de Ferré 1960, Kwei and Lee 1963) . 
Source: 

Published—Hu and Cheng 1948. 


SECT. PARRYA Mayr 


SUBSECT. CEMBROIDES Engelm. 


Pinus cembroides Zuce. MAP 15 
Mexican pinyon (see also Map 17) 
Pinus cembroides Zucc., K. Bayer. Akad. Wiss. 
Miinchen, Abhand]. Math.-Phys. 1: 392. 1832; 

Flora [Jena] 15 (2), Beibl. 93. 1832. 


Mexican pinyon is widespread at low elevations in the 
mountains bordering the arid plateau of northern Mexico. 
It ranges from southeastern Arizona, southwestern New Mex- 
ico, and western Texas through northern Mexico and south 
to Puebla and México. It overlaps the closely related Pinus 
edulis in southern Arizona and New Mexico. The reported 
occurrence of P. cembroides in northern Baja California (Shaw 
1909, p. 6; Martinez 1948, p. 84) is not shown on the map. 
Additional sources: 

Published—Goldman 1951; Johnston 1943 (Coah.) ; G. C. 
Rzedowski 1960 (S.L.P.) ; Zobel and Cech 1957 (N.L.). 

Unpublished—C. E. Blanco 1964 (Dgo., Coah.); F. Medel- 
lin-Leal 1964 (S.L.P.); O. Zarzosa L. 1964 (Dgo.). 


Pinus edulis Engelm. MAP 17 


pinyon 
Pinus edulis Engelm. in Wisliz., Mem, Tour 
North. Mex. 88. 1848. 


Pinyon is widespread in the southwestern United States, 
just entering Wyoming and extending south through Utah, 
Colorado, and New Mexico to the western end of Oklahoma, 
west Texas, and the northern border of Chihuahua; extend- 
ing west throughout much of Arizona to an outlier in eastern 
California. Its range overlaps that of the Mexican pinyon, 
Pinus cembroides, in southern Arizona and New Mexico. P. 
edulis also overlaps, and may occasionally intergrade with, P. 
monophylla at the western edge of its range. The relative 
distribution of these two species has been confused by the 
prevalence of a single-leaved form of P. edulis in central 
Arizona south of the Mogollon Rim, but it has been estab- 
lished that this form is a variation of P. edulis (Little 1950, 


[Ds 74) ie 


Additional sources: 

Published—Deaver and Haskell 1955 (Ariz.); Jack 1900 
(Colo.) ; Leiberg et al. 1904 (Ariz.); Merriam 1890 (Ariz.); 
Peterson 1962 (Wyo.); Plummer and Gowsell 1904 (N. Mex.); 
Sudworth 1900a (Colo.); Weber 1961 (Colo.); Wolf 1938 
(Calif.). 

Thesis—W. D. Stanton 1931. 
flora of the Henry Mountains of Utah. 
Young Univ., 63 pp., illus. 

Unpublished—W. P. Cottam 1964. 


A preliminary study of the 
M.A. thesis, Brigham 


Pinus quadrifolia Parl. MAP 16 
Parry pinyon 
Pinus quadrifolia Parl. ex Sudw., U. S. Dept. Agr. 
Div. Forestry Bul. 14: 17. 1897. 


Parry pinyon is confined to low elevations in the dry 
mountains of southern California and northern Baja Cali- 
fornia from the San Jacinto Mountains south to the Sierra 
San Pedro Martir. Its range overlaps that of Pinus mono- 
phylla, but no intermediates have been reported. All of the 
sources listed below pertain to Baja California. 

Additional sources: 
Published—Goldman 1916; Wiggins 1944. 
Unpublished—Reid Moran 1963, 1964; J. Olmsted 1962. 


Pinus monophylla Torr. & Frém. MAPS 16 AND 17 
singleleaf pinyon 
Pinus monophylla Torr, & Frém. in Frém., Rpt. 
Explor. Exped. Rocky Mts. 319, t. 4. 1845: 


: ‘monophyllus. 2 


Singleleaf pinyon is widespread and common at low ele- 
vations in the isolated mountain ranges of the Great Basin, 
ranging from southern Idaho, western Utah, and northwest- 
ern Arizona through most of Nevada and eastern and central 
California to northern Baja California. It overlaps Pinus 
edulis in southwestern Utah, northwestern Arizona, and the 
New York Mountains of southern California, and intermedi- 
ate forms are known. This species was once thought to 
extend into central Arizona (Munns 1938), but the single- 
leaved pinyon of that region is now considered a variation of 
P. edulis (Little 1950, p. 12). The range of P. monophylla 
has recently been extended more than 100 miles south, in Baja 
California (Anonymous 1963; Reid Moran 10319) . 
Additional sources: 

Published—Anonymous 1963; Bailey and Bailey 1941; 
Goldman 1916 (Baja Calif.) ; Merriam 1893; Wiggins 1940 
(Baja Calif.). 

Thesis—C. McMillan 1948. A taxonomic and ecological 
study of the flora of the Deep Creek Mountains of central 
western Utah. M.S. thesis, Univ. Utah, 99 pp., illus. 

Unpublished—W. C. Bullard 1963 (Calif.); W. P. Cot- 
tam 1964 (Utah); G. K. Griffith 1963 iNew): L. W. Hoskins 
1963 (Nev.); M. Humphreys 1963 (Nev.); F. D. Johnson 1964 
(Idaho); M. A. McColm 1963 (Nev.); if H. Thomas 1964 
(Baja Calif.); W. W. Wagener 1963 (Calif.) . 


Pinus culminicola Andresen & Beaman MAP 18 


Potosi pinyon 
Pinus culminicola Andresen & Beaman, Arnold 
Arboretum Jour. 42: 438, fig. 2-4, 1961. 

This recently named species is the only high-elevation 
pinyon. It has been found only on Cerro Potosi, possibly 
the highest mountain in Nuevo Leén, where it grows as a 
timberline shrub (Andresen and Beaman 1961) and in the 


forest understory. It had previously been identified as Pinus 
flexilis (Muller 1939). 


Pinus maximartinezii Rzedowski MAP 15 
Martinez pinyon 
Pinus maximartinezii Rzedowski, Ciencia 23: 17, 
fig. 1-3, t. 2. 1964 

This remarkable new pinyon is known only from its 
type locality, the Cerro de Pifiones in the southern part of 
Zacatecas, Mexico, near the town of Juchipila. Discovered 
only in 1963, it is notable among the pinyons for its very 
large cones (Rzedowski 1964) . 


Pinus pinceana Gord. MAP 18 
Pince pinyon 
Pinus pinceana Gord., Pinet. 204. 1858. 

This pinyon is restricted to a few areas in northeastern 
and eastern Mexico from central Coahuila to central Hidalgo, 
and is nowhere common. It is often associated with Pinus 
cembroides, the common pinyon of Mexico. 
Additional sources: 

Published—Johnston 1943 (Coah., Zac. 
(S.L.P.); McVaugh 1952 (Qro.). 


); Diaz Luna 1962 


Pinus nelsonii Shaw MAP 18 
Nelson pinyon 
Pinus nelsonii Shaw, Gard. Chron. Ser. 3, 
36: 122, fig. 49. 1904; “nelsoni” 

This pinyon is confined to a few scattered localities in 
northwestern Mexico, from southern Coahuila, southern 
Nuevo Leon, and western Tamaulipas to San Luis Potosi. 
One of the rarest of the Mexican pines, it is associated with 
the common Mexican pinyon, Pinus cembroides. 

Additional sources: 


Published—Crespo 1963 (Tamps.) ; Miroy 1961 (Tamps.). 


SUBSECT. GERARDIANAE Loud. 


Pinus gerardiana Wall. MAP 19a 
chilgoza pine 
Pinus gerardiana Wall. ex D. Don in Lamb., Desc. 


Genus Pinus. Ed. 3 (8°), v. 2, unnumbered page 
between p. 144 and p. 145, t. 79. 1832. 

This species, noted chiefly for its large edible seeds, is 
confined to the mountains of eastern Afghanistan, parts of 
Pakistan, and scattered localities in the dry inner valleys ol 
the northern Himalayas. It may be more widely distributed 
than the map shows in Wazciristan, along the Afghanistan- 
Pakistan border. It is separated from Pinus bungeana, ms 
only close relative, by the 1,300-mile-long Tibetan highland 
Sources: 


Published—Aitchison 1879, 1881-1882 \fehanistan 
Burkill 1909 (Baluchistan); Clarke 1957 \fghanistan); 
Duthie 1898 (Pakistan); Linchevsky and Prozorovsky 1949 
(Afghanistan); Schweinfurth 1957; Stebbing 1906 (Baluchis 
tan). 

Unpublished—E. Nasir 1963 (Pakistan R. J. Rodin 


1963 (Pakistan); G. L. Webster 1963 (Kashmn 


9 


ana Zuece. MAP 19b 
ingeana Zucc. in Endl., Synops. Conif. 
166. 1847. 
rk pine grows in northern China from northern 
nd Hupeh north to Shansi and Hopeh. Notable 
its peculiar Platanus-like bark, this pine is com- 
rare throughout its range. 


ublished—Cheng 1939 (Szechuan); H. F. Chow 1934 

peh) ; Hers 1922 (Honan); Hu 1935 (Honan); H. Smith 

5 (Shansi); Sowerby 1937; ‘Tang 1931 (Shansi) ; Teng 1947 
su); Wilson 1907 (Hupeh). 

Unpublished—Wang Chi-Wu 1964 (Shensi). 


SUBSECT. BALFOURIANAE Shaw 


Pinus balfouriana Grev. & Balf. MAP 20 
foxtail pine (see also Map 21) 
Pinus balfouriana Grev. & Balf. in A. Murr., Bot. 
Exped. Oreg. | Rpt. No. 8] No. 618, t. 1853. 


Foxtail pine is confined to two high-elevation areas about 
300 miles apart: the southern Sierra Nevada, Calif., and the 
higher mountains of northwestern California from the South 
Yolla Bolly Mountains north to the Marble Mountains. The 
stands on the east slope of the Sierra Nevada are about 20 
miles from stands of Pinus aristata in the Inyo Mountains, 
only the width of the arid Inyo Valley separating the two 
species at this point. A northern outlier of the Sierra Nevada 
distribution reported by Sudworth (1908, p. 41) on the 
divide between the South Fork of the San Joaquin River and 
the North Fork of the Kings River has never been verified, 
and is not included in the distribution shown on the map. 
Additional sources: 

Published—Howell 1951. 

File reports—“Survey of foxtail pine in northern Cali- 
fornia” by T. H. Harris and Benton Howard, August 28, 1956. 
Forest Service, Region 5, San Francisco, Calif.; “Blister rust 
occurrence survey in the Marble Mountains, Klamath Na- 
tional Forest’”’ by Benton Howard, October 4, 1956. Forest 
Service, Region 5, San Francisco, Calif. 

Unpublished—D. V. Hemphill 1964 (northern Calif.) . 


MAP 21 
(see also Map 20) 
Pinus aristata Engelm. in Parry & Engelm., Amer. 
Jour. Sci. and Arts, Ser. 2, 34: 331. 1862. 


Bristlecone pine ranges from eastern California through 
Nevada, Utah, and northern Arizona to Colorado and north- 
ern New Mexico. It grows at high elevations and frequently 
extends to timberline. In many places it is very localized, 
but in others, such as the White Mountains of eastern Cali- 
fornia, it forms extensive stands. It is probably more widely 
distributed in the poorly known mountain ranges of Nevada 
than the map indicates. 

Additional sources: 

Unpublished—W. P. Cottam 1964 (Utah); Margaret M. 
Douglass 1962 (Colo.); C. W. Ferguson 1962 (Calif.); R. D. 
Wright 1962 (Calif.). 


Pinus aristata Engelm. 
bristlecone pine 


10 


SUBGEN. PINUS 
SECT. TERNATAE Loud. 


SUBSECT. LEIOPHYLLAE Loud. 


Pinus leiophylla Schiede & Deppe MAP 22 


Chihuahua pine 
Pinus letophylla Schiede & Deppe in Schlecht. & 
Cham., Linnaea 6: 354. 1831. 

Chihuahua pine ranges from southern and central Mexico 
through western Mexico to southwestern New Mexico and 
southern and central Arizona. Included here is Pinus leio- 
phylla var. chihuahuana (Engelm.) Shaw, sometimes consid- 
ered a separate species, P. chihuahuana Engelm. P. leiophylla 
is unique among American pines in having cones that require 
three seasons to mature. Its northernmost known occurrences 
are just south of the Mogollon Rim in central Arizona (7 
miles west of Pinedale—Sudworth 1927; about 25 miles east 
of Payson—M. B. Applequist, personal communication, 1964). 
Additional sources: 

Published—Goldman 1951; Guzman and Vela 
(Zac.) ; J. IT. Marshall 1957 (Son., Chih.) ; Shaw 1909. 

Thesis—James H. Maysilles 1959. Floral relationships 
of the pine forest of western Durango, Mexico. Ph.D. thesis, 
Univ. Michigan, 165 pp., illus. 

Unpublished—M. B. Applequist 1964 (Ariz.); B. Hall- 
berg 1964 (Oax.); E. Larsen 1962; N. Sanchez Mejorada to 
L. W. Bryan 1963 (Mich.); O. Zarzosa L. 1964 (Dgo.). 


1960 


Pinus lumholtzii Robins. & Fern. MAP 23 
Lumholtz pine 
Pinus lumholtzii Robins. & Fern., Amer. Acad. 
Proc. 30: 122. 1895. 

This species grows at relatively low elevations in north- 
western Mexico, from Sonora and Chihuahua south to Jalisco. 
It is notable for its pendent but stiff needles. 

Additional sources: 

Published—Gentry 1942, 1946 (Sin.); Guzman and Vela 
1960 (Zac.); Shaw 1909 (Nay.). 

Thesis—James H. Maysilles 1959. Floral relationships of 
the pine forests of western Durango, Mexico. Ph.D. thesis, 
Univ. Michigan, 165 pp., illus. 

Unpublished—C. E. Blanco 1964 (Dgo.) ; E. Larsen 1962; 
O. Zarzosa L. 1964 (Dgo.). 


SUBSECT. CANARIENSES Loud. 


Pinus canariensis C. Smith MAP 24b 
Canary Island pine 
Pinus canariensis C. Smith in Buch, Phys. Beschr. 
Canar. Ins. 159. 1825. 

Canary Island pine is confined to the central and western 
Canary Islands, off the coast of northwestern Africa. Its only 
close relative, Pinus roxburghii, grows in the Himalaya 
Mountains more than 5,000 miles away. 

Sources: 

Published—Ceballos and Ortuno 1951. 

Unpublished—L. Ceballos to N. T. Mirov (n.d.) (Gran 
Canaria). 


Pinus roxburghii Sarg. MAP 24a 
chiy pine 
Pinus roxburghti Sarg., Silva No. Amer. 11: 9. 1897. 

This species, long known as P. longifolia Roxb., is con- 
fined to the monsoon belt of the outer Himalayas from Bhutan 
to the northeastern part of West Pakistan. It grows at low 
elevations (generally below 7,000 feet). Early reports to the 
contrary, it apparently has not been found in Afghanistan 
except in cultivation (Aitchison 1891). Its distribution in 
Nepal and Bhutan is only sketchily known. It has not been 
reported east of Bhutan, but this part of India is almost un- 
explored botanically (Schweinfurth 1957) . 

It is separated by more than 5,000 miles from its closest 
relative, Pinus canariensis. 

Sources: 

Published—Banerji 1958 (Nepal); Biswas 1933; Burkill 
1910 (Nepal); R. E. Cooper 1933 (Bhutan); Gammie 1898 
(India); Gorrie 1933 (India); Kawakita 1956 (Nepal); Malik 
1962 (Pakistan); Osmaston 1922 (India); Pirson 1962 
(India) ; Polunin 1950 (Nepal); Schweinfurth 1957; S. Singh 
1929 (Kashmir); Smith and Cave 1911 (Sikkim); Tilman 
1952- (Nepal); Troup 1921 (India); L. H. J. Williams 1953 
(Nepal). 

Unpublished—R. J. Rodin 1963 (Pakistan) . 


SUBSECT. PINEAE Shaw 


Pinus pinea L. MAP 25 


Italian stone pine 
Pinus pinca L., Sp. Pl. 1000. 1753. 


Italian stone pine is widespread on the Iberian peninsula 
(Spain and Portugal), ranges along the northern and eastern 
shores of the Mediterranean Sea, and occurs at a few places 
near the north coast of Turkey. It is the only Eastern Hemi- 
sphere species with cones that require three seasons to develop 
to maturity. It is also notable for its edible seeds, and has 
been widely planted for many centuries. Its natural range 1s 
highly uncertain (Rikli 1926) , but we have omitted those 
stands known to be artificial in origin, such as the well-known 
forest near Ravenna, Italy. 

Sources: 

Published—Briquet 1910 (Corsica); Buffault 1931 
(Greece) ; Chodat 1913 (Portugal); Feinbrun 1959 (Lebanon); 
Gaussen 1953-1954 (France); Heske 1959a (Albania); Holm- 
boe 1914 (Cyprus); Kayacik 1957 (Turkey); Knoche 1921 
(Balearic Islands); Philippson I8957(Greece)s 1919) (Turkey) 
Rechinger 1943 (Greece): Rikli 1926; Touring Club Italiano 
1958 (Italy); Turrill 1929. 

Unpublished—Director-General, Forest Services of Portu- 
gal, 1964; H. Gaussen 1964 (France); General-Director of 
Forestry, Turkey, 1964; Instituto Forestal de Investigaciones 
Vy Experiencias, pa 1962; B. Kasapligil to N. T. Mirov 
1961 (T urkey) ; Kayacik 1964 (Turkey); E. Magini to 
N. T. Mirov hie (Italy); C. Moulopoulos 1964 (Greece); A. 
de Philippis 1964 (Italy). 


SECT. PINUS 


SUBSECT. SYLVESTRES Loud. 


Pinus resinosa Ait. MAP 26 
red pine 
Hort. Kew. 3: 367. 1789 


from New- 


Pinus resinosa Ait., 


Red pine ranges through southern Canada 
foundland to southeastern Manitoba, and south through the 
Lake States and the northeastern United States, with outliers 
in Illinois and West Virginia. This species is the only rep- 
resentative of this typically Eurasian group of pines on the 
North American mainland. Its closest relative is probably 
Pinus nigra of southern Europe. 

Additional sources: 

Published—Brenneman 1956 (Ill.); Cook, Smith, and 
Stone 1952 (N.Y.) ; Haddow 1948a and 1948b (Ont.). 

Unpublished—T. C. Brayshaw and A. E. Porsild 1964 
(Canada) ; Gary Saunders (N.F.). 


Pinus tropicalis Morelet MAP 26 
tropical pine 
Pinus tropicalis Morelet, Rev. Hort. Cote d’Or 
Le? 106: W851 


This species is confined to the Isle of Pines and the low 
mountains of Pinar del Rio Province, western Cuba. It is 
separated from Pinus resinosa, the only other Western Hemi- 
sphere representative of this group, by more than a thousand 
miles. 


Sources: 

Published—E. E. Smith 1954; Marie-Victorin and Léon 
1942. 
Pinus nigra Arnold MAP 27 


Austrian pine 
Pinus nigra Arnold, Reise Mariazell 8, t. 1785. 

Austrian pine ranges widely through southern Europe 
from Spain to Austria and throughout much of the Balkan 
Peninsula, east to southern Russia and south to Turkey: on 
the islands of Cyprus, Sicily, and Corsica, with outliers in Al- 
geria and Morocco. Its variability in nature, recently de- 
scribed by Rohrig (1957), is reflected in the great taxonomic 
complexity of this species (Schwarz 1936). 

Austrian pine is Bee ociatha with several related species in 
various parts of its range, including the closely similar Pinus 
heldreichii and the widespread P. sylvestris. Within the 
limited range of P. heldreichii, that species and P. nigra are 
sometimes confused. 

Reports of Pinus nigra on the Mediterranean islands of 
Crete and Rhodes are apparently in error (Rechinger 1951). 
Sources: 

Published—Akademia Nauk SSSR 1954 (U.S.S.R.); Anié 
1957 (Yugoslavia): Bernhard 1931 (Turkey): Buftault 1931 
(Greece) ; Comité de Geographie du Maroc 1957 (Morocco) ; 
Debazac 1963 (France), 1964 (Corsica): Dudié 1953 (Yugo- 
slavia); Fekete 1959 (Rumania); Fekete and Blattny 1913-1914 
(Yugoslavia); Fukarek 1958; Gaussen (n.d.) and 1953-1954 


(France, Spain); Giacobbe 1937 (Italy); Heske 1954 (Tur- 
key), 1959a (Greece) ; Jovantevié 1961 (Yugoslavia) ; Kayacik 
1954 (Turkey): Kosanin 1929 (Yugoslavia) ; yore and 
Peverimhoff 1927 (Algeria); Markgraf 1932 Albania 
Miletié 1959 (Yugoslavia) ; Pejoski 1956 (Yugoslavia Philip. 
pson 1895 (Greece); Rechinger 1951 (Greece); Rubnet and 
Reinhold 1953, fig. 74 (Crimea): Salvador 1927 (Spain) 


Scharfetter 1988 (Austria); Vidakovié 1957 (Yugoslavia) 


{—Forestry Research Institute, Rumania, 
irector of Forestry, Turkey, 1964; Instituto 
vestigaciones y Experiencias, Spain, 1962; B. 

. IT. Mirov 1961 (Turkey); H. Kayacik 1964 
Magini to N. T. Mirov 1960 (Italy); E. D. 
(961 (Cyprus); R. Morandini 1964 (Italy); A. de 
61 and 1964 (Italy); E. Rohrig 1964 (Austria) . 


sinus heldreichii Christ MAP 28 
leldreich pine 
Pinus heldreichii Christ, Naturf. Gesell. 
Basel Verhandl. 3: 549. 1863. 

This pine is confined to high elevations on the Balkan 
peninsula (central Yugoslavia to northern Greece and south- 
western Bulgaria) and in southern Italy. Since it is sometimes 
confused with its much more widespread associate, Pinus 
nigra, its range is not yet fully established. Within the past 
few years its known distribution in Montenegro (Laku&i¢ 
1961) and Serbia (ToSi¢ 1959) has been considerably en- 
larged. 

Sources: 

Published—Delevoy 1948; Fukarek 1941; Gourlay 1938 
(Greece) ; Heske 1959a (Albania); Jedlowski 1959 (Italy); 
Lakusié 1961 (Yugoslavia); Longhi 1956 (Italy); Markgraf 
1931; Tosié 1959 (Yugoslavia) . 


Unpublished—E. Magini to N. T. Mirov 1960 (Italy: 
Campania); C. Moulopoulos 1964 (Greece). 
Pinus mugo Turra MAP 29 


Swiss mountain pine 

Pinus mugo ‘Vurra, Gior. Ital. (Grisilini) 1: 152. 1764. 

Pinus mugo (P. montana Mill.) grows at high elevations 
in the mountains of central and southern Europe (Alps, 
Pyrenees, Carpathians, Balkans), ranging east and south to 
the U.S.S.R., Rumania, Bulgaria, and central Italy, north to 
Germany and Poland, and west to eastern Spain, where the 
species reaches its southern limit. It ranges in form from a 
dwarfed bush to a small tree. The tree form, which is preva- 
lent in the Pyrenees and the western Alps, is sometimes 
segregated as Pinus uncinata Ramond. The unusually com- 
plex nomenclature of this pine is based primarily on_ its 
variable growth form and cones (Willkomm 1861, Kirchner 
et al. 1908). 

Pinus mugo is associated with the closely related P. 
sylvestris in many parts of its range, and the two are believed 
to hybridize in nature (Fitschen 1930, p. 430). 

Sources: 

Published—Adamovié 1909; Akademia Nauk SSSR 1954 
(U.S.S.R.) ; Beck 1901 (Yugoslavia) ; Ceballos 1941 (Spain) ; 
- Cermak et al. 1955 (Czechoslovakia); Fekete and Blattny 
1913-1914; Fukarek 1959 (Yugoslavia); Gaussen 1923 
(France), 1956 (France, Spain); Giordano 1962b (Bulgaria) ; 
Kirchner et al. 1908; KoSanin 1929 (Yugoslavia) ; Markgrat 
1932 (Albania); Merxmiiller 1954; Scharfetter 1938 (Austria) ; 
Schmid 1949 (Switzerland); Touring Club Italiano 1958 
(Italy); Willkomm 1861. 

Unpublished—Forestry Research Institute, Rumania, 
1964; JF. Lacaze 1963 (France); R. Morandini 1964 (Italy) ; 
S. Bialobok 1964 (Poland). 


Pinus pinaster Ait. MAP 30 


maritime pine 
Pinus pinaster Ait., Hort. Kew. 3: 367. 1789. 
Maritime pine ranges from the Iberian peninsula (Spain, 
Portugal) through southern France to the west coast of Italy, 


12 


in northern Africa from Morocco to Tunisia, and to the is- 
lands of Corsica and Sardinia. Its natural distribution has 
been somewhat obscured by the widespread planting of this 
species. We have excluded the extensive planted stands in 
southwestern France and northwestern Spain, but a portion 
of the distribution of this species in Portugal is certainly of 
artificial origin. 

Sources: 

Published—Comité de Geographie du Maroc 1957 (Mo- 
rocco); Desole 1960 (Sardinia); Gaussen (n.d.) (France, 
Spain) , 1953-1954 (France); Gaussen and Vernet (n.d.) (Al- 
geria, Tunisia); Guinier 1952; Rikli 1943-1946; Servicgo de 
Reconhecimento e de Ordenamento Agrario 1962 (Portugal) ; 
Touring Club Italiano 1931 (Italy). 

Unpublished—H. Gaussen 1964 (France); Instituto For- 
estal de Investigaciones y Experiencias 1962 (Spain); A. de 
Philippis 1961 and 1964 (Italy). 


Pinus halepensis Mill. MAP 31 
Aleppo pine 
Pinus halepensis Mill., Gard. Dict. Ed. 8, 
Pinus No. 8. 1768. 


Aleppo pine is widely distributed in the Mediterranean 
region, ranging from Morocco to Tunisia and Libya (Cyre- 
naica) in North Africa, with outliers extending to the northern 
edge of the Sahara Desert; from eastern Spain through south- 
ern France and Italy to the Adriatic coast of Yugoslavia and 
to Greece; at the eastern end of the Mediterranean Sea from 
Israel and Jordan north to Turkey. Pinus brutia replaces 
this species in the northeastern part of the Mediterranean 
region, and the two are known to hybridize in the region of 
overlap (see P. brutia). The natural occurrence of P. 
halepensis on the island of Corsica is uncertain; Nahal 
(1962b) and H. Gaussen (personal communication, 1964) 
indicate that it occurs in one place, but Briquet (1910) states 
that it is not native to the island. 

Sources: 

Published—Beck 1901 (Yugoslavia); Comité de Geo- 
graphie du Maroc 1957 (Morocco); Feinbrun 1959 (Leb- 
anon); Francini 1953 (Italy); Gaussen 1953-1954 (France) ; 
Giordano 1962a (Lebanon); Heske 1959a (Greece) ; Kasapli- 
gil 1956 (Jordan); Kayacik 1954 (Turkey): Maire 1926 (AI- 
geria, Tunisia); Markgraf 1932 (Albania); Mikesell 1961 
(Morocco) ; Nahal 1962a (Syria), 1962b (France, Corsica); 
Touring Club Italiano 1931 (Italy); Turrill 1929 (Greece); 
Zohary 1947 (Israel, Jordan). 

Unpublished—Instituto Forestal de Investigaciones y 
Experiencias, Spain, 1962; H. Kayacik 1962, 1964 (Turkey) ; 
H. G. Keith 1961 (Libya); E. Magini to N. T. Mirov 1960 
(Italy); R. Morandini 1964 (Italy); J. Papaioannou 1964 
(Greece); A. de Philippis 1961 and 1964 (Italy); M. Vidako- 
vic 1964 (Yugoslavia). 


Pinus brutia Ten. MAP 31 


Pinus brutia Ten., Prodr. Fl. Nap. Ixxii. 1811. 


This close relative of Pinus halepensis largely replaces 
that species in the eastern Aegean region, Turkey, the Levant, 
on the Mediterranean islands of Crete and Cyprus, and in 
the Black Sea region. The ranges of the two overlap in 
northeastern Greece, where they hybridize (Papajoannou 
1936), in the Levant (Feinbrun 1959, Nahal 1962a), and in 
southeastern Turkey (Kayacik 1954). 

Variants of Pinus brutia in the Black Sea region are 
sometimes called P. pithyusa Stev. and P. eldarica Medw._ P. 
pithyusa grows along the northern and northeastern shores 
of the Black Sea, P. eldavica on a single low mountain (Eilar- 
Ugi) in the central Transcaucasus. 


Pinus brutia or one of its variants has long been thought 
to occur naturally in Iran and Afghanistan, and Papaioan- 
nou’s map of the distribution of this species (in Nahal 1962b) 
shows it in these countries. Planted in both countries, it is 
especially common in Afghanistan (Aitchison 1891, Clarke 
1957). But comparatively intensive recent botanical explora- 
tions of these two countries have failed to find any natural 
stands of this pine (Bobek 1951; Kitamura 1960; Linchevsky 
and Prozorovsky 1949; Zohary 1963; J. Papajoannou, personal 
communication, 1964). Aitchison (1891) attempted to track 
down rumors of natural forests of this pine, but found only 
planted trees. The source of seed of this common Afghanistan 
pine remains an intriguing question, since the nearest natural 
stands of the species are almost 1,000 miles from Afghanistan. 
Sources: 

Published—Akademia Nauk, SSSR, 1954 (U.S.S.R.); 
Alemdag 1962 (Turkey); Bernhard 1931 (Turkey) ; Chatur- 
vedi 1961 (Syria); Czeczott 1954; Feinbrun 1959 (Lebanon) ; 
Giordano 1962a (Lebanon); Heske 1954 (Turkey), 1959a 
(Greece) , 1959b (Iraq) ; Kayacik 1954 (Turkey) ; Ketskhoveli 
1960) (U:S:S:R.); Maleéjeff 1929 (U.S.S.R.); Nahal 1962a 
(Syria); Radde 1899 (U.S.S.R.); Rechinger 1951 (Greece) ; 
Rubner and Reinhold 1953 (U.S.S.R.); Selik 1959 (Turkey). 

Unpublished—General Director of Forestry, Turkey, 
1964; H. Kayacik 1964 (Turkey); E. D. Michaelides 1961 
(Cyprus) ; J. Papajoannou 1964. 


Pinus sylvestris L. 
Scotch pine 
Pinus sylvestris L., Sp. Pl. 1000. 1753. 

The most widely clistributed of the pines, Scotch pine 
grows throughout northern Eurasia from Scotland, southern 
Spain, northern Greece, and northern Turkey to northern 
Manchuria and the Sea of Okhotsk. In Europe it is associated 
with several related species, including Pinus mugo and P. 
nigra. A principal source of information for our map is the 
excellent distribution map by Steven and Carlisle (1959) . 
Sources: 


MAP 32 


Published—Akademia Nauk, SSSR, 1954 (U.S.S.R.); 
Bernhard 1931 (Turkey); Cermak et al. 1955 (Czechoslo- 


vakia) ; Coutinho 1939 (Portugal); Fekete and Blattny 1913- 
1914; Fenzel 1932-1933 (Manchuria); Gaussen 1953-1954 
(France), 1956 (France, Spain); Georgescu 1939 (Rumania) ; 
GulisaSvili 1951 (Transcaucasus) ; Heske 1954 (Turkey) ; Hes- 
mer and Feldman 1954 (West Germany); Hultén 1950 
(Northern Europe); Karlberg 1960 (Turkey) ; Kayacik 1954 
(Turkey) ; Ketskhoveli 1960 (Transcaucasus); Murzaev 1955 
(Manchuria); Rubner and Reinhold 1953 (Crimea); Seifriz 
1935 (Transcaucasus); Shimaniuk 1962 (Siberia); Stefanovi¢ 
1958 (Yugoslovia) ; Steven and Carlisle 1959; Touring Club 
Italiano 1958 (Alps) . 

Unpublished—S., Bialobok 1964 (Yugoslavia); H. Gaussen 
1964 (France); General Director of Forestry, Turkey, 1964; 
Instituto Forestal de Investigaciones y Experiencias, Spain, 
1964; B. Kasapligil to N. T. Mirov 1961 (Turkey); H. Kayacik 
1964 (Turkey); JF. Lacaze 1964 (France); E. Magini to N. T. 
Miroy 1960 (Italy); R. Morandini 1964 (Italy) ; C. Moulopou- 
los 1964 (Greece); A. de Philippis 1961, 1964 (Italy); Pilar 
Rifé to N. T. Mirov 1961 (Spain); M. Vidakovic 1964 
(Yugoslavia); J. W. Wright 1960. 


Pinus densiflora Sieb. & Zuce. MAP 33 
Japanese red pine 
Pinus densiflora Sieb. & Zucc., FI. Jap.°2s22, 
t. 112. 1844. 
Pinus densiflora, including P. funebris Komarov, is widely 
distributed in Japan from northern Honshu to Yaku Shima, 


south of Kyushu; from Korea to eastern Manchuria and ad- 
jacent U.S.S.R.; and on the Shantung Peninsula of China. 


Hayashi (1952) maps it at a single locality in southern 
Hokkaido, but this stand was probably planted (M. Kusaka, 
personal communication, 1964) and has been omitted from 
our map. The distribution of P. densiflora on the mainland 


of Asia, especially in southern Manchuria and eastern China, 
is still uncertain, partly because this species has been con- 
fused with several other east-Asian hard pines. Wu (1956) 
shows its range extending westward to include the Liaotung 
Peninsula of southern Manchuria, but Takahasi (1944) states 
that P. densiflora is confined to a small area in the southern 
part of eastern Manchuria. We have omitted the Liaotung 
region from the range shown on the map. 
Sources: 

Published—Akademia Nauk, SSSR, 1954 (U.S.S.R.); Fen- 
zel 1932-1933 (Manchuria) ; Hayashi 1952 (Japan); Liu 1928 
(Shantung); Loesener 1920 (Shantung); Uyeki 1926 (Korea). 

Unpublished—M. Kusaka 1964 (Hokkaido, Korea); N. 
T. Mirov 1961 (Korea). 


Pinus thunbergiana Franco MAP 34 
Japanese black pine 
Pinus thunbergiana Franco, Lisboa Inst. 


Super. Agron. An. 16: 130. 1949. 


Japanese black pine, a maritime species, grows on three 
of the principal islands of Japan—Honshu, Kyushu, and 
Shikoku—and in southern Korea. Its southern limit, accord- 
ing to Hayashi (1952), is Takara Island in the Tokara group 
at about 29° N. latitude, but Wilson (1920) reports that 
Pinus luchuensis is the only pine in this island group. P. 
thunbergiana is very similar to P. nigra of southern Europe 
and the Mediterranean region. This species was long known 
as P. thunbergii Parl. (not Lamb.) . 

Sources: 
Published—Hayashi 1952 (Japan); Uyeki 1926 (Korea). 


Pinus massoniana Lamb. MAP 35 


Masson pine 
Pinus massoniana Lamb., Descr. Genus Pinus 
175 t-b2, 1803: 

Pinus massoniana is widely distributed in China, ranging 
from Shantung, Kiangsu, and northern Taiwan west to 
Szechuan and south to northern Vietnam. A collection from 
the Amoy hills in Merrill’s list of Hainan plants (Merrill 
1927) is probably wrongly attributed to Hainan; F. P. Met- 
calf (1942) lists the same specimen for Fukien Province, 
where the city of Amoy is located. 

We have followed Wu (1956) in including here the little- 
known Pinus henry: Masters (P. henry 
(Masters) Wu). Shaw (1914) considered this pine, found 
in Hupeh and adjoining areas, a variant of P. tabulaeformrs. 

Pinus massoniana is widely planted, and through much of 
its range is the only species of pine present. On many of the 
mountains of eastern and central China it is replaced at 
higher elevations by P. hwangshanensis. 

Sources: 

Published—Bui 1962 (Vietnam); Cheng 1932. (Kwet- 
chow), 1939 (Szechuan); Handel-Mazzetti 1927 and 1929-1956 
(Kweichow) ; Law 1947 (Kiangsu); Liu et al. 1960 (Honan 
F. P. Metcalf 1942 (Fukien); Pendleton 1933 (Kwangtung); 
Steward and Cheo 1935 (Kwangsi):; Wang 1961; Wilson 1913 
(Hupeh); Wu 1956. 

Unpublished—Forest Research Institute, Taiwan, 
T. Mirov 1961 (Taiwan); Kuo Pao-chang 1963 (Shantung). 


massoniana var. 


mensis Hayata MAP 36 


ad pine 

iavwanensis Hayata, Tokyo Col. Sci. Jour. 

30)(Axcts I) 307. LOE 
»ine grows at middle and high elevations in the 
is of central Taiwan. It is closely related to Pinus 
nsis, of the Ryukyu Islands northeast of Taiwan, and 

vangshanensis, of mainland China. 
Source: 

Unpublished—Forest Research Institute, Taiwan, to N. 
Tr. Mirov 1961. 


Pinus luchuensis Mayr MAP 36 


Luchu pine 
Pinus luchuensis Mayr, Bot. Centralbl. 
58: 149, fig. 1894. 

This low-elevation species is confined to the Ryukyu 
Islands between Japan and Taiwan. It is the only pine on 
the islands from Yonaguni, the southwesternmost of the 
Ryukyus, to Amami-Oshima, northeast of Okinawa. This 
species is also the only pine on the Tokara Islands north of 
Amami-Oshima according to Wilson (1920), but Hayashi 
(1952) reports that Pinus thunbergiana occurs as far south as 
Takara, one of the southernmost islands of this group (see 
map 34). 

Sources: 

Published—Wilson 1920. 

Unpublished—F. R. Fosberg 1963; S. Hatusima 1962; E. 
H. Walker 1963. 


Pinus hwangshanensis Hsia MAP 36 


Hwangshan pine 
Pinus hwangshanensis Hsia in ‘Tsoong, Peiping 
Natl. Acad. Inst. Bot. Contrib. 4: 155. 1936. 

This pine has a discontinuous distribution in the higher 
mountains of eastern and central China, extending from 
Chekiang and Anhwei west to Hupeh and Kweichow and 
south to Fukien and Hunan. For many years this species 
was confused with P. tabulaeformis of northern and western 
China (e.g., Law 1947), although it appears to be much more 
closely related to P. luchuensis, the low-elevation pine of the 
Ryukyu Islands. 

Source: 
Published—Wu 1963. 


Pinus tabulaeformis Carr. MAPS 37 AND 38 


Chinese pine 
Pinus tabulaeformis Carr., Traité Gen. Conif. 
Bai yo lONeL8 67: 

This widespread and variable pine occupies a broad, dis- 
continuous belt in China from southeastern Tibet and north- 
ern Yunnan to Inner Mongolia, southern Manchuria, and 
Shantung. It usually grows at moderate elevations, and is 
one of the characteristic trees of the isolated mountain ranges 
in the semiarid northwest of China. The common form of 
this species at high elevations in western Szechuan and north- 
ern Yunnan is Pinus tabulaeformis var. densata (Mast.) Rehd. 
Wu (1947: see Thesis below) has suggested that this variant 
may be a natural hybrid between P. tabulaeformis and P. 
yunnanensis, Dut it extends at least 100 miles north and 
northwest of the northern limits of P. ywnnanensis (Cheng 
1939, Ku and Cheo 1941). 

Sources: 

Published—Cheng 1939 (Szechuan); Chow 1947 (Kansu) ; 
Clark and Sowerby 1912 (Shansi); Cowdry 1922 (Hopeh) ; 
Fox 1949 (Inner Mongolia); Handel-Mazzetti 1927 and 1929- 
1936 (Yunnan, Szechuan); Hao 1938 (Kansu); Hers 1922 


(Honan); Hu 1935 (Honan); Kabanoyv 1962; Ku and Cheo 
1941 (Szechuan); Kung and Wang 1934 (Hopeh) ; Limpricht 
1922; Liou 1931 (Hopeh); Liu 1928 (Hopeh); Marquand 
1929 (Tibet); Orr 1933 (Yunnan); Rehder 1923; Rehder 
and Wilson 1929 (Kansu); Rock 1933 (Kansu); H. Smith 
1925 (Shansi); Takahasi 1936 (Hopeh), 1944 (Liaoning); 
Wang 193 \(Shans1); Waylor ) 1947 @uibes) sa. Memneualoay 
(Kansu) ; Walker 1941 (Kansu, Inner Mongolia); Wang 1961; 
Ward 1936b and 1941a (Tibet); Wilson 1913 (Szechuan); 
Wu 1956; Yang 1937 (Hopeh). 

Thesis—Chung-Lwen Wu 1947. The phytogeographical 
distribution of pine in China. M.F. thesis, Yale Univ., 74 
pp. ilus. 

Unpublished—Wang Chi-Wu 1964 (Hopeh) . 


Pinus yunnanensis Franch. MAP 37 


Yunnan pine 
Pinus yunnanensis Franch., Jour. de Bot. 13: 253, 1899. 


This pine is confined to Yunnan, southern Szechuan, and 
the western parts of Kwangsi and Kweichow. It is closely 
allied to Pinus insularis, which grows at generally lower ele- 
vations to the south and west of P. ywnnanensis, and to the 
variable P. tabulaeformis to the north. 

Sources: 

Published—Cheng 1939; Handel-Mazzetti 1927, 1929- 
1936; Hsu 1950; Ku and Cheo 1941; Rock 1947; Schweinfurth 
1957; Teng 1940; Wang 1961; Ward 1924; Wilson 1926. 

Unpublished—Wang Chi-Wu 1964. 


Pinus insularis Endl. MAP 38 
Khasi pine (see also Map 37) 
Pinus insularis Endl., Synops. Conif. 157. 1847. 


This pine grows in most of the higher parts of southeast 
Asia, from the Khasi Hills of Assam, India, the valleys of 
Zayul in southeastern Tibet, and southwestern Yunnan, to 
the highlands of southern Vietnam and the mountains of 
northern Luzon in the Philippine Islands. Its distribution is 
incompletely known in many parts of southeastern Asia, es- 
pecially the mountains of Laos and northern Vietnam. The 
limits of the species in China are uncertain because of its 
close similarity to Pinus yunnanensis. Wu (1956) considered 
the two a single species, but we have followed the usage of 
most of the botanists familiar with them in the field and 
treated them separately (Handel-Mazzetti 1929-1936; Wang 
1961; Wilson 1926). Intermediates have been reported be- 
tween this species and the allied P. merkusi in the Zambales 
region of Luzon (Perkins 1904) . 

Sources: 

Published—Biswas 1933; Bor 1938 (Assam); Bui 1962 
(Laos) ; Philippine Islands, Bureau of Forestry, 1935 (Luzon) ; 
Burkill 1924 (Burma, Yunnan); Chevalier 1919 (Vietnam); 
Credner 1935 (Thailand); Deb 1960 (Manipur); Ferrars 1875 
(Burma) ; Fischer 1938 (Assam); Foxworthy 1911 (Luzon); 
Handel-Mazzetti 1929-1936 (Yunnan); Larsen 1962 (Thai- 
land); J. G. F. Marshall 1901" (@hailand) = “Men illgg22 
(Luzon); Milton 1960 (Burma); Ogawa et al. 1961 (Thai- 
land); Pendleton 1962 (Thailand); Rao and Panigrahi 1961 
(Assam); Schweinfurth 1957 | (Wibet); PB. Simei si 
(Burma) ; Suvatabandhu 1961 (Thailand); Teng 1948 (Yun- 
nan); Troup 1921 (India, Burma); Vidal 1960 (Laos); Wang 
1961 (Yunnan); Ward 1934 (Tibet), 1936a (Assam), 1941c 
(Burma), 1949 (Burma), 1952a (Manipur), 1959 (Burma) ; 
Whitford 1911 (Luzon); Wilson 1926 (Yunnan, Burma). 

Unpublished—M. Schmid to N. T. Mirov 1961 (Viet- 
nam); Silviculturist, Burma, 1963 (Burma); Wang Chi-Wu 
1964 (Yunnan). 


Pinus merkusii Jungh. & de Vriese MAP 39 


Merkus pine 


Pinus merkusw Jungh. & de Vriese in de Vriese, 
Pl. Nov. Ind. Bat. 5, t. 2. 1845. 


Pinus merkusii ranges from eastern Burma, northern 
Laos, northern Vietnam, and the island of Hainan south to 
Cambodia, southern Vietnam, and the island of Sumatra, and 
east to Luzon and Mindoro in the Philippine Islands. This 
species is the only member of the genus that extends south 
of the Equator. It reaches its southern limit at about 2° S. 
latitude on Sumatra. The distribution of P. merkusii is still 
not completely known, especially on the Asian mainland. Its 
natural occurrence on the island of Hainan is questionable; 
Merrill (1927) indicates that it may have been planted there, 
but both Hosokawa (1940) and Wu (1956) show Hainan as 
part of the natural range of this species. P. merkusii grows 
at low elevations throughout its range, and in many areas is 
associated with and replaced elevationally by P. insularis. 
These two closely related species may hybridize on Luzon 
(see P. insularis) . 

Sources: 

Published—Bui 1962; Philippine Islands, Bureau of For- 
estry, 1935 (Mindoro); Burkill 1924 (Burma, Thailand); 
Chevalier 1944 (Vietnam); Ferrars 1875 (Burma); Luytjes 
1924 (Sumatra); Maurand 1943; Merrill 1927 (Hainan); 
Royal Forest Department, Thailand, 1962 (Thailand) ; Ryan 
and Kerr 1911 (Thailand); P. Singh 1913 (Burma); van 
Steenis 1958 (Sumatra); Suvatabandhu 1961 (Thailand); 
Vidal 1960 (Laos); Wilson 1926 (Burma, Thailand) . 

Unpublished—W. S. Astle to N. T. Mirov 1961 (Cam- 
bodia); N. T. Mirov 1961 (Luzon, Sumatra); M. Schmid to 
N. T. Mirov 1961 (Vietnam); Silviculturist, Burma, 1963 
(Burma) . 


SUBSECT. AUSTRALES Loud. 


Pinus palustris Mill. MAP 40 


longleaf pine 


Pinus palustris Mill., Gard. Dict. Ed. 8, 
Pinus No. 14. 1768. 


Longleaf pine is limited to the Coastal Plain of south- 
eastern United States from southeastern Virginia to central 
Florida and west to eastern Texas. It hybridizes in nature with 
at least two of its principal associates, Pinus taeda (Chapman 
1922) and P. elliottii (Mergen 1958). 


Pinus taeda L. MAP 41 


loblolly pine 
Pinus taeda L., Sp. Pl. 1000. 1753. 


Loblolly pine ranges through the Coastal Plain and Pied- 
mont of eastern and southeastern United States from Dela- 
ware south to central Florida, west to eastern Texas, and 
north in the Mississippi Valley to southeastern Oklahoma, 
Arkansas, and southern Tennessee. It is associated with sev- 
eral related species, and is believed to hybridize in nature 
with at least four of them—Pinus palustris (Chapman 1922) 
P. echinata (Zobel 1953), P. serotina (Wenger 1958), and P. 
rigida’. 

Additional source: 

Published—Bethune 1963 (Fla.). 


6 Committee on Southern Forest Tree Improvement Newsletter IV (1). 


May 1955. (Processed) . 


Pinus echinata Mill. MAP 42 
shortleaf pine 
Pinus echinata Mill., Gard. Dict. Ed. 8, 
Pinus No. 12. 1768 

Shortleaf pine extends from Long Island, New York, and 
New Jersey to Pennsylvania, southern Ohio, southern Illinois, 
and southern Missouri, south to eastern Oklahoma and east- 
ern Texas, and east to northern Florida and Georgia. The 
most widespread of the southern pines, this species occupies 
large areas of Coastal Plain, Piedmont, and mountains from 
the Appalachians to the Ozarks. It crosses naturally with at 
least two associated pines, Pinus rigida (Austin 1928) and P. 
taeda (Zobel 1953). 
Additional sources: 

Published—Aughanbaugh 1950 (Pa.); Haney 1955; Lim- 
ing 1946 (Mo.); Perry 1924 (Pa.); Shafer and Chisman 1957 
(Paine 


Pinus glabra Walt. MAP 43 


spruce pine 
Pinus glabra Walt., Fl. Carol. 237. 1788. 

Spruce pine is confined to the Coastal Plain of southeast- 
ern United States from South Carolina to northern Florida 
and west to southeastern Louisiana. A minor constituent of 
the pine forests, it is associated with Pinus taeda, P. elliottit, 
and others. 


Pinus rigida Mill. MAP 44 
pitch pine 
Pinus rigida Mill., Gard. Dict. Ed. 8, 
Pinus No. 10. 1768. 


Pitch pine ranges from central Maine to New York, ex- 
treme southwestern Quebec, and extreme southeastern On- 
tario, south to Pennsylvania, Ohio, and Delaware, and south 
in mountains to eastern Tennessee, northern Georgia, and 
western South Carolina. From near sea level northward, this 
species extends southward into higher altitudes along the 
entire length of the Appalachian Mountains. It often grows 
on poor sites, and is the characteristic tree of the pine barrens 
of New Jersey. 

It is associated with several related pines in the southern 
part of its range, and is thought to cross in nature with Pinus 
echinata (Austin 1928) and P. taeda’. Its closest relative, 
however, is P. serotina. These two very similar pines inter- 
grade in Delaware’ and southern New Jersey (Clausen 1939), 
but it is uncertain whether this intergradation is due to hy- 
bridization or to incomplete divergence of the two. 

Additional sources: 

Published—Grandtner 1961 (Que.); Illick and Aughan- 
baugh 1930 (Pa.); Rouleau 1955 (Que.). 

Unpublished—T. C. Brayshaw and A. E. Porsild 1964 
(Canada). 


Pinus serotina Michx. MAP 44 


pond pine 
Pinus serotina Michx., Fl. Bor.-Amer. 2: 205. 18053. 

Pond pine is restricted to the Coastal Plain of south- 
eastern United States from southern New Jersey and Delaware 
south to central and northwestern Florida and central Ala- 
bama. 

It is thought to cross naturally with one of its commonest 
associates, Pinus taeda (Wenger 1958). At the northern end 
of its range it overlaps and intergrades with P see J 
rigida). 


7 Committee on Southern Forest Tree Improvement Newsletter TV (1). 


May 1955. (Processed) . 


ns Lamb. MAP 43 
n pine 
bungens Lamb., Ann. Bot. 2: 198. 1805. 


\fountain pine is largely confined to the Appala- 
untains of eastern United States from eastern and 
“ansyivania southwest to eastern Tennessee and 
ern South Carolina, with outposts in Kentucky and 

n Georgia. This species is remarkable among the 
pines of this series for its heavily armed cones. 


Pinus elliottii Engelm. MAP 45 
slash pine 
Pinus elliottti Engelm., Acad. Sci. St. Louis Trans. 
4: 186, t. 1-3. 1880. 

Slash pine is restricted to the Coastal Plain of southeast- 
ern United States from southern South Carolina to southern 
Florida, including the Florida Keys, and west to southeastern 
Louisiana. A major constituent of the lower Coastal Plain 
pine forests, slash pine has two geographical varieties, both 
mapped. South of the widespread typical variety, (var. el- 
liotti), South Florida slash pine (Pinus elliottii var. densa 
Little & Dorman) extends through central and southern 
Florida to the Keys. This species is separated from the nearest 
stands of the closely related P. caribaea, on Grand Bahama 
Island, by about 70 miles. 

Among the several related pines with which it is as- 
sociated, slash pine is believed to hybridize naturally with 
Pinus palustris (Mergen 1958). Perry and Wang (1957) have 
speculated that P. elliottii var. densa, notable for the modified 
grass stage of its seedlings, may have originated through in- 
trogressive hybridization of these two species. 

Additional sources: 

Published—Little and 
(Fla.). 


Dorman 1954; Langdon 1963 


Pinus caribaea Morelet MAP 46 
Caribbean pine 
Pinus caribaea Morelet, Rev. Hort. Céte d’Or 


LO als oil: 


Caribbean pine occurs on several of the Bahama Islands, 
in Pinar del Rio Province of western Cuba, on the Isle of 
Pines, and along the Caribbean seaboard of Central America 
from British Honduras to Nicaragua. As formerly inter- 
preted, this species included slash pine of the southeastern 
United States, now considered distinct as P. elliottii (Little 
and Dorman 1954). 

In its island distribution the only other pine with which 
Caribbean pine is associated is Pinus tropicalis—in western 
Cuba and on the Isle of Pines. In interior Honduras and 
Nicaragua, its range overlaps that of P. oocarpa. The inland 
form of P. caribaea in this region tends to resemble P. oocarpa, 
suggesting the possibility of natural hybridization (Denevan 
1961, p. 276; L. Williams 1955). 

The southernmost outlier of Pinus caribaea on the east 
coast of Nicaragua, near the town of Bluefields, is the most 
southerly locality reached by any pine in the Western Hemi- 
sphere. 

The presence of Pinus cavibaca in Quintana Roo, Mexico 
(Standley and Steyermark 1958), has not been verified and is 
not shown on the map, 

Sources: 

Published—Barrett and Golfari 1962; Denevan 1961 
(Nicaragua); Directorate of Overseas Surveys 1958 (British 
Honduras); Helbig 1957 and 1959 (Honduras); Little and 
Dorman 1954 (Bahama Islands); Lundell 1961 (Guatemala); 
March 1949 (Bahama Islands); Marie-Victorin and Léon 1942 
(Cuba) ; E. E. Smith 1954 (Cuba); Standley and Steyermark 
1958 (Guatemala). 


16 


Thesis—Jeffrey Radley 1960. The physical geography 
of the east coast of Nicaragua. M.A. thesis, Univ. of Calif., 
Berkeley, Calif., 188 pp., illus. 

Unpublished—C. L. Ludwig (undated map of Guate- 
mala) . 


Pinus occidentalis Sw. MAP 46 
West Indian pine 
Pinus occidentalis Sw., Nov. Gen. Sp. Pl. 103. 1788. 
This species is restricted to the island of Hispaniola, 
where it is the only pine, and the Sierra Maestra of eastern 


Cuba. It grows at a wide range of elevations, but is more 
common at high elevations. 
Sources: 

Published—Ciferri 1936 (Dominican Republic); Denevan 


1961 (Haiti); E. E. Smith 1954 (Cuba); Street 1960 (Haiti). 


Pinus cubensis Griseb. MAP 46 


Cuban pine 
Pinus cubensis Griseb., Amer. Acad. Mem., 
Ser. 2, 8: 530. 1862. 


This low-elevation pine is limited to the eastern part of 
Oriente Province, Cuba. It may extend to the Sierra Maestra 
of this province, where the closely related Pinus occidentalis 
is widespread, but this extension has not been verified (E. E. 
Smith 1954) and is not shown on the map. 

Source: 
Published—E. E. Smith 1954. 


SUBSECT. PONDEROSAE Loud. 


Pinus ponderosa Laws. MAP 47 
ponderosa pine 
Pinus ponderosa Laws., Agr. Man. 354. 1836. 


Ponderosa pine, including its variants Pinus ponderosa 
var. scopulorum Engelm. and P. ponderosa var. arizonica 
(Engelm.) Shaw (P. arizonica Engelm.), extends from south- 
ern British Columbia east to southwestern North Dakota and 
central Nebraska, and south to southern California, northern 
Durango, and San Luis Potosi. Within this enormous range 
this variable yellow pine is unexplainably absent from a large 
area that includes southern Idaho, western Wyoming, south- 
western Montana, and part of the Great Basin. In California 
ponderosa pine extends south almost to the Mexican border 
but has not been found in northern Baja California (Dufheld 
and Cumming 1949). 

In the Sierra Madre Occidental of northwestern Mexico, 
Pinus ponderosa var. arizonica grows with other yellow pines 
(P. engelmanni, P. durangensis) which link the P. ponderosa 
complex to the group of yellow pines concentrated in central 
Mexico (P. montezumae, P. pseudostrobus, and others). In 
California ponderosa pine is associated with P. jeffrey: at 
upper elevations, and the two hybridize in nature (Haller 
1962). Ponderosa pine may also intergrade with P. washoen- 
sis (Haller 1961). 

Our map is based primarily upon a detailed map of the 
species prepared by R. Z. Callaham. 

Additional sources: 

Published—Ayres 1900a and 1900b (Mont.); Bailey and 
Bailey 1941; Costello and Schwan 1946 (Colo.); Deaver and 
Haskell 1955 (Ariz.); Haller 1962 (Calif.); Johnston 1943 
(Coah.) ; Leiberg 1900b (Oreg.), 1904a (Mont.); J. T. Mar- 
shall 1957 (Son., Chih.) ; Merriam 1893 (Nev.); Potter and 
Green 1964 (N. Dak.); Rogers 1953; G. C. Rzedowski 1960 
(S.L.P.); J. Rzedowski 1956 (S.L.P.); Sudworth 1900a and 
1900b (Colo.); ‘Townsend 1895 (N. Mex.); Zobel and Cech 
1957 QNDE 


Unpublished—W. D. Billings 1961 (Nev.); T. C. Bray- 
shaw and A. E. Porsild 1964 (B.C.); W. P. Cottam 1964 
(Utah); J. D. Curtis 1964 (Utah) ; J. F. Franklin 1964 (Oreg.); 
J. R. Haller 1964 (Calif.); F. D. Johnson 1964 (Idaho); F. 
Kramer to N. T. Mirov (n.d.) (Nev.); O. V. Matthews 1964 
(Ores) eRe AY Read? 1964; R> Schmidt 1964 ((B:G.); W. 1: 
Stein 1964; R. F. Wagle 1962 (Ariz.); O. Zarzosa L. 1964 


(Dgo.). 
Pinus washoensis Mason & Stockwell MAP 48 
Washoe pine 
Pinus washoensis Mason & Stockwell, Madrono 
8: 62. 1945. 


Not distinguished until 1945, Washoe pine is known only 
from limited areas in three mountain ranges at the western 
edge of the Great Basin in Nevada and California. It oc- 
cupies a few square miles on the east slopes of Mt. Rose, Nev., 
and has recently been discovered in the southern Warner 
Mountains of northeastern California (Haller 1961) and on 
the Bald Mountain Range (Calif.), about 20 miles northwest 
of Mt. Rose (Critchfield and Allenbaugh 1965). In all three 
localities it grows at higher elevations than Pinus ponderosa, 
its closest relative. According to Haller, it may also be oc- 
casional in a variable population of P. ponderosa and P. 
jeffreyi near Hobart Mills, Calif., about 14 miles west of 
Mt. Rose. 

Sources: 

Published—Critchfield and Allenbaugh 1965; Haller 1961; 

Mason and Stockwell 1945. 


Pinus jeffreyi Grev. & Balf. MAP 48 
Jeffrey pine 
Pinus jeffreyi Grev. & Balf. in A. Murr., Bot. 
Exped. Oreg. [ Rpt. No. 8] 2, t. 1853. 

Primarily a California species, Jeffrey pine extends into 
southwestern Oregon, western Nevada, and northern Baja 
California. It usually grows at higher elevations than the 
closely related Pinus ponderosa, but the two overlap broadly 
and hybridize in nature (Haller 1962). Our map is based 
on a distribution map prepared by D. F. Roy and modified 
from the sources of information listed below. 

Additional sources: 

Published—Haller 1962. 

Thesis—Donald V. Hemphill 1952. The vertebrate 
fauna of the boreal areas of the southern Yolla Bolly Moun- 
MMe California. Ph.D. thesis, Oregon State College, 340 pp., 
illus. 

Unpublished—B. K. Ford 1963 (Calif.) ; J. R. Griffin 1964 
(Calif.); D. V. Hemphill 1964 (Calif.); J. L. Jenkinson 1963 
(Calif., Oreg.); Reid Moran 1964 (Baja Calif.). 


Pinus engelmannii Carr. MAP 49 
Apache pine 
Pinus engelmannii Carr., Rev. Hort., Sér. 4, 
3: 227. 1854; “engelmanni.” 

Apache pine extends from southeastern Arizona and ex- 
treme southwestern New Mexico through the Sierra Madre 
Occidental of Mexico to Zacatecas and Aguascalientes. 

This species has also been known as Pinus latifolia Sarg., 
P. apacheca Lemm., and P. macrophylla Engelm. It usually 
grows at lower elevations than the related yellow pines of this 
region (P. ponderosa, P. durangensis, P. cooper). 

Additional sources: 

Published—Gentry 1946 (Sin.); Guzman and Vela 1960 
(Zac., Ags.) ; J. 1. Marshall 1957 (Son., Chih.). 

Thesis—James H. Maysilles 1959. Floral relationships of 
the pine forests of western Durango, Mexico. Ph.D. thesis, 
Univ. Michigan, 165 pp., illus. 

Unpublished—O, Zarzosa L. 1964 (Dgo.); C. E. 
1964 (Dgo.). 


Blanco 


Pinus durangensis Martinez MAP 50 
Durango pine 
Pinus durangensis Martinez, Mex. Inst. Biol. An. 
ligre24, fio. 1-4. 1942 

This species is confined to Mexico, ranging from eastern 
Sonora to southern Durango, with outliers in Michoacan and 
Nuevo Leon. It is an important constituent of the yellow 
pine forests in the Sierra Madre Occidental. 
Additional sources: 

Published—J. T. Marshall 1957 
(Chih.) ; Zobel and Cech 1957 (N.L.). 

Thesis—James H. Maysilles 1959. Floral relationships 
of the pine forests of western Durango, Mexico. Ph.D. thesis, 
Univ. Michigan, 165 pp., illus. 

Unpublished—Mexico, Instituto Nacional de Investiga- 
ciones Forestales 1964 (Ags.); N. Sanchez Mejorada to L. W. 
Bryan 1963 (Mich.) ; O. Zarzosa L. 1964 (Dgo.). 


Wicht 1949 


5on.); 


Pinus cooperi C. E. Blanco MAP 51 


Cooper pine 
Pinus cooperi C. E. Blanco, Mex. Inst. Biol. An. 
202,185; -fig; 1_ 1949: 

This species grows at moderate elevations in the exten- 
sive pine forests of southern Chihuahua and western Durango. 
It is associated with several other species in the western yel- 
low pine group, including Pinus ponderosa (P. arizonica 
Engelm.) , P. engelmannii, and P. durangensis. P. coopert was 
originally named P. lutea C. E. Blanco (not Walt.) . 
Additional sources: 

Unpublished—C, E. Blanco 1964 (Dgo.); O. Zarzosa L. 
1964 (Dgo.); E. Hernandez X. and J. Vasquez Soto 1964 
(Chih.). 


Pinus montezumae Lamb. MAP 49 


Montezuma pine 
Pinus montezumae Lamb., Descr. Genus Pinus. Ed. 
3m(8 2) ls 59, tei22: 1832. 

This variable species is a major constituent of the pine 
forests of central Mexico, ranging from Jalisco to Veracruz 
and south to Oaxaca, Chiapas, and Guatemala. It is also 
present in the Sierra Madre Oriental, and its northernmost 
known occurrence is in an outlier of this range, the Sierra 
Picachos in Nuevo Leon (Zobel and Cech 1957). In various 
parts of its range it is associated with many of the other Mex- 
ican yellow pines, and it may intergrade with some of them 
(Loock 1950, p. 180) . 

Additional sources: 

Published—Aguilar 1961 (Guatemala); Goldman 1951; 
Hernandez X. et al. 1951 (Tamps.); Leavenworth 1946 
(Mich.) ; Loock 1950 (Mich.); Schwerdtfeger 1953 (Guate- 
mala) ; Zobel and Cech 1957 (N.L.). 

Unpublished—B. Hallberg 1964 (Oax.); E. Larsen 1962; 
F. Medellin-Leal 1964 (S.L.P.); N. Sanchez Mejorada to L. 
W. Bryan 1963 (Mich.). 


Pinus hartwegii Lindl. MAP 51 
Hartweg pine 

Pinus hartwegit Lindl., Bot. Reg. v. 25, Misc. 62. 1839. 

Pinus hartwegtt, including the doubtfully distinct P. 
rudis Endl., has a markedly discontinuous distribution at high 
elevations from Chihuahua and Nuevo Leén south to Guate- 
mala and El Salvador. It often reaches timberline, but at 
lower elevations it overlaps and may intergrade with P 
zumae (Standley and Steyermark 1958) . 
Additional sources: 

Published—Aguilar 1961 (Guatemala); Hinds and Lar 
sen 1961 (Mex.); Leavenworth 1946 (Mich Loock 1950 


anda 1957 (Chis.); Schwerdtfeger 1953 (Guate- 

909 (Chih.); Sociedad Botanica de México 1960 

‘ley and Steyermark 1958 (Guatemala); Zobel 
CNEWEE 

lished—P. H. Allen 1962 (EI Salvador); B. Hall- 

{ (Oax.); Mexico, Instituto Nacional de Investiga- 

restales 1964 (Gro.); E. Larsen 1962; R. McVaugh 


Lo). 


i michoacana Martinez MAP 50 
Michoacan pine 
Pinus michoacana Martinez, Mex. Inst. Biol. An. 
libel tro 4s Oa 

This pine grows at relatively low elevations in central 
and southern Mexico, from southern Durango and San Luis 
Potosi southeast to Chiapas. It is sometimes associated with 
its close relative, Pinus montezumae. 
\dditional sources: 

Published—Guzman and Vela 1960 

(Mich.) . 

Unpublished—L. W. Bryan 1964  (Oax.); B. Hallberg 
1964 (Oax., Chis.); E. Larsen 1962 (Mich., Dgo.) ; N. Sanchez 
Mejorada to L. W. Bryan 1963 (Mich.). 


(Zac.) ; Loock 1950 


Pinus pseudostrobus Lindl. MAP 52 


Pinus pseudostrobus Lindl., Bot. Reg. v. 25, 
Misc. 63. 1839. 


Pinus pseudostrobus, including the doubtfully distinct 
P. tenuifolia Benth. (not Salisb.) and P. oaxacana Mirovy, is 
widespread at moderately high elevations from Sinaloa and 
Nuevo Leon through central and southern Mexico, and in 
Central America as far south as northwestern Nicaragua. This 
variable species complex extends farther south than any other 
western yellow pine. In the western part of its range it over- 
laps the closely related P. douglasiana. 

Additional sources: 

Published—Aguilar 1961 (Guatemala); Allen 1955 (Hon- 
duras); Castanos 1962 (Oax.); Denevan 1961 (Nicaragua) ; 
Gentry 1946 (Sin.); Goldman 1951 (Mich.); Leavenworth 
1946 (Mich.); Loock 1950 (Mich.); Miranda 1961 (Chis.); 
Miranda and Sharp 1950 (Pue.); Pfeifer 1960 (Honduras) ; 
Shaw 1909; Sociedad Botanica de México 1960 (Ver.); Vogel 
1954 (Honduras) ; Wicht 1949 (Mich.); Zobel and Cech 1957 
(NES 

Unpublished—P. H. Allen 1962 (El Salvador); B. Hall- 
berg 1964 (Oax.); N. Sanchez Mejorada to L. W. Bryan 1963 
(Mich.) . 


Pinus douglasiana Martinez MAP 532 


Douglas pine 
Pinus douglasiana Martinez, Madrono 7: 4, t.l. 1943. 


This rather uncommon pine is widely scattered at middle 
elevations in western Mexico from extreme southern Sonora 
to Michoacan and México. Its range slightly overlaps that of 
its closest relative, the highly variable Pinus pseudostrobus. 
Additional sources: 

Unpublished—B. Hallberg 1964 (Oax.) ; 
1964 (Dgo.). 


Cc. E. Blanco 


Pinus teocote Schiede & Deppe MAP 33 
Pinus teocote Schiede & Deppe in Schlecht. & Cham., 
Linnaea 5: 76. 1830. 

Pinus teocote, including the weakly segregated P. herrerai 
Martinez, is one of the most widely distributed Mexican 
pines, ranging from Chihuahua and Coahuila south to 
Chiapas, and just entering western Guatemala. In southern 
Mexico it overlaps P. lawsonii, a related species. 


18 


Additional sources: 

Published—Aguilar 1961 (Guatemala); Matuda 1950 
(Chis.) ; J. Rzedowski 1956 (S.L.P.); G. C. Rzedowski 1960 
(S.E-P.);) Shaw) 1909 (SE: P);" Wicht7 1949) 5 ((Michs)p- rele VValle 
liams 1939 (Oax.). 

Thesis—James H. Maysilles 1959. Floral relationships 
of the pine forests of western Durango, Mexico, Ph.D. thesis, 
Univ. Michigan, 165 pp., illus. 

Unpublished—C. E. Blanco 1964 (Coah.); B. Hallberg 
1964 (Oax.); F. Medellin-Leal 1964 (S.L.P.); O. Zarzosa EL. 
1964 (Dgo.). 


Pinus lawsonii Roezl MAP 53 
Lawson pine 
Pinus lawson Roezl ex Gord., Pinet. Sup. 64. 1862. 


This pine is confined to southern Mexico from Jalisco 
to Veracruz and south to Oaxaca. Its range overlaps that 
of the related Pinus teocote. 

Additional sources: 

Published—Loock 1950 (Mich.) . 


Unpublished—B. Hallberg 1964 (Oax.); N. Sanchez 
Mejorada to L. W. Bryan 1963 (Mich.) . 
SUBSECT. SABINIANAE Loud. 
Pinus sabiniana Dougl. MAP 54 


Digger pine 
Pinus sabiniana Dougl. ex D. Don in Lamb., Descr. 
Genus Pinus. Ed. 3 (8°), v. 2, unnumbered p. 
between p. 144 and p. 145, t. 80. 1832; 
Dougl., Linn. Soc. London Trans. 16: 749. 1833. 


Digger pine is confined to California. It 1s characteristic 
of the dry foothills of the Sierra Nevada and the Coast Range. 
Sources: 

Published—Griffin 1964. 

Unpublhished—]. R. Griffin 1964. 


Pinus coulteri D. Don MAP 55 


Coulter pine 
Pinus coultert D. Don, Linn. Soc. London Trans. 
17: 440. 1836. 

Coulter pine is restricted to the coastal mountains of 
central and southern California and northern Baja California. 
It overlaps two related species, Pinus jeffreyi and P. sabiniana, 
and is known to hybridize in nature with P. jeffrey: (Zobel 
1951). The distribution of the species in California is based 
entirely on Vegetation-Type survey information. ‘The sources 
listed below all pertain to Baja California. 

Additional sources: 
Published—Goldman 1916; Martinez 1948. 
Unpublished—Reid Moran 1961; E. L. Sleeper 1964. 


Pinus torreyana Parry MAP 55 
Torrey pine 
Pinus torreyana Parry ex Carr., Traite Gen. 
Conif. 326. 1855. 

Torrey pine has a very restricted range in southern 
California. On the mainland it is confined to the low coastal 
bluffs flanking the Soledad Valley north of San Diego.  Else- 
where it occurs only as a small grove on Santa Rosa Island. 
The mainland distribution shown on the map is based on 
Vegetation-Type mapping done in 1931; the present range 
of the species may be slightly less than that shown. 
Additional source: 

Unpublished—E. R. Blakley 1961 (Santa Rosa Island). 


SUBSECT. CONTORTAE Little & Critehfield ~ 


Pinus banksiana Lamb. MAP 56 
jack pine 
Pinus banksiana Lamb., Descr. Genus Pinus 
Le Fs Tee iat 1tetO) 3}. 

Primarily a Canadian species, jack pine extends from 
north of 65° N. latitude along the Mackenzie River in west- 
ern Canada south and east to the Lake States, northern New 
York and New England, and the Maritime Provinces of 
Canada. In Alberta, where this species overlaps the closely 
related Pinus contorta, natural hybridization is common (Moss 
1949, Mirov 1956). 

Additional sources: 

Published—Baldwin 1961 (N.H.); Bentley and Smith 
1960 (N.S.); Rudolf and Schoenike 1963 (U.S.); Rudolph, 
Libby, and Pauley 1957 (Minn.); Schoenike 1962 (Minn.). 

Thesis—Roland E. Schoenike 1962. Natural variation 
in jack pine (Pinus banksiana Lambert). Ph.D. thesis, Univ. 
Minn., 252 pp., illus. 

Unpublished—T. C. Brayshaw and A. E. Porsild 1964 
(Canada); E. W. Littlefield 1952 (N.Y.). 


Pinus contorta Dougl. MAP 56 
lodgepole pine 
Pinus contorta Doug]. ex Loud., Arb. Frut. Brit. 
e292 es 22 1022111838. 

One of the most widespread of the American pines, lodge- 
pole pine extends from central Yukon in western Canada 
south to southern Colorado and northern Baja California. 
Within these limits lodgepole pine grows at a wider range of 
elevations than any other pine. Common just above sea 
level along the Pacific coast, it reaches about 11,000 feet in 
the southern Sierra Nevada and Rocky Mountains. The 
morphological diversity of this wide-ranging species has been 
investigated by Critchfield (1957). 

The known range of lodgepole pine in western Canada 
has only recently been extended to the Mackenzie District, 
where it has been found in the Liard Mountains and else- 
where in the southwestern part of the district (Jeffrey 1959; 
T. C. Brayshaw and A. E. Porsild, personal communication, 
1964) . Another outlier, in the Birch Mountains of northeast- 
ern Alberta, was also discovered recently (A. H. Marsh, per- 
sonal communication, 1964) . 

The only close relative of lodgepole pine in western 
North America is Pinus banksiana. The two overlap and 
hybridize in Alberta (see P. banksiana), and introgression 
into lodgepole pine is likely (Mirov 1956; Critchfield 1957). 
Additional sources: 

Published—Jeffrey 1959 
(Oreg.). 

Unpublished—T. C. Brayshaw and A. E. Porsild 1964 
(Canada); J. F. Franklin 1964 (One; Wash.)i:) J. de. eee 
son 1963 (Calif., Oreg.); F. D. Johnson 1964 (Idaho); A. 
Marsh 1963 (Alta.); O. V. meters [964 (Oren. |G O! cae 
1964 (Calif.) . 


(Canada); Leiberg 1900b 


*Subsect. Pinus subgen. Pinus sect. Pinus subsect. Contortae Little 
& Critchfield, subsect. nov. Folia 2 in fasciculo, brevia (2-9 cm. longe) , 
hypodermide biforme, ductis resiniferis maxime ex parte medialibus. 
Ramuli verni multinodales. Strobili parvi (3-8 cm. longe) , symmetricales 
vel obliqui, plerumque clausi vel serotini aperti, longe  persistentes, 
squamae plerumque cum aculeo persistente. Holotypus: Pinus contorta 
Dougl. ex. Loud., Arb. Frut. Brit. 4: 2292, fig. 2210-2211. 1838. 


Pinus virginiana Mill. MAP 57 
Virginia pine see also Map 
Pinus virginiana Mill., Gard. Dict. Ed. 8, 
Pinus No. 9. 1768 

Virginia pine is widespread in and near the Appalachian 
Mountains and the Piedmont region east the Appalachians, 
ranging from Long Island, New York, south and east to cen- 
tral Alabama, western Tennessee, and southern Indiana and 
Ohio. It is separated from the closely related Pinus clausa 


about 200 miles. 


by a 100-mile gap and from P. banksiana 
Additional source: 
Published—Ross 1951 (Ind.) 


Pinus clausa (Chapm.) Vasey MAP 57 
sand pine (see also Map 5 
Pinus clausa (Chapm. ex Engelm.) Vasey ex Sarg., 
U. S. Census, 10th, 1880, v. 9 (Rpt. Forests 
No. Amer.): 199. 1884. 

Sand pine grows on the sandy plains of Florida, just 
extending into the southern tip of Alabama. It is separated 
from the southernmost outliers of its closest relative, Pinus 
virginiana, by about 100 miles. 

Additional source: 
Published—R. W. Cooper 1957. 


SUBSECT. OOCARPAE Little & Critchfield 


Pinus radiata D. Don MAP 58 
Monterey pine 
Pinus radiata D. Don, Linn. Soc. London Trans. 
442. 1836. 

The most widely planted of the pines, Monterey pine is 
restricted in nature to three coastal localities in central Cali- 
fornia. A related form; Pinus radiata var. binata Lemm., 
grows on Guadalupe Island, more than 150 miles from the 
coast of Baja California and nearly 500 miles from the nearest 
mainland population of P. radiata. Unlike mainland P. 
radiata, the island form grows at mederate elevations (1,300- 
4,000 feet according to Reid Moran, personal communica- 
tion, 1961). On the mainland, P. radiata occurs with P. at- 
tenuata and P. muricata and is reported to hybridize occasion- 
ally with both (Stebbins 1950, p. 209-210). 

Additional source: 
Unpublished—Reid Moran 1961 (Guadalupe Island) . 


Pinus attenuata Lemm. MAP 58 
knobcone pine 
Pinus attenuata Lemm., Mining and Sci. Press 
64: 45. 1892. 

Knobcone pine, remarkable for its universally closed 
cones, is distributed patchily at low and middle elevations in 
the mountains of California and southwestern Oregon, with 
a single known locality in northern Baja California. It over- 
laps the closely related coastal species, Pinus radiata, at 
place in central California, and occasional natural hybrids 
have been observed in this locality (Stebbins 1950, p. 209 


9Subsect. Pinus subgen. Pinus sect. Pinus subsect 


& Critchfield, subsect. nov. Folia plerumque 3 (2-5) in fas 

dermide plerumque biforme, ductis resiniferis maxime ex 

bus, interdum internalibus vel septalibus. Ramuli verni m: dales vel 
uninodales. Strobili plerumque obliqui, clausi, longe persistentes, squamac 
cum aculeo vel protuberantes. Holotypus: P Schiede it 


Schlecht., Linnaea 12: 491 IS3s 


IYCES: 


—Howitt and Howell 1964 (Calif.) . 
rene B. Newcomb 1962. Geographic variation 
cata Lemm. Ph. D. thesis, Univ. Calif., Berke- 
917 pps) ulus: 
rlished—B. K. Ford 1963 (Oreg.); J. F. Franklin 
reg.); J. R. Griffin 1963, 1964 (Calif.); J. L. Jenkin- 
; O. V. Matthews 1964 (Oreg.); R. B. Vasey 1963 


Pinus muricata D. Don MAP 59 
bishop pine 
Pinus muricata D. Don, Linn. Soc. London Trans. 
17: 441. 1836. 

Included here are Pinus remorata Mason and P. muricata 
var. cedrosensis Howell, although the status of the latter pine 
in particular is uncertain (Fielding 1961). Bishop pine, an 
extremely variable species complex, occurs in seven rather 
widely separated localities on the coast of California and 
northern Baja California, and on three islands situated 15 to 
30 miles offshore. Its distribution may overlap that of the 
closely related P. attenwata near the coast of Mendocino 
County, Calif., and the two species grow less than 10 miles 
apart in the vicinity of San Luis Obispo, Calif. On the 
Monterey peninsula near Monterey, Calif., P. muricata is 
associated with P. radiata, and natural hybrids have been 
observed (Stebbins 1950, p. 209). A reported occurrence of 
P, muricata near Crescent City, Calif., based on a single tree 
found in the coastal forest, is not shown on the map (Mason 
1949; H. L. Mason, personal communication, 1964) . 
Additional sources: 

Published—Epling and Robison 1940 (Baja Calif.) ; 
Howell 1949 (Calif.); Mason 1949 (Calif.); W. Metcalf 1921 
(Calif.); Muller 1962 (Baja Calif.). 

Theses—John W. Duffeld 1951.  Interrelationships of 
the California closed-cone pines with special reference to 
Pinus muricata D. Don. Ph.D. thesis, Univ. Calif., Berkeley, 
Calif, 77 pp. illus.; George O. Hale 1941. A survey of the 
vegetation of Cedros Island, Mexico. M.A. thesis, Univ. 
Calif., Los Angeles, Calif., 96 pp., illus. 

Unpublished—E. R. Blakley 1961 (Santa Cruz and Santa 
Rosa Islands); W. J. Libby 1964 (Cedros Island); P. Zinke 
1961 (Calif.) . 


Pinus patula Schiede & Deppe MAP 60 
Mexican weeping pine 
Pinus patula Schiede & Deppe in Schlecht. & Cham., 
Linnaea 6: 354. 1831. 
Pinus patula is restricted to eastern Mexico from Tamau- 
lipas to Oaxaca. Its presence in Tamaulipas was reported 
only recently (Hernandez X. et al. 1951). 


A variant in Chiapas referred tentatively to var. longi- 


pedunculata Loock may be identical with P. oocarpa var. 
ochoterenai Martinez (E. L. Little, Jr., 17957, 10 km. S. E. of 
San Cristobal de Las Casas: B. Hallberg, personal communica- 
tion, 1964). The range of the Chiapas highlands pine is 
shown separately on Map 61. 
Additional sources: 
Published—Hernandez X. et al. 1951 
1949 (Hgo.). 
Unpublished—B. Hallberg 1964 (Oax., Chis.). 


(Tamps.); Wicht 


Pinus greggii Engelm. MAP 60 
Gregg pine 
Pinus greggii Engelm. ex Parl. in DC., Prodr. 
16 (2). 396. 1868. 

This uncommon species‘is confined to eastern and north- 
eastern Mexico from southeastern Coahuila to Hidalgo. Its 
range overlaps that of the closely related Pinus patula in 
eastern Hidalgo. 

Additional sources: 

Published—Johnston 1943 (Coah.) . 

Unpublished—J. W. Andresen 1964 (N.L.); E. Larsen 
(Hgo.); F. Medellin-Leal 1964 (S.L.P.). 


Pinus oocarpa Schiede MAP 61 
Pinus oocarpa Schiede in Schlecht., Linnaea 12: 491. 1838. 


One of the most widespread Latin American pines, Pinus 
oocarpa ranges in Mexico from Sonora southeast to Hidalgo 
and Chiapas, and in Central America east and south to 
British Honduras and northwestern Nicaragua. It grows at 
relatively low elevations, and extends farther south than any 
other American pine except P. caribaea. P. oocarpa var. 
ochoterenai Martinez is shown separately because of its un- 
certain status (see P. patula). 

Pinus tecumumani of Guatemala, which was proposed 
without a Latin diagnosis by Schwerdtfeger (1953), is con- 
sidered by Standley and Steyermark (1958) a variant of P. 
oocarpa, and is not shown separately on the map. 

Additional sources: 

Published—Denevan 1961 (Nicaragua); Gentry 1946 
(Sin.) ; Goldman 1951; Hunt 1962 (British Honduras) ; Lauer 
1959 (Honduras) ; Loock 1950 (Mich.); Miranda 1961 (Chis.); 
Schwerdtfeger 1953 (Guatemala); Shaw 1909; Standley and 
Steyermark 1958 (Guatemala); Vogel 1954 (Honduras); Wag- 
ner 1962 (Chis.). 

Thesis—Carl L. Johannessen 1959. The geography of 
the savannas of interior Honduras, Ph.D. thesis, Univ. Calif., 
Berkeley, Calif., 283 pp., illus. 

Unpublished—C, E. Blanco 1964 (Dgo.); B. Hallberg 
1964 (Oax., Chis.) ; E. Larsen 1962; N. Sanchez Mejorada to 
L. W. Bryan 1963 (Mich.); M. D. Moore 1963 (EI Salvador) ; 
O. Zarzosa L. 1964 (Dgo.). 


Pinus pringlei Shaw MAP 61 
Pringle pine 
Pinus pringle: Shaw in Sarg., Trees and Shrubs 
1: 211, t. 100. 1905. 

This distinctive low-elevation pine is confined to southern 
Mexico, ranging from Michoacan to Puebla and Oaxaca. It 
is sometimes associated with the related Pinus oocarpa. 
Additional sources: 

Published—Shaw 1909; Loock 1950 (Mich.). 

Unpublished—Hallberg 1964 (Oax.); Larsen 1962 
(Mex.). 


Anonymous. 

1963. [Pinus monophylla in Baja California.] 
Nat. Hist. Mus. Bul. 354, Apr. 1, 1963. San 
Diego, Calif. 1 p. 

Adamovic, L. 

1909. Die Vegetationsverhaltnisse der Balkan- 
lander. 567 pp., illus. (Die Vegetation der 
Erde, Vol. XI, ed. A. Engler and O. Drude) 
Leipzig: Wilhelm Engelmann. 

Aguilar G., J. Ignacio. 

1961. Pinos de Guatemala. 33 pp., illus. 

Guatemala. Min. Agr., Dir. Gen. Forest. 
Aikman, J. M., and Hayden, Ada. 

1938. Iowa trees in winter. Iowa State Col. 

Ext. Cir. 246, 72 pp., illus. 
Aitchison, J. E. T. 

1879. The vegetation of the Kuram and Har- 

iab Valleys. Indian Forester 5: 179-188. 


1881-1882. On the flora of the Kuram Valley, 
etc., Afghanistan. Linn. Soc. Jour. Bot. 18: 
1-113; 19: 139-200, illus. 


1891. Notes to assist in a further knowledge 
of the products of western Afghanistan and 
of north-eastern Persia. Bot. Soc. Edinburgh 
Trans. 18: 1-266. 

Akademia Nauk, SSSR. 


1954. [Vegetation map of U.S S.R.]_ 1: 
4000,000, 8 sheets. Bot. Inst. Imeni L. Kom- 
arova. [In Russian.] 


Alemdag, Seref. 

1962. [Development, yield and management 
rules of red pine (Pinus brutia Ten.) for- 
ests in Turkey.] Ormancilik Arastirma En- 
stitusi Yayinlari, Teknik Biilten Serisi 11, 
160 pp., ilfus. [In Turkish. English sum- 
mary.] 

Allen, P. H. 

1955. The conquest of Cerro Santa Barbara, 

Honduras. Ceiba 4: 253-270, illus. 
Andresen, John W., and Beaman, John H. 

1961. A new species of Pinus from Mexico. 

Arnold Arboretum Jour. 42: 437-441, illus. 
Anic, Milan. 

1957. [Austrian pine (Pinus nigra Arn.) on 
the northern part of Velebit (Croatia) .] 
Glasnik za Sumske Pokuse 13: 461-507, illus. 
[In Serbo-Croatian. English summary.] 

Aughanbaugh, J. E. 

1950. Shortleaf pine in Pennsylvania. Pa. 

Forests and Waters 2: 82-85, 93. 
Austin, Lloyd. 

1928. Breeding pines for more rapid growth. 

Jour. Hered. 19: 288-301, illus. 
Ayres, H. B. 

1900a. The Flathead forest reserve. U.S. 
Geol. Survey 20th Ann, Rpt., Part V, pp. 
245-316, illus. 


1900b. Lewis and Clark forest reserve, Mon- 
tana. U.S. Geol. Survey 2lst Ann. Rpt., Part 
V, pp. 27-80, illus. 
Bacigalupi, Rimo. 
1933. Pinus flexilis in the upper Kern River 
watershed. Madrono 2: 107-108. 
Bailey, Harold E., and Bailey, Virginia Long. 
1941. Forests and trees of the western na- 
tional parks. U.S. Dept. Int. Conserv. Bul. 
6, 129 pp., illus. 
Bailey, Vernon. 
1913. Life zones and crop zones of New Mex- 
ico. U.S. Dept. Agr. Bur. Biol. Survey, No. 
Amer. Fauna 35, 100 pp., illus. 


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illus. 

West, Erdman, and Arnold, Lillian E. 

1956. The native trees of Florida. Rev. ed. 

218 pp., illus. Gainesville: Univ. Fla. Press. 


White, J. H., and Hosie, R. C. 


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Forests. 


Whitford, H. N. 

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———_—_——— and Craig, Roland D. 


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Wicht, C. L. 
1949. Tree seed hunting in Mexico. So. 


African Forestry Assoc. 
illus. 
Wiggins, Ira L. 
1940. Yellow pines and other conifers ob- 
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Gard. Jour. 41: 267-269, illus. 


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Williams, L. H. J. 

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Williams, Llewlyn. 
1939. Arboles y arbustos del Istmo de Te- 
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1955. Pinus caribaea. 
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Wilson, Ernest Henry. 


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1907. A letter from China. Gard. Chron. 
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Wooton, E. O., and Standley, Paul C. 

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3: 97-118, illus. 
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1924. Natural vegetation. In Atlas of Amer- 
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illus. U.S. Dept. Agr., Bur. Agr. Econ. 


Ban Me 
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INDEX OF SCIENTIFIC AND COMMON NAMES 


Text page numbers and map numbers are given for the accepted 
specific names in Pinus and for the English common names under pine 
and pinyon. Cross references are added for scientific names mentioned 
or combined as synonyms. 


xine, Aleppo, 12, Map 31 ponderosa, 16, Map 47 durangensis, 17, Map 50 
| PP 1) I p g P 


Apache, 17, Map 49 Pringle, 20, Map 61 echinata, 15, Map 42 7 
Armand, 7, Maps 11 and 12 red, 11, Map 26 edulis, 8, Map 17 | 
Austrian, 11, Map 27 sand, 19, Map 57 (Map 56) eldarica, see P. brutia | 
Balkan, 7, Map 10 Scotch, 13, Map 32 elliottii, 16, Map 45 

bishop, 20, Map 59 shortleaf, 15, Map 42 engelmannii, 17, Map 49 

blue, 7, Map 12 (Map 14) Siberian stone, 5, Map 4 excelsa, see P. griffithit 

bristlecone, 10, Map 21 (Map 20) slash, 16, Map 45 fenzeliana, 8, Map 14 

Canary Island, 10, Map 24b southwestern white, 7, Map 8 (Map 9) flexilis, 6, Map 8 

Caribbean, 16, Map 46 spruce, 15, Map 43 flexilis var. macrocarpa, see P. strobiformis 

Chihuahua, 10, Map 22 sugar, 6, Map 7 flexilis var. reflexa, see P. strobiformis 


chilgoza, 9, Map 19a 
Chinese, 14, Maps 37 and 38 
chir, 11, Map 24a 

Cooper, 17, Map 51 
Coulter, 18, Map 55 

Cuban, 16, Map 46 

Digger, 18, Map 54 
Douglas, 18, Map 52 
Durango, 17, Map 50 
eastern white, 6, Map 6 
foxtail, 10, Map 20 (Map 21) 
Gregg, 20, Map 60 
Hartweg, 17, Map 51 
Heldreich, 12, Map 28 
Hwangshan, 14, Map 36 
Italian stone, 11, Map 25 
jack, 19, Map 56 

Japanese black, 13, Map 34 
Japanese red, 13, Map 33 


Japanese stone, 5, Map 3 (Map 4) 


Japanese white, 8, Map 13 
Jeffrey, 17, Map 48 
Khasi, 14, Map 38 (Map 37) 
knobcone, 19, Map 58 

- Korean, 5, Map 2b (Map 11) 
lacebark, 10, Map 19b 
Lawson, 18, Map 53 
limber, 6, Map 8 
loblolly, 15, Map 41 
lodgepole, 19, Map 56 
longleaf, 15, Map 40 
Luchu, 14, Map 36 
Lumholtz, 10, Map 23 
maritime, 12, Map 30 
Masson, 13, Map 35 
Merkus, 15, Map 39 
Mexican weeping, 20, Map 60 
Mexican white, 7, Map 9 
Michoacan, 18, Map 50 
Monterey, 19, Map 58 
Montezuma, 17, Map 49 
pitch. 15, Map 44 
pond, 15, Map 44 


96 


Swiss mountain, 12, Map 29 
Swiss stone, 5, Map 4 
Table-Mountain, 16, Map 43 
Taiwan red, 14, Map 36 
‘Taiwan white, 8, Map 14 
Torrey, 18, Map 55 

tropical, 11, Map 26 
Virginia, 19, Map 57 (Map 56) 
Washoe, 17, Map 48 

West Indian, 16, Map 46 
western white, 6, Map 6 
whitebark, 5, Map 5 
Yunnan, 14, Map 37 


Pinus, 5, Map 1 
Pinus albicaulis, 5, Map 5 


anhweiensis, see P. armandii 
apacheca, see P. engelmannii 
aristata, 10, Map 21 (Map 20) 
arizonica, see P. ponderosa 
armandii, 7, Maps 11 and 12 
attenuata, 19, Map 58 
ayacahuite, 7, Map 9 


ayacahuite var. brachyptera, see P. 


formis 
balfouriana, 10, Map 20 (Map 21) 
banksiana, 19, Map 56 
brutia, 12, Map 31 
-bungeana, 10, Map 19b 
canariensis, 10, Map 24b 
caribaea, 16, Map 46 
cembra, 5, Map 4 
cembroides, 8, Map 15 (Map 17) 
chihuahuana, see P. leiophylla 
chylla, see P. griffithii 
clausa, 19, Map 57 (Map 56) 
contorta, 19, Map 56 
cooperi, 17, Map 51 
coulteri, 18, Map 55 
cubensis, 16, Map 46 
culminicola, 9, Map 18 
dalatensis, 8, Map 14 
densiflora, 13, Map 33 
douglasiana, 18, Map 52 


funebris, see P. densiflora 
gerardiana, 9, Map 19a 

glabra, 15, Map 43 

greggtt, 20, Map 60 

griffithii, 7, Map 12 (Map 14) 
halepensis, 12, Map 31 
hartwegii, 17, Map 51 
heldreichii, 12, Map 28 

henryi, see P. massoniana 
herrerai, see P. teocote 
himekomatsu, see P. parviflora 
hwangshanensis, 14, Map 36 
insularis, 14, Map 38 (Map 37) 
jeffreyi, 17, Map 48 

koraiensis, 5, Map 2b (Map 11) 
krempfii, 5, Map 2a 
kwangtungensis, see P. fenzeliana 
lambertiana, 6, Map 7 
latifolia, see P. engelmannit 
lawsonii, 18, Map 53 
leiophylla, 10, Map 22 
longifolia, see P. roxburghii 
luchuensis, 14, Map 36 
lumholtzii, 10, Map 23 

lutea, see P. cooperi 
macrophylla, see P. engelmannit 
massoniana, 13, Map 35 
maximartinezii, 9, Map 15 
merkusii, 15, Map 39 
michoacana, 18, Map 50 
monophylla, 9, Maps 16 and 17 
montana, see P. mugo 
montezumae, 17, Map 49 
monticola, 6, Map 6 
morrisonicola, 8, Map 14 
mugo, 12, Map 29 

muricata, 20, Map 59 

nelsonti, 9, Map 18 

nigra, 11, Map 27 

oaxacana, see P. pseudostrobus 
occidentalis, 16, Map 46 
oocarpa, 20, Map 61 

palustris, 15, Map 40 


parviflora, 8, Map 13 
patula, 20, Map 60 


pentaphylla,see P. parviflora 


peuce, 7, Map 10 
pinaster, 12, Map 30 
pinceana, 9, Map 18 
pinea, 11, Map 25 
pithyusa, see P. brutia 
ponderosa, 16, Map 47 
pringlei, 20, Map 61 
pseudostrobus, 18, Map 52 
pumila, 5, Map 3 (Map 4) 
pungens, 16, Map 43 
quadrifolia, 9, Map 16 
radiata, 19, Map 58 
reflexa, see P. strobiformis 
remorata, see P. muricata 


resinosa, 11, Map 26 

rigida, 15, Map 44 

roxburghii, 11, Map 24a 

rudis, see P. hartwegit 
sabiniana, 18, Map 54 

serotina, 15, Map 44 

sibirica, 5, Map 4 

strobiformis, 7, Map 8 (Map 9) 
strobus, 6, Map 6 

sylvestris, 13, Map 32 
tabulaeformis, 14, Maps 37 and 38 
taeda, 15, Map 41 

taiwanensis, 14, Map 36 
tenuifolia, see P. pseudostrobus 
teocote, 18, Map 53 
thunbergiana, 13, Map 34 
thunbergii, see P. thunbergiana 


yy U. S. GOVERNMENT PRINTING OFFICE: 1965—O 764-741 


torreyana, 18, Map 55 
tropicalis, 11, Map 26 


uncinata, see P. mugo 


virginiana, 19, Map 57 (Map 56) 


wallichiana, see P. griffithii 
wangii, 8, Map 14 
washoensis, 17, Map 48 


yunnanensis, 14, Map 37 


pinyon, 8, Map 17 
pinyon, Martinez, 9, Map 15 
Mexican, 8, Map 15 (Map 17) 


Nelson, 9, Map 18 
Parry, 9, Map 16 

Pince, 9, Map 18 
Postosi, 9, Map 18 


singleleaf, 9, Maps 16 and 17 


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