Citation
The relationships of the salamanders of the genus Plethodon

Material Information

Title:
The relationships of the salamanders of the genus Plethodon
Creator:
Highton, Richard Taylor, 1927- ( Dissertant )
Grobman, Arnold B. ( Thesis advisor )
Wallace, H. K. ( Reviewer )
Bader, Robert S. ( Reviewer )
Edwards, R. ( Reviewer )
Carr, Archie ( Reviewer )
Place of Publication:
Gainesville, Fla.
Publisher:
University of Florida
Publication Date:
Copyright Date:
1956
Language:
English
Physical Description:
214 leaves. ; 28 cm.

Subjects

Subjects / Keywords:
Abdomen ( jstor )
Amphibians ( jstor )
Coastal plains ( jstor )
Counties ( jstor )
Pigmentation ( jstor )
Pigments ( jstor )
Salamanders ( jstor )
Species ( jstor )
Stripes ( jstor )
Vertebrae ( jstor )
Biology thesis Ph. D
Dissertations, Academic -- Biology -- UF
Salamanders ( lcsh )
City of Jasper ( local )
Genre:
bibliography ( marcgt )
non-fiction ( marcgt )

Notes

Abstract:
Abbreviated introduction: The North American genus Flethodon presently has more recognised forms than any other genus of salamanders. It is the type genus of the lungless family Plethodontidae, reviewed so ably by Dunn in 1926. At present, this is the most successful salamander family; over half of the living species of the Order Caudata are plethodontids. North America is the center of the distribution of the family, but one genus, Hydromantes , also occurs in Europe, and members of several genera enter the Neotropical region. Salamanders of this family occupy habitats ranging from strictly aquatic cavernicoles (lyphlomolge and haideotriton ) and mountain stream dwellers ( Leurognathus ) to others like Plethodon , which are completely terrestrial, even to the ex-bent of laying their eggs on land. Dunn believed that Plethodon is the most primitive genus in the attached-tongue branch of the family, and that Hemidactyiium, Ensatina, Batrachoseps , and Aneides are more specialised derivtives of a Plethodonlike ancestor. In 1926, only eleven forms of Plethodon were known to Dunn. In 1943 Bishop listed 17 species (one has since been removed from the genus) and two subspecies (both are now considered different species). In 1944, Grobman reviewed the distribution and relationships of the eastern section of the genus, including some sixteen species and subspecies. There are now twenty-five forms recognized in eastern North America, with seven more in the western United States and Canada. Grobman (1944: 266) divided the eastern forms into two groups, the Large Plethodons and the Small Plethodons. He suggested that these groups might actually represent distinct genera or subgenera, but he reserved judgement in this matter until the relationship of the western species with the eastern forms could be determined. One of the purposes of the present study is to determine these relationships by a comparative study of the morphology of all the species of the genus Plethodon. Of the twenty-five genera in the family Plethodontidae, only two others show as great or greater disjunctions in their distributions as that found in Plethodon , These are hydromantes , with two forms in Europe and three species in California, and Aneides , with four species in western North America and one in the Appalachian Mountains of the eastern United States. In the past two decades, several new plethodons have been discovered in both eastern and western North America. A review of the relationships of these forms offers a valuable opportunity to study the evolution of this important group of North American salamanders. The importance of examining plethodons in life has been emphasized by most recent workers. Many species, particularly the Large Eastern forms, are extremely difficult to identify after years of preservation. In few other vertebrates are differential characters between species so rare. For this reason, a special attempt was made to obtain living specimens of each form. Twenty-five of the thirty-two forms recognized herein have been studied alive.
Thesis:
Thesis -- University of Florida, 1956.
Bibliography:
Bibliography: leaves 198-213.
General Note:
Manuscript copy.
General Note:
Vita.

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University of Florida
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University of Florida
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Copyright [name of dissertation author]. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
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021499998 ( AlephBibNum )
13110546 ( OCLC )
ACW6856 ( NOTIS )

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THE RELATIONSHIPS OF THE SALAMANDERS

OF THE GENUS PLETHODON










By
RICHARD HIGHTON


A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF
THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY











UNIVERSITY OF FLORIDA
June, 1956











TABLE OF CONTENTS


INTRODUCTION . . . . . . . . . .

MiTHODS ..... . ........... .

THE RELATION PETWEEI! NUITIER OF COSTAL GROOVES AND TRUNK
VERTEBPAE I LEODON . . . . . . .

PIGLMITA TION IN THE GENUS PLETHODON . . . ..

KEY TO TIH StAtLUt1DiRS OF THE GEMUS PLTHODON . ..

SYSTE.MTICS . . ... . . . . . . ...

The Western Plethodons ...... . . .


Plethodon vandykei Group . . . .

Plethodon vandykei vandykei Van Denburgh

Plethodon vandykei idahoensis Slater and S

Plathodon vandykei larselli Burns . .

Plethodon vehiculun Group . . . . .

Plethodon dunni Bishop .... . .

Plethodon vehiculum (Cooper) . . .

Plethodon products Group . . . ..

Plethodon products, new name . .

Plethodon neomexicanus Group . . ..

Plethodon neoRexicanus Stebbins and Riemer

The Eastern Small Plethodons . .. .

Plethodon welleri Group ...........

Flethodon welleri Walker . . . .


* pp

Lipp


9





9

9


*

*


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*





.*

*

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*

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9


Page

. 1

.9 5


8

12

. l

.. 20

. 29

3L
. 34

35

. 37

37

* 38



. 3
. 41



. 83
43






46
S. 7


51

51


Plethodcn richmrondi richmondi Netting and Miittlean

Plethodon richriondi papei pighton and Grobrian .

Plethodon richrondi netting Green . . .


,-~"


*


*


*









Plethodon cinereus Group .............. 65

Plethodon dorsalis dorsalis Cope . . .. 68

Plethodon dorsalis angusticlavius Grobman . .. 71

Plethodon cinereus cinereus (Green) . . . 76

Plethodon cinereus serratus Grobman . . .. 78

Plethodon cinereus polycentratus Highton and Grobman 79

The Eastern Large Plethodons . . . ..... . 80

Plethodon wehrlei Group ............... 90

Plethodon wehrlei wehrlei Fowler and Dunn . . 93

Plethoden wehrlei dixi Pope and Fowler . .. 96

Plethodon wehrlei jacksoni NeQ . . . . 96

Plethodon yonahlossee Group ........ ... 97

Plethodon yonahlossee Dnn ........... 100

Plethodon ouaehitae Dunn and Heinse . . . 103

Plethodon caddoensis Pope and Pope . . . 105

P1ethodon glutinosus Group . .......... 106

Plethodon jordani jordani Blatchley . . . 129

Plethodon jordani metcalfi Brimley . . . 132

Plethodon jordani shermani Stejneger . . . 134

Plethodon jordani unicoi, new subspecies . . 135

Plethodon jordani melaventris Pope and Hairston 136

Plethodon jordani rabunensis Pope and Hairston 138

Plethodon jordani teyahalee Hairston . .. 1 0

Plethodon jordani clemsonae Brimley . . ... 141

Plethodon glutinosus glutinosus (Green) . . 193

Plethodon glutinosus albagula Grobman . . . 196

SUKMRY .................. ....... .. 197

LITERATURE CITED ................ ..... 198






iii


LIST OF FIGURES

Figure Page

1. Lateral and dorsal views of the trunk vertebrae of
representatives of the three major groups of the
genus Plethodon. . . . . . . . . . 23

2. Suggested phylogeny of the major subdivisions of the
genus Plethodon. . . . . . . . . . 26

3. Suggested phylogeny of the Western Plethodons . . . 33

4. The distribution of the subspecies of Plethodon
vandykei in Washington, Oregon, and Idaho. . . 36

5. The distribution of Plethodon dunni in Oregon and
southwestern Washington, . .. ... .. . 0

6. The distribution of Plethodon vehiculum in Oregon,
Washington, and BritishColumbia . . . . . 42

7. The distribution of Plethodon products in California
and Oregon . . . . . . . . . . . 45

8. The distribution of Plethodon neomexicanus in New
Mexico . . . . . . . . . ... 45

9. Suggested phylogeny of the Eastern Small Plethodcns . 52

10. The distribution of Plethodon welleri in North
Carolina, Tennessee, and Virginia. . . . . .. 52

11. The distribution of the subspecies of Plethodon
richmondi. . . . . . . . . . . 63

12. The distribution of the subspecies of Plethodon
dorsalis . . . . . . . 70

13. The distribution of the subspecies of Plethodon
cinereus . . . . . . . ..... .. 77

14. Vomerine teeth of 269 peninsula Florida P. glutinosus
plotted against snout-vent length. .. . . . 83

15. Vomerine teeth of 45 P. glutinosus from the Coastal
Plain of Virginia and North Carolina plotted
against snout-vent length. ... . ........ 83

16. Vomerine teeth of 115 P. j. jordani plotted against
snout-vent length .. . . . . . .. 8










Figure Page

17. Vomerine teeth of 60 P. w. wehrlei plotted against
snout-vent length. .. .. . .......... 84

18. Vomerine teeth of 39 P. caddoensis plotted against
snout-vent length. . . . . . . . 8

19. Suggested phylogeny of the Eastern Large Plethodans . 8

20. The distribution of the subspecies of Plethodon wehrlei 95

21. The distribution of the Plethodon yonahlossee Group . 102

22. The distribution of the subspecies of Plethodon ordani 112

23. The distribution of the dark-chinned (glutinosus) and
light-chinned (albagula) populations of P. glutinosus
in central Texas . -. . . . . . . 189

2h. The distribution of Plethodon glutinosus in eastern
United States . . . . . . . . . 195









INTRODUCTION


The North American genus Plethodon presently has more recog-

nized forms than any other genus of salamanders. It is the type genus

of the lungless family Plethodontidae, reviewed so ably by Dunn in 1926.

At present, this is the most successful salamander family; over half of

the living species of the Order Caudata are plethodontids. North Amer-

ica is the center of the distribution of the family, but one genus,

Hydromantes, also occurs in Europe, and members of several genera enter

the Neotropical region. Salamanders of this family occupy habitats,

ranging from strictly aquatic cavernicoles (Typhlomolge and Haideotriton)

and mountain stream dwellers (Leurognathus) to others like Plethodon,

which are completely terrestrial, even to the extent of laying their eggs

on land. Dunn believed that Plethodon is the most primitive genus in the

attached-tongue branch of the family, and that Hemidactylium, Ensatina,

Batrachoseps, and Aneides are more specialized derivatives of a Plethodon-

like ancestor.


In 1926, only eleven forms of Plethodon were known to Dunn.

In 1943, Bishop listed 17 species (one has since been removed from the

genus) and two subspecies (both are now considered different species).

In 1944, Grobrnan reviewed the distribution and relationships of the

eastern section of the genus, including some sixteen species and sub-

species. There are now twenty-five forms recognized in eastern North

America, with seven more in the western United States and Canada.


Grobman (1914: 266) divided the eastern forms into two groups,

the Large Plethodons and the Small Plethodons. He suggested that these






-2-


groups might actually represent distinct genera or subgenera, but he

reserved judgement in this matter until the relationship of the west-

ern species with the eastern forms could be determined. One of the

purposes of the present study is to determine these relationships by

a comparative study of the morphology of all the species of the genus

Plethodon.


Of the twenty-five genera in the family Plethodontidae, only

two others show as great or greater disjunctions in their distribu-

tions as that found in Plethodon. These are Hydromantes, with two

forms in Europe and three species in California, and Aneides, with

four species in western North America and one in the Appalachian Moun-

tains of the eastern United States. In the past two decades, several

new plethodons have been discovered in both eastern and western Iorth

America. A review of the relationships of these forms offers a valu-

able opportunity to study the evolution of this important group of

North American salamanders.


The importance of examining plethodons in life has been em-

phasized by most recent workers. Many species, particularly the Large

Eastern forms, are extremely difficult to identify after years of pres-

ervation. In few other vertebrates are differential characters between

species so rare. For this reason, a special attempt was made to obtain

living specimens of each form. Twenty-five of the thirty-two forms

recognized herein have been studied alive.


The list of persons who have contributed specimens to this

project is long and I wish to express to each of them my sincerest

thanks and appreciation for their valuable aid: Charles G. Adams,






-3-


Walter Auffenberg, Herbert Barden, Albert P. Blair, E. E. Brown,

Jerry and Esther Brown, Douglas M. Burns, Archie Carr, A. H. Chaney,

Roger Conant, John W. Crenshaw, J. C. Dickinson, John Dolan, Herndon

Dowling, Philip C. Dumas, Helen T. Gaige, M. Ruth Gilreath, Arnold B.

Grobman, T. P. Haines, Keith L. Hansen, Robert Hellman, Albert H.

highton, Thelma Howell, Leslie Hubricht, Robert Humphries, Richard M.

Johnson, James Kezer, J. D. Kilby, James Knepton, H. D. Leake, Edwin

H. McConkey, John S. Mecham, Sherman A. Minton, Wilfred T. Neill,

Howard T. Odum, Larry H. Ogren, David Pettus, John Quinby, George

Rabb, Neil D. Richmond, William Riemer, Bernard Roseman, Doug Rossman,

Albert Schwastz, Robert S. Simmons, Ralph Sinclair, William Sloan,

Peter Smith, Bette Starner, Charles J. Stine, Walter Stone, Virginia

Switzer, Sam R. Telford, G. M. Thorp, Gordon Thurow, Don Tinkle,

Arnold and Rusty Van Pelt, Charles F. Walker, Tilliam Witt, and Harry

Yeatman.


Most of the material given me by these persons has been

deposited in the University of Florida collection. This collection

has formed the nucleus of the material on which this study is based.

Some additional specimens have been borrowed from other collections,

but I did not attempt to follow the course of the conventional monog-

rapher and examine every specimen available in museums, simply for the

sake of completeness. The material borrowed from other museums in-

cludes the following: Western Plethodons from the University of

California; Texas P. glutinosus from Bryce C. Brown, Ottys Sanders,

John S. Mecham, and the Strecker LAseurm; P. d. angusticlavius, P.

glutinosus, P. ouachitae, and P. caddoensis from the University of

Arkansas; P. c. serratus, P. w. wehrlei, and P. w. dixi from the









Chicago Natural History Museum; P. glutinosus, P. w. wehrlei, and P.

richmondi from the Carnegie useum; P. cinereus from the Museum of

Comparative Zoology; P. richmcndi from the United States national

Museum; P. glutinosus and P. 1. rabunensis from the Charleston Euseum;

P. dorsalis from the University of Georgia; P. richmondi and P. glutin-

osus from the Cincinnati Society of Natural History; P. dorsalis from

the Ross Allen-'ilfred T. Neill collection; and the entire Flethodon

collection of the Great Smoky Mountains National Park.


For the loan of material in their care, I wish to thank the

following museum officials: Neil D. Richmond, Carnegie museum (CM);

Albert Schwartz, Charleston Museum (ChM); Robert Inger, Chicago Natural

History Vhseum (CNHM); Ralph Dury, Cincinnati Society of Natural History

(CSNH); Arthur Stupka, Great Smoky Mountains National Park (GSUNP);

Arthur Loveridge, Museum of Comparative Zoology (MCZ); woilfred T. Heill,

Ross Allen Reptile Institute (ERA-TTN); Bryce C. Brown, Strecker Museum

(SM); Ierndon Dowling, University of Arkansas (UA); Bernard Martof,

University of Georgia (UG); William Riemer, University of Florida (UF);

Charles F. Walker, University of Michigan, Museun of Zoology (U1:Z);

and Doris 1. Cochran, United States National Museum (USNM). The fol-

lowing individuals have generously let me borrow material in their

private collections: Bryce C. Brown (BCB); John S. Mecham (JSM); and

Ottys Sanders (OS).


In addition I have greatly benefitted from stimulating dis-

cussion of some of the problems which have arisen during the course of

this study with the following persons: Drs. Robert Bader, Pierce

Brodkorb, Archie Carr, John W. Crenshaw, J. C. Dickinson, Richard A.


- 4-










Edwards, Coleman J. Goin, howard T. Odum, UVilliam RLeraer, H. K.

Wallace, and Messrs. Walter Auffenberg, Robert Hellman, Richard M.

Johnson, and Tilfred T. Neill. 1y wife, Anne, besides being patient

and sympathetic while undergoing considerable personal inconvenience

during many long hours of field work in the various hot, wet, cold,

and mosquito-infested places where these animals live, has also mate-

rially contributed to the project by typing, filing, proofreading, and

especially by taking practically all the color notes on geographic

variation of the P. glutinosus group. The Chairman of nm Graduate

Supervisory Conmittee, Dr. Arnold B. Grobmrn, has been a constant

source of intellectual stimulation, encouragement and helpful advice

and criticism.



METHODS


The importance of studying living plethodons, because of the

usual loss and alteration of their pigments in preservatives, has al-

ready been mentioned. It is almost impossible to study geographic var-

iation in pigmentation without considerable field work, and it was ini-

tially hoped that this could be done with all the forms in the genus.

Unfortunately, several of the forms have not been seen alive and so

our knowledge of their geographic variation has not been augmented.

However, I have been able to do a considerable amount of field work

with some of the Eastern Large Plethodons, and the result has been a

considerable amount of new information concerning them. I believe

that when similar studies of other species are made, they will be

found to vary as much as glutinosus and jordani in pigmentation char-

acters.






-6-


A perusal of the literature on this genus will disclose

that most workers have given considerable weight to variation in the

number of costal grooves., A careful study of the literature indi-

cates, however, that there is not agreement in the reports of number

of costal grooves in many species. The reason for this was apparent

to the writer when I failed to obtain consistent counts on the same

specimens counted at different times. A method for accurately deter-

mining the true variation in body segmentation would be a necessary

preliminary to the use of this character in studying relationships.

Since the costal grooves show a direct correlation with the number of

vertebrae, a method of counting the number of trunk vertebrae by the

use of X-ray photographs was devised. As an outgrowth of this study,

a method for counting costal grooves which will accurately reflect

the body segmentation of salamanders of this genus will be reported

below.


'any other characters have been used in the study of the

systematics of this genus. As far as possible, these have been re-

studied using much new material that has recently become available.

Several nomenclatorial changes will be suggested based on an attempt

to best indicate the biological relationships of these organisms.


The results of this study have clearly indicated that there

are three major natural subdivisions of the genus Plethodon. These

would probably be considered subgenera in most animal groups, but

herpetologists have rarely used subgenera in their classification.

Although the use of subgenera would be helpful in this case, conmon

names will be used for these groups in conformity with current herpe-

tological practices. They will be referred to as the eastern n Pleth-






-7-


odons," "Eastern Small Plethodons," and "Eastern Large Plethodons."

Each of these subdivisions of the genus is further divided into

species groups consisting of species which are more closely related to

each other than they are to any other similar group. In a few cases,

species which are morphologically quite distinct from any other species

in their section of the genus have been segregated into a species

group consisting of a single species.


For each taxon, information on the morphology, variation,

and distribution is given, followed by a synonomy, data on the type

specimen, diagnosis of the form, and a description of the pigmentation,

segmentation, number of vomerine teeth, and size. Detailed descrip-

tions of several hundred specimens were made during the course of this

study. For most forms, similar data are available in Bishop (1943) or

Stebbins (1951) or in the original description. It is unnessary to

repeat these here, since the concern is variation of populations, not

detailed descriptions of individual specimens.

Dunn (1926) presents fairly complete synonomies for each of

the forms known at that time. Therefore the large number of refer-

ences on the distribution and habits of the more common forms are not

included here. An attempt has been made to include the first refer-

once to each combination of names, as well as most of the papers deal-

ing'.with the systematics of each form. Check lists have usually been

omitted, except when now combinations of names or other new information

is presented. Consistancy has not been a goal, and some papers deal-

ing with habits or distribution have been included in the synonomies

of little known species, while similar papers on well known forms






- 8 -


that do not contribute new information have been omitted.


The locality maps have been prepared mainly as an aid to

the discussion of the distribution of the various species groups.

They are based on literature records as well as specimens examined

by the writer. Maps showing the distribution of many of these forms

are available elsewhere, but there have been many changes in the

taxonomy since the publication of some of these, and it seems desir-

able to include them here in spite of the fact that there is some

repetition. New information on the geographic distribution of sever-

al of the forms is included on some of the maps.


Seventeen subspecies are recognized in this paper. An

examination of the distribution maps indicates that several of the

polytypic species exhibit terminal raciation and most of these

races are isolated from their nearest relatives by areas of unin-

habited country. This is true of the races of P. vandykei, P.

dorsalis, P. cinereus, and to some extent, P. glutinosus. The sub-

species of P. wehrlei are poorly defined, and need more study. There

do not appear to be natural barriers between some of the subspecies of

P. richmondi and P. jordani, and there is evidence that intermediate

populations are present between some of the races. A more detailed

consideration of raciation in both of these species is given below.


THE RELATION BETWEEN NUMBER OF COSTAL GROOVES AND

TRUNK VERTEBRAE IN PLETHODON


Radiographs of series of specimens of all the species in

this genus have indicated that the number of trunk vertebrae in a






-9-


single population is remarkably constant. (Since there is little

differentiation of the salamander body vertebrae into regions, Francis

(1934) suggests that all precaudal vertebrae, except the atlas and the

sacrum, be called trunk vertebrae.) There is never a variation of

more than three trunk vertebrae in a single population and with few

exceptions, the intermediate figure has a very high frequency of

occurrence. The value of this character in studying variation becomes

apparent when it is found that a difference of a single vertebra be-

tween two populations is easily detectable and has been used success-

fully as a key character to separate a high proportion of specimens of

two contrasted forms.


A method of counting costal grooves that would accurately

reflect the number of trunk vertebrae would be much more effective in

the taxonoTic study of this genus. At present, their use is rather

limited, since they are rarely counted consistently by different

workers. Pope (1950: 102) correctly summarizes the existing situa-

tion by stating that "variation of a magnitude of one or two grooves

will be meaningless unless the same worker has made all the counts

and done so .ith great care." The method described here has been

found to correctly correspond with the number of trunk vertebrae with

over 90% accuracy.

The atlas is not related to any costal grooves. The first

trunk vertebra is located just anterior to the gular fold and its rib

does not appear to be associated with the gular fold or with any of

the costal grooves. The rib of the second trunk vertebra is located

in the ryoseptum of the first costal groove. The external position

of this groove varies somewhat, but is usually close to the front






- 10 -


limb insertion. If the first costal groove is poorly defined or absent,

as often occurs when it is located directly over the front limb inser-

tion, it should be counted, as the space obviously corresponds to a ver-

tebra whether the groove is visible or not. The individual variation in

the position of the first costal groove is probably due to the variation

in the position of the pectoral girdle relative to the vertebrae. The

girdle may be located opposite the second trunk vertebra, between the

second and third trunk vertebrae, or opposite the third trunk vertebra.

The ribs of the third trunk vertebra extend posteriorly so that the

second coastal groove is always posterior to the front limbs.


There is a one-to-one relation between the remaining body ver-

tebrae and the costal grooves, except in the case of rare aberrations

(one vertebra possessing two or more pairs of ribs, or one vertebra

possessing a rib on only one side, in which case a corresponding costal

groove is present only on that side). The last pre-sacral vertebra

(usually the only trunk vertebra which does not possess ribs) is repre-

sented by the groove just anterior to the insertion of the hind limbs.

Often this groove joins ventrally the one just preceding it so that on

the lower sides there is only onegroove which is forked dorsally. Both

grooves should be counted, since they correspond to two separate verte-

brae. The groove present over the hind limb insertion (sometimes weakly

developed or absent) corresponds to the sacral vertebra, and each groove

on the tail corresponds to a caudal vertebra.


There is much more individual variation in the position of the

sacral vertebra in relation to the pelvic girdle than there is in the

relation of the shoulder girdle to the second and third trunk vertebrae.






- 11 -


The sacral vertebra may be located so far anterior to the pelvic girdle

that the first caudal vertebra is opposite the hind limbs. It may also

be slightly anterior, opposite, or slightly posterior to the pelvic

girdle. Occasionally when it is posterior to the girdle, the costal

groove corresponding to the last trunk vertebra is located over the

hind limb. This is the major source of error in attempting to accurately

correlate the number of costal grooves with the number of trunk verte-

brae. Fortunately, however, fewer than 10% of the specimens examined

were in this category. Using this method, costal groove counts were

made on 85 specimens of several species of the genus Plethodon (includ-

ing specimens possessing 15 to 22 trunk vertebrae), and 93% of the

counts accurately represented the number of trunk vertebrae obtained

from X-ray photographs. Since the first trunk vertebra is not asso-

ciated with a costal groove, the number of costal grooves is always one

less than the number of trunk vertebrae. It would therefore appear

that this method can be used advantageously to obtain an accurate es-

timate of the number of trunk vertebrae of all the species of the genus

Plethodon.


Occasionally the sacral rib may attach to one vertebra on one

side of the animal and to the following vertebra on the other side.

Rarely, there are two sacral ribs issuing from two successive vertebrae

on the same side of the animal. The suggested method of counting costal

grooves is of no value in detecting such abnormalities.





- 12 -


PIGMENTATION IN THE GENUS PLETHODON


Pigmentation in living specimens was studied with the aid of

a dissecting microscope. Although there is great variety in the color-

ation of the animals included in this genus, the actual different types

of pigments are few. No histological or biochemical studies of the

pigments have been made, thus similarity in the appearance of the pig-

mentation is the basis for the above statement. Three main types were

identified and are called melanophores, guanophores, and lipophores,

following Ctebbins (1951).


Melanophores are present in all the forms in the genus. They

are responsible for the dark ground color of these salamanders. The

other pigments generally occur in gaps in the melanophore ground color.


Guanophores are responsible for the white and brassy spots,

present on many of the species. There appears to be little difference

in the structure of the different colored guanophores, but mainly a

difference in the amount and color of light reflected from the spots.

The iridescence characteristic of animals with "brassy" "dorsal spots

appears to be centered in small round crystalline structures found at

intervals along the pseudopodia of the guanophores. The number of these

crystals seems to determine the amount of the iridescence that has been

variously described by different workers as "brassy flecking," metallic

golden spotting," "golden blotches," "bronzy mottling," and "frosting."

This type of guanophore pigmentation is present in the iris of most of

the Eastern Small Plethodons and the Western Plethodons, and is present

on the dorsum of many of the species. It is responsible for the brassy

dorsal flecks of glutinosus, ouachitae, dixi, clemsonae, popei, netting,







- 13 -


cinereus, polycentratus, dorsalis, welleri, and vehiculum. In welleri,

these guanophores are concentrated to form dense clusters. In Lluti-

nosus, they are usually associated with other white guancphores. In

the other forms they are scattered over the back and are not clumped

into spots. Brassy flecks are rarely found other than in the iris or

on the dorsum of salamanders of this genus.


Guanophores that lack the brassy iridescence are co-mon on

the belly and sides of many spocics, and are also present on the dorsum,

occasionally occurring there with the brassy type. They have a much

greater tendency to be clumped together to form larger spots than do the

brassy flecks, but are occasionally found singly. In glutinosus, for

example, it is often possible to see, on the same animal, every type

of intermediate between the brassy type and those which lack the brassy

iridescence.


The white guanophores are characteristically found on the

dorsum of glutinosus, ouachitae, cinereus, polycentratus, popei,

dorsalis, and vehiculum. They are present on the sides of almost all

of the eastern species (except metcalfi and melaventris), and on the

bellies of all the Eastern Small Plethodons. Often these lateral and

ventral spots have a yellowish color, but this is not due to the presence

of brassy flecks.


Lipophores do not have the structure of guanophores in that

they lack the pseudopodia vhich can easily be observed in the guano-

phores. The red color of jordani, shermani, wehrlei, yonahlossee,

ouachitae, cinereus, polycentratus, serratus, dorsalis, and vehiculum





- 14 -


appears to be due to the same type of lipophore pigment. The dorsal

band of yonahlossee is a darker color because of the additional pres-

ence of melanophore pigment. Lipophores may also be yellow in color,

as in some cinereus, vehiculum, and dunni.


The variety of colors present in the genus seems to be due

entirely to variation in the abundance of these pigments or various

combinations of the three. In some forms, the lipophores or guanophores

or both are lacking. There is also variation in the concentration of

melanophores. These pigmentation characters may vary somewhat withinn a

species, both individually and geographically, but are fairly uniform in

most forms, enabling a person familiar with living specimens to easily

identify most salamanders by the color pattern alone. The phylogenetic

significance of the distribution of these pigments in the various pleth-

odons is discussed below in the accounts of the various forms.



KEY TO THE SALAMANDERS OF THE GENUS PLETHODON


The key is based mainly on the average number of trunk verte-

brae occurring in each of the forms. Before using this key, it is sug-

gested that the section on the method of counting costal grooves that

accurately reflects the number of trunk vertebrae be read. A small per-

centage of specimens of each form may not be correctly identified on

this basis, but a small series will usually key out correctly. Ranges

are included and locality data may proove more helpful in identifying

preserved specimens than pigmentation characters.


1 a Costal grooves usually 14 ................. 2

b Costal grooves 15 or more . . . . . . .... 4








2 a Ventral color reddish (Multnomah County, Oregon, and Ska-

mania County, Washington) . . ..... P. v. larselli

b Ventral color not reddish ... ... ... ..... 3

3 a Yellow or orange dorsal stripe contrasts sharply with the
lateral black ground color; the proximal segments of the

limbs dark in color (Kootenai County, Idaho) P. v. idahoenois

b Ground color light, not contrasting sharply with the dor-

sal stripe; yellow lipophore pigment similar to that found

on the dorsum present on the proximal segment of the

limbs (western Jashington) . .... P. v. vanykei

4 a Costal grooves usually 15 (western Oregon and southwestern
Washington) . . . . . . . . P. dunni

b Costal grooves 16 or more .... ... . . . 5

5 a Costal grooves usually 16 .. .... .. ... 6

b Costal grooves 17 or more . . . ........ 20

6 a Belly mottled with yellow or red, white and black; size small,

usually under 50 mm. snout-vent length; often a red, tan,

or yellow dorsal stripe (southwestern British Columbia, in-

cluding Vancouver Island, western '",ahirnton and western

Oregon) ................... P. vehiculum

b Belly usually dark, at least posteriorly, or with scattered

small white ventral spots; size usually larger (except

welleri and caddoensis); dorsal stripe usually absent

(except yonahlossee and ouachitae) (eastern United States) 7

7 a Red pigment present on back, legs, or cheeks . . ... 8

b No red pigment present on animal . .... ...... 11


- 15 -





- 16 -


8 a Red pigment confined to sides of head or legs in adults 9

b Red pigment largely restricted to dorsum . . .. 10

9 a Red pigment most abundant on sides of head (Great Smo1ky

Fountains of Tennessee and North Carolina) . jordani

b Red pigment most abundant on legs (Mantahala Mountains,

North Carolina) . . . . ..... P. sherani

10 a White pigment lacking in dorsal stripe (Blue Ridge Aoun-

tains of southwestern Virginia, northeastern Tennessee,

and northwestern North Carolina) . . . P. yonahlossee

b Abundant white pigment occurring within the dorsal stripe

(Ouachita mountains of Arkansas and Oklahoma) P. ouachitae

11 a Size small, adults not over 50 Im. snout-vent length;

back with large coalescing iridescent brassy spots,

usually covering about half the area of the dorsum

(Blue Ridge Province of southwestern Virginia, north-

eastern Tennessee and northwestern Ntorth Carolina) P. wolleri

b Back without dorsal brassy spots, or if present, they

are small in size and cover less than one quarter of

the area of the dorsur . .. ....... ... 12

12 a Body entirely black, guanophores and lipophores absent . 13

b Dorsum and/or sides with guanophore spotting . . .. 1

13 a Belly much lighter than dorsum (mountains of western North

Carolina and adjacent Tennesece and Virginia) P. j. metcalfi

b Belly almost as dark as dorsum (southwestern North

Carolina) . . . . . . .. .. P. j. melaventric





- 17 -


14 a Back with large conspicuous white or brassy dorsal spots 15

b Back without large conspicuous guanophore spotting, or if

present, the spots are of very small size . . . 17

15 a Melanophore pigmentation on chin greatly reduced, compared

to belly ....................... 16

b Melanophore pigmentation on chin similar to that on belly

(eastern United States from southern New York to northern

Florida, west to eastern Louisiana, Illinois, Lissouri,

eastern Oklahoma, and the Balcones Escarpment in Texas)

. . . . . . . . . . . lutincsus

16 a Dorsum with white spots only (Balcones Escarpment in

east central Texas) . . . . ... . P. albagula

b Dorsum with large white spots and smaller brassy flecks

(Caddo Mountains of Arkansas) .. . . . P. caddoensis

17 a Dorsum with very small white or brassy spots . . . 18

b Dorsum black, without any guanophore spotting . . . 19

18 a Dorsum covered with tiny brassy flecks (vicinity of

Jocassee, South Carolina) . . . . P. J. clemsonae

b Dorsum with very tiny scattered white spots (Tusquitee

and Snowbird Mountains of western North Carolina)

. . . . . . . . . . P. P. teyahalee

19 a Belly much lighter than ground color of back (Unicoi

Mountains of western North Carolina and eastern

Tennessee) . . . . . . . . P. j. unicoi

b Belly nearly as dark as dorsal ground color (mountains

of northeastern Georgia . . .... . P. rabunensis






- 18 -


20 a Costal grooves usually 17 . . .. .. . ... . 21

b Costal grooves 18 or more . . . . . . . .. 23

21 a Large red spots on dorsum of adult (southwestern Virginia)

* * *. * . . . . . . . P. w. jacksoni

b Red spots absent from dorsum of adult . . . . . 22

22 a Dorsum with numerous white and brassy spots (Roanoke

County, Virginia) . . . . . .... P. w. dixi

b Dorsum with very small white or brassy spots (Cattaraugus

County, New York, south through western Pennsylvania,

adjacent Ohio, and West Virginia) . . . P. w. wehrlei

23 a Costal grooves usually 18 . . . ....... .. 2L

b Costal grooves 19 or more . . . . . . . . 27

24 a Size large, up to 66 mm. snout-vent length (northwestern

California and southwestern Oregon) . . . P. products

b Size small, up to 45 mm. in snout-vent length (eastern

United States) . . . . . . . . . 2

25 a Belly mottled with red, yellow, white, and black pigment 26

b Belly black, with small white spots (Cheat Mountains of

West Virginia) . . . . . . . . P. r. nettingi

26 a Dorsal stripe very narrow, less than 1/3 the width of the

body (southwestern Missouri, northwestern Arkansas, and

adjacent Oklahoma) . . . . . d. angusticlavius

b Dorsal stripe much wider than 1/2 the width of the body,

edges of stripe very irregular (southern Illinois, Indiana,

and southern Ohio, south through Kentucky and Tennessee to

northern Alabama and northwestern Georgia) P. d. dorsalis






- 19 -


27 a Costal frooves usually 19 .............. . 28

b Costal grooves 20 or more . . . . . ..... 30

28 a Fifth too on hind foot usually with one phalanx (Jemez

fountains Now 'oxico) . . . . P. neomexicanus

b Fifth too on hind foot with two phalanges (eastern North

Amorica) . . . . . . . . . 29

29 a Dorsal red stripe with straight edges (southern Canada,

south to torth Carolina, eastern Kentucky, southern

Illinois, and eastern Missouri) . . . P. c. cinereus

b Dorsal red stripe with serrations corresponding to each

costal re (western Arkansas and eastern Okla-

homa) . . . .... . ... .. P. c. serratus

30 a 7;: pi rent present on sides and dorsum (northwestern

-r: ia) .............. P. c. polycentratus

b No red :i -cnt on animal ........... .. . 31

31 a Costal grooves usually 21 or more (western Pennsylvania,

south ni Ohio, eastern Kentucky, south to northwestern

Vir,i .) . . . . . . . . P. r. richmondi

b Costal r .ves usually 20 (southwestern Virginia, north-

western north Carolina, and adjacent Kentucky) P. r. popei





- 20 -


SYSTEMATICS


Plethodon Tschudi

Plethodon Tschudi (1838: 58). Genotype (by original designation):

Salamandra glutinosa Green.

Phatnomatorhina Bibron in Bonaparte (1839). (Substitute name)

Sauropsis Fitzinger (18h3: 33). (non Sauropsis Agassiz, 1832, Jahrb.

Min., 3: 1l2). Genotype (by original designation):

Salamandra erythronota Green.


Diagnosis;- Plethodontidae with tongue attached in front; premaxillae

separate; teeth on posterior portion of maxillae; tail not constricted

at base; five toes on the hind feet; no palmar tubercles; terminal

phalanges normal; and no aquatic larval stage.


According to Dunn (1926), the closest relatives of Plethodon

are Batrachoseps, Ensatina, Aneides, and Hemidactylium. All four of

these genera differ from Plethodon in several fundamental characteristics.

Batrachoseps and Hiemidactylium possess only four toes on the hind feet.

Ensatina and Ilemidactylium possess a basal constriction of the tail.

Ensatina has palmar tubercles. The premaxillae are fused in Aneides and

Batrachoseps (except B. wrighti. Hemidactylium has an aquatic larval

stage. Aneides lacks teeth on the posterior portion of the maxilla and

has expanded terminal phalanges. Dunn (1926: 22) believed that Plethodon

is the most primitive genus in this group of genera and that the others,

with the possible exception of Ensatina, are more specialized than

Plethodon. The genus Plethodon is the largest in number of species, and

there is more divergence within the genus than in the other related

genera. This might be considered evidence for, but not necessarily

proof of, a greater age for this genus.






- 21 -


Grobnn (IPo9: 266) suggests that the relationship between

the Eastern : -r Small Plethodons is not close and lists size

and costal groove c forenccs which distinguish them. Actually there

is some c.r-r. bet cen the Eastern Large and Small Plethodons in

these characters, -ut there are other important differences between the

two groups. nectal gland is much better developed in male Eastern

Large Plethodcn~ th n in the nastern Small Plethodons. The worm-like

body and shooter 13 of the sternn Small Plethodons is characteristic.

The Eastern -. lthodons have fewer teeth and there is a great deal

of variation in te number of trunk vertebrae within the group (range

16-2h), heres the ;astern Large Plethodons show very little variation

(range 16-19). Tiere is more webbing on the toes of the small forns

than in the 1- 7 cies. The Eastern Large Flethodons have an un-

pigmented --rietal eritoneun, while in the srall species the peri-

tonoum is :1 '"- rith melanophores. The differences in the struc-

ture of the ver ebrae are discussed below.


One .iore, P. wehrlei, is usually included with the Eastern

Large 'lethodonr on the basis of size, but in several ways it is inter-

mediate between the two groups. It has more costal grooves and fewer

teeth than other Saster Large Plethodons. Its toes are webbed as in

the -astern :.ll lethodons and the peritoneum has a few melanophores.

Kere it not for the intermediate character of this species, the two

groups should probably be recognized as distinct genera, but the some-

what inter. ei te nature of wehrlei would seem to indicate that they

have not "cc ne sufficiently distinct to justify such an action. The

two -rc. could best be regarded as subgenera, but to conform with

current prtise, the common names, Eastern Large Plethodons and Eastern






- 22 -


Small Plethodons, will be used in this paper. Although P. wehrlei

possesses more characters that would link it with the Eastern Small

Plethodons than do any of the other Eastern Large Plethodons, it is

probably more closely related to members of the latter group. There-

fore, it is included as a separate species group under the Eastern

Large Plethodons. Its intermediate position is important, however, in

linking the two groups.


During studies on the osteology of this genus, certain dif-

ferences in the vertebrae of the two eastern sections of the genus

have been noted. The height of the vertebrae of the Eastern Small

Plethodons is proportionately less and the vertebrae usually lack the

neural spines that are present on those of the large eastern species

(see figure 1). P. wehrlei is not intermediate in this regard, but

closely resembles the other Eastern Large Plethodons. The vertebrae

of all the eastern species have been examined except for caddoensis

and richmondi.


To investigate the relationship of the western forms with

those in eastern North America, each of the above differential charac-

ters was studied in all five of the western species. As with the east-

ern forms, the size varies within the group, but four of the five west-

ern species are as large as most of the Eastern Large Plethodons. The

number of trunk vertebrae varies considerably in the western forms

(range 15-20). The mental gland is poorly developed in all the Western

Plethodons. The body form is variable, ranging from short and stout in

vandykei to very elongate in neomexicanus. The number of vomerine teeth

is low in the western forms. The toes of two species (dunni and vehi-






- 23 -


B
I mm


c


Figure 1. Lateral and dorsal views of the trunk vertebrae of rep-
resentatives of the three major groups of the genus

Plethodon. A. P. glutinosus. L. P. welleri

C. P. dunni


1!


1-m






- 2h -


culum) are ver; slightly webbed, while the other chree species have

webbed toes. .' 2ritoneum of all of the westernr n Plethodons is

pigmented vith nel nophores. The vertebrae of the western forms are

quite different from both eastern types (P. neomexicanus has not been

examined). h- are proportionately longer and have longer transverse

processes than of the eastern species. As a group, the western

Plethodons are more variable, and in at least one character, the

structure of the vertebrae, they are very distinct from both eastern

sections of the gerus. It would appear that the v:estern and Eastern

Plethodons have been separated for a long time, and that both have

undergone considerable speciation during this period.


In several ways, the Eastern Small Plethodons resemble the

Western Flethodons more closely than do the Eastern Large Plethodons.

The red or yellow dorsal stripe is more common in both groups than it

is in the Eastern Large Plethodons. Both have a great amount of

vari;.tion in the number of trunk vertebrae and degree of elongation of

the body. Loth h've low vo-lerine teeth counts and a pigmented peri-

toneui. Both h-ve a less well developed mental gland than the Eastern

Large "lethodons (except for dorsalis). It would appear that the

Eastern tarre ilethodons are the more specialized fonas, with a larger

number of vonerine teeth, loss of pigmentation in the peritoneum, and

increase in the development of the mental gland. The fact that P.

wehrlei possesses characteristics of both groups would indicate that

both v.ere derived frovi a common ancestor. This conron ancestor was

probably nore like the Eastern Small Plethodons and the Western Pleth-

odons in ost of its characteristics, since it would be unlikely that

these two groups would have independently converged toward each other






- 25 -


in so many ways from an ancestor that was similar to the Eastern Large

Plethodons as suggested by Dunn. The relationships of the three groups

would appear to be best indicated by a phylogeny similar to that shown

in figure 2.


The relationship of the Western Plethodons with the other

plethodontid genera in western North America needs further study. It

may be that the Western Plethodons are more closely related to Batra-

choseps, Ensatina, or western Aneides than they are to any of the

Eastern Plethodons. The western plethodontid salamanders may have been

isolated from their eastern relatives for a long period of time. The

fact that two groups now included in the genus Plethodon, one in east-

ern North America and the other in the western part of the continent,

have both retained many primitive characteristics, does net necessarily

mean that they are still generically related to one another. The fact

that each of these groups (the Western and Eastern Plethodons) has more

species than any other plethodontid genus in its region, as well as the

the fact that these species are so diverse, would support the view that

they have been separated for a long period of time. A review of this

entire problem is needed, but it would involve an investigation of the

characteristics of the genera Aneides, Batrachoseps, Ensatina, and

Hlemidactylium, and is beyond the scope of this study. Such an inquiry

should be completed before erecting a new genus for the Western Pletho-

dons, but this arrangement seems to be indicated by the present incom-

plete information.


The systematic arrangement of the genus Plethodon suggested

in this paper is as follows:







- 26 -


Eastern Large
Plethodons


Figure 2.


Eastern Small
Plethodons


Suggested phylogeny of the major subdivisions of the


genus Plethodon.


Western
Plethodons





- 27 -


Western Plethodons

Plethodon vandykei Group

Plethodon vandykei vandykei Van Denburgh

Plethodon vandykei idahoensis Slater and Slipp

Plethodon vandykei larselli Burns

Plethodon vehiculum Group

Plethodon dunni Bishop

Plethodon vehiculum (Cooper)

Plethodon products Group

Plethodon products new name

Plethodon neomexicanus Group

Plethodon neomexicanus Stebbins and Riemer

Eastern Small Pletho'ons

Plethodon welleri Group

Plethodon welleri alker

Plethodon richmondi richmondi Netting and Miittle:nan

Plethodon richmondi popei llighton and Grobman

Plethodon richmondi nettingi Green

Plethodon cinereus Group

Plethodon dorsalis dorsalis Cope

Plethodon dorsalis angusticlavius Grobman

Plethodon cinereus cinereus (Green)

Plethodon cinereus serratus Grobman

Plethodon cinereus polycentratus Ilighton and Grobman






- 28 -


Eastern Large Plethodons

Plethodon wehrlei Group

Plethodon wehrlei wehrlei Fowler and Dunn

Plethodon wehrlei dixi Pope and Fowler

Plethodon wehrlei jacksoni Newman

Plethodon yonahlossee Group

Plethodon yonahlossee Dunn

Plethodon ouachitae Dunn and IIeinze

Plethodon caddoensis Pope and Pope

Plethodon glutinosus Group

Plethodon jordani jordani Blatchley

Plethodon jordani metcalfi Brimley

Plethodon jordani shermani Stejneger

Plethodon jordani unicoi new subspecies

Plethodon jordani nelaventris Pope and Hairston

Plethodon jordani rabunensis Pope and llariston

Flethodon jordani teyahalee Hairston

Plethodon jordani clensonae Brimley

Plethodon glutinosus glutinosus (Green)

Plethodon p:lutinosus albagula Grobnan




- 29 -


The Western Plethodons


The Western Plethodons include five species, none of which,

except for dunni and vehiculum, appear to have close affinities. P.

vandykei, with three presently recognized subspecies, vandykei, larselli,

and idahoensis, occurs in coastal Washington, northwestern Oregon, and

northern Idaho. P. dunni is known from southwestern Washington and

coastal Oregon. P. vehiculum ranges on the coast from central Oregon to

southern British Columbia, including Vancouver Island. P. products

(formerly P. elongatus) occurs in a small area in northwestern Cali-

fornia and southwestern Oregon. P. neonexicanus is known only from the

Jemez Mountains of New Mexico.


Stebbins (1951) has recently summarized the knowledge of the

amphibians of western North America, and has given detailed descriptions

of all five species in life. Little new information on individual or

geographic variation can bo offered here since I have had no field ex-

perience with these animals and only four forms (idahoensis, dunni,

vehiculum, and products) have been examined in life. Reference should

be made to Stebbins' book for further information on these forms.


Radiographs of a series of each of the forms exceptt the re-

cently described P. v. larselli) were taken and the number of trunk

vertebrae in each form is shomn in table I. The usual number of trunk

vertebrae in each species of I.estern Plethodon is different, and none

of the known forms possess 18 trunk vertebrae as the modal number.


Except for vehiculum, all of the stern Flethodons are fairly

large in size. All but neomexicanus possess a stripe phase in the adult,

and all but dunni and vehiculum have webLed toes. The vomerine tooth






- 30 -


TABLE I


THE NUMBER OF TRUNK VERTEBRAE IlN -ST''I"J FORMS OF THE GFIIUS PLETHODON


Number of trunk vertebrae


Form


15 16 17 18 19 20


v. vandykei

v. idahoensis

dunni

vehiculum

Droductus

neomexicanus


10 32


1 15






- 31 -


counts of all of the western forms are low compared to Eastern Large

Plethodons of similar size; vehiculum and products have especially

low vomerine counts.


P. vandykei is unusual in possessing a parotid gland and in

having the lowest number of vertebrae in the genus. P. neomexicanus

is unusual in being the only species in the genus that has a reduced

fifth toe on the hind limbs.


Plethodon dunni and Plethodon vehiculum appear to be rather

closely related. P. dunni attains a larger size than P. vehiculum and

has more vomerine teeth. P. vehiculum usually has one more trunk ver-

tebra than dunni. P. dunni has more aquatic habits than vehiculum. The

color of the dorsal lipophores in the striped phase of the two forms is

different. The stripe of dunni is usually greenish yellow, while that

of vehiculum varies from reddish tan to yellow. The toes are very

slightly webbed in both and most other structural features are similar.


The other western Plethodons possess a considerable amount of

webbing on their toes. The amount of webbing on the toes does not ap-

pear to be correlated with aquatic tendencies, since the two most aqua-

tic forms, dunni and vandykei, represent extremes in the absence and

presence of webbing in the western forms. The forms vehiculum and ro-

ductus, both with more terrestrial tendencies, also differ greatly in

amount of webbing between the toes. Moreover, in the eastern United

States, some of the forms have webbed toes, while others do not, yet all

are terrestrial.


There is no doubt that dunni and vehiculum should be regarded





- 32 -


as distinct species, since the two occur sympatrically through most of

the range of dunni without any evidence of interbreeding. A study of

this relationship and the possible differences in the habitat niches of

the two forms, suggested by Stebbins (1951: 65), is much to be desired.


The forms, vandykei, products, and neomexicanus all seem to

differ from each other as much or more than species groups in the Eastern

Flethodons. The fact that they are so distinct morphologically makes it

difficult to determine-their relationships. In all probability, these

forms have been differentiating for a long period of time. There has

probably been a considerable amount of extinction of humid forest dwell-

ing western N'orth American amphibians due to climatic changes which have

occurred during or since the Tertiary. The present discontinuous dis-

tribution of this genus in the area, with only two species recorded in

the Rocky fountains, separated by almost a thousand miles, supports

this view. Without doubt there are probably other undiscovered forms

of this genus in the western United States. Many salamanders of the

region are difficult to collect, except during extremely favorable con-

ditions, and some may have been overlooked in spite of intensive col-

lecting. There remain many areas in which there has been very little

collecting, and these nay be profitably searched for members of this

genus. Lowe (1955: ?50), for example, believes that plethodontid sal-

amanders will be found in the higher mountains of Arizona because con-

ditions there are similar to those where they have been taken in other

areas.


The relationships of these salamanders would seem to be best

indicated by the phylogeny shown in figure 3.







- 33 -


dunni vehiculum products neomexiconus


Sug -ested phylogerW of the Vestem plethodons.


Figure 3.






- 34 -


Plethodon vandykei Group

Plethodon vandykei

This species is koon from the Coast Range, Olympic Mountains,

and Cascade Mountains of western ;ashington; Coeur d'Alene Lake, in

northern Idaho; and from Larch Mountain, iultnomah County, Oregon. The

Idaho population was described as a distinct species (idahoensis) by

Slater and Slipp (19O0: 38), but has been considered a subspecies of

vandykei by Lowe (1950: 93)-and more recent workers. Although the Idaho

and I.ashington populations of this species are usually believed to be

isolated from one another, Savage (1952: 183) suggests that they may be

connected by a corridor of high humid country along the international

boundary between Y.ashington and British Columbia. The two forms, idaho-

ensis and vandykei, apparently differ mainly in color characters.


In 1954, Burns discovered Plethodon vandykei in Ifultnomah

County, Oregon. Two specimens from Larch Mountain were unusual in pos-

sessing red bellies. A series of eleven specimens collected on the north

side of the Columbia River, at Archer Falls, Skamania County, Washington,

also agreed in coloration with the Oregon specimens and this red-bellied

form was described as the third subspecies of vandykei, P. v. larselli.


Stebbins (1951: 80-1) states that there are two color phases

of P. v. vandykei. The light phase has the color of the belly and sides

very similar to that of the dorsal stripe, while the dark phase has a

darker lateral and ventral pigmentation.

This species is usually found in very damp situations and thus

replaces P. dunni ecologically, as well as geographically. It has re-






- 35 -


cently been shown that their ranges do overlap slightly in Multnomah

County, Oregon (Burns, 1954: 85) and Pacific County, Washington (Storm,

1955: 64-5).

Plethodon vandykei vandykei Van Denburgh

Plethodon vandykei Van Denburgh (1906: 61). Dunn (1926: 151-3). Slater

(1933: 44). Bishop (1943: 275-8). Storm (1955: 64-5).
Plethodon vandykei vandykei Van Denburgh. Lowe (1950: 93). Stebbins

(1951: 80-4). Stebbins (1954: 56-7). Slater (1955: 132-3).

Type:- CAS 6910 (now destroyed), collected at Paradise Valley, Mount

Ranier National Park, -ashington, by E. C. Van Dyke, in July, 1905.

Diagnosis:- A Western Plethodon with 15 trunk vertebrae. It differs from

idahoensis in having a vider dorsal stripe, lighter ground color, and by

presence of yellow pigment like that found on the dorsum on the proximal

segment of the limbs.

Range:- Known from western 'ashington, from Clallam County south to Pa-

cific County and east to Pierce County (see figure 4).

Description:- This subspecies has not been seen in life and only four

preserved specimens were examined. Stebbins (1951: 80-1) gives detailed

color notes on this form.

Costal grooves usually number 14 and the trunk vertebrae 15.
In the available specimens, vonerine teeth range from 7 to 10 in a series.


The largest individual is 56 rm. in snout-vent length.





-36-


Figure h. The distribution of the subspecies of Plethodon vandykei
in Washington, Oregon, and Idaho.






- 37 -


Plethodon vandykei idahoensis Slater and Slipp

Plethodon idahoensis Slater and Slipp (1940: 38). Slater (1941: 81, 85,

103). Bishop (1943: 259-61). Hilton (19h8: 120).
Plethodon vandykei idahoensis Slater and Slipp. Lowe (1950: 93).

Stebbins (1951: 80-4). Stebbins (195s: 56-7).

Type:- USNi 110o5o, an adult male, collected at the northeast corner of

Coeur d'Alene Lake, Kootenai County, Idaho, at an elevation of 2160 feet,

by James R. Slater, on September 13, 1939.

Diagnosis:- A race of Plethodon vandkei in which the yellow or orange

dorsal stripe is narrower than in P. v. vandykei and contrasts sharply

with the lateral black ground color, and the proximal segments of the

limbs are dark in color.

Range'- Known only from the type locality in northern Idaho (figure 4).

Description:- The back has a dorsal orange or yellow lipophore stripe.

The rest of the animal is pigmented with melanophores, except for nel-

anophore gaps on the chin where there are yellow lipophores. There are

also brassy guanophores in the iris, and a few scattered white guanophores

on the belly and sides.

The costal grooves usually number lh, the trunk vertebrae

15. Vomerine teeth range from 5 to 12 in a series. Of 15 specimens
examined, the largest is 56 mm. in snout-vent length.


Plethodon vandykei larselli Burns
Plethodon vandykei larselli Bums (19h4: 83-7).





- 38 -


Type:- USNM 1Jh129, an adult male, collected on the north slope of
Larch Mountain, three miles from the summit on the Multnomah Falls

Trail, Multnomah County, Oregon, on May 24, 1953, by Douglas M. Burns.

Diagnosist- A race of Plethodon vandykei in which the ventral color is

cardinal red to reddish orange.

Range:- Known only from the type locality and from Archer Falls, Ska-

mania County, Uashington (figure 4).

Description:- Specimens of this subspecies have not been examined and

reference should be made to the original description for details of

structure and coloration.



Plethodon vehiculum Group

Plethodon dunni Bishop

Plethodon dunni Bishop (1934: 169). Jewett (1936: 71). Fitch (1936:

637-8). Graf, Jewett, and Gordon (1939: 102). Gordon (1939: 55-6).

Slater (1939: 154). Bishop (1943: 242-6). Stebbins (1951: 68-72).

Stebbins (1954: 54-5). Storm (1955: 64-5). Dumas (1955: 65).

Slater (1955: 132).

Type:- USNM 95196, an adult female, collected just outside the city

limits of Portland, Clackanas County, Oregon, by Stanley G. Jewett, Jr.,

on January 13, 1934.

Diagnosis:- A Western Plethodon with 16 trunk vertebrae.

Range:- Curry County, in southeastern Oregon, north to Pacific County,






- 39 -


Washington, east to the western slope of the Cascade fountains (figure

5).

Description:- The dorsal stripe is made up of yellowish green lipo-

phores. This pigment also occurs in abundance on the sides, but is re-

duced on the belly. On the lower sides and belly there are a few

yellowish guanophores., There are brassy guanophores in the iris.


The costal grooves usually number 15, the trunk vertebrae 16.

The medial end of the vomerine series projects posteriorly toward the

parasphenoid patches, so that the two vomerine series form a V. Vomer-

ine teeth range from 4 to 13 in a series. The largest specimen examined

is 65 mm. in snout-vent length.


P. dunni is closely related to P. vehiculum, but differs in

size, coloration, number of vomerine teeth and number of body segments.

It occurs from southwestern Washington south through coastal Oregon.

This species occurs sympatrically with P. vandykei in southwestern Wash-

ington, with P. products in southwestern Oregon, end with P. vehiculum

throughout most of its range.


P. dunni is apparently the most aquatic Plethodon. Stebbins

(1951: 70) states that it is almost invariably found in places that are

saturated with water, and that it will often take to the water in an

attempt to escape capture.


Most individuals possess a dorsal stripe that is greenish

yellow in color, but Stebbins (1951: 69) reports that nelanistic speci-

mens lacking the dorsal stripe have been found in Bcnton County, Oregon.






- hO -


S. 0o 0o








Figure 5. The distribution of Plethodon dunni in Oregon and south-
western V'ashing'mn.






- 41-


Plethodon vehiculum (Cooper)
Ambystoma vehiculum Cooper (1860: pl. 31, fig. 4).
Plethodon intermedius Baird (in Cope, 1867: 209). Cope (1869: 100).
Strauch (1870: 72). Boulenger (1882: 57). Cope (1883: 23).
Garman (1884: 38). Cope (1889: 15l-7). Cox (1907: 52). Van
Denburgh (1916: 218-9). Fowler and Dunn (1917: 25). Hardy
(1926: 22). Kermode (1926: 35). Dunn (1926: 154-6). Slevin
(1928: 52-5). Logier (1932: 317-8). Slater (1933: i4). Svihla
(1933: 39). Slater (1934: 1l0-1). Slevin (1934: 46). Cowan
(1937: 18).
Plethodon vehiculus (Cooper). Bishop (1934: 171). Jewett (1936: 71).
Watney (1938: 89). Slater (1939: 154). Graf, Jewett, and Gordon
(1939: 10-1). Brown and Slater (1939: 9).
Plethodon vehiculum (Cooper). Slater (190L: 43). Slater and Brown
(1941: 75-7). Bishop (1913: 278-81). Stebbins (1951: 84-7).
Stebbins (1954: 57-9). Logier and Toner (1955: 17). Slater

(1955: 133-4).

Type:- Apparently no longer in existence. The type locality is Astoria,
Oregon.

Diagnosis:- A Western Plethodon with 17 trunk vertebrae.

Range:- From Coos County, in southwestern Oregon, north to southwestern
British Columbia, including Vancouver Island (figure 6).

Description:- Lipophores in the region of the dorsal stripe may be ab-

sent, producing a uniformly colored phase, or present, resulting in a






















































Figure 6. The distribution of Plethodon vehiculum in Oregon, Wash-

ington, and British Columbia.


- 2 -






- 43 -


striped phase. The color of the dorsal stripe is quite variable, vary-

ing from light yellow, through yellow, orange, and red to brown. The

lipophores are absent from the sides, but are present on the belly. The

sides are black with only a few small white guanophore spots. These are

also present on the belly, and along with the melanophore and lipophore

pigmentation give it a mottled appearance. Small white guanophore spots

similar to those on the sides, as well as smaller brassy flecks, are

present on the dorsum of the dark phase individuals. Brassy guanophores

are present in the iris.


Costal grooves usually number 16, trunk vertebrae 17. Vomer-

ine teeth range from 3 to 7 in a series and the two series converge

posteriorly to form a V as in dunni. The largest specimen 49 examined

is 51 mn. in snout-vent length.

This species is the smallest Western Plethodon and has the

largest range. It is the most abundant Plethoden over most of its

range. P. vehiculum is superficially quite similar in appearance to

the eastern P. cinereus. Dark and striped phases are present and the

belly is mottled in both species. The similarities between the two are

probably due to convergence or parallel evolution because in all other

characters, P. vehiculum is morphologically more similar to the other

Western Plethodons.


Plethodon products Group


Plethodon products, new name

Plethodon elongatus Van Denburgh (1916: 216-8). (non) Salamandra elon-

gata Valenciennes in Dumeril and Bibron (1854: 84). Grinnell and









Camp (1917: 134). Storer (1925: 21, 10). Dunn (1926: 156-8).

Slevin (1928: 55-7). Slevin (1934: 46-53). Wood (1934: 191).

Fitch (1936: 638). Wood (1939: 110). Gordon (1939: 13, 30, 56).

Bishop (1943: 246-9). Hilton (1946: 45). Stebbins and Reynolds

(1947: 41-2). Stebbins (1951: 72-6). Stebbins (1954: 55-6).

Type:- CAS 29096, collected at Requa, Del Norte County, California, by
J. R. Slevin, May 22-26, 1911.

Diagnosis:- A Western Plethodan with 19 trunk vertebrae.

Range:- Southwestern Oregon and northwestern California (figure 7).

Description:- The belly of this species is very dark with a few sact-

tered white guanophore spots. The orange to reddish brown dorsal lipo-

phore stripe is usually brighter in juveniles than in adults, where it

is often reduced or absent. Often it is divided into a right and left

dorsolateral stripe by the presence of medial melanophore pigment.

The costal grooves usually number 18, the trunk vertebrae 19.

Vomerine teeth range from 4 to 7 in a series. This is a large species,

the largest of 18 specimens examined is 66 mn. in snout-vent length.

This is the most elongate of the Western Plethodons in the

Pacific Northwest. Its greater number of body segments distinguishes

it from the species vanXkei, dunni, and vehiculum, and its webbed toes

also distinguish it from the last two. It is probably more closely re-

lated to P. neomexicanus than the other Western Plethodons. P. products
has a rather limited distribution. It is known only from Trinity, Hum-

boldt, and Del Norto Counties, C'lifornia; and Curry County, Oregon.



































Figure 7. The distribution of Plethodon products in California and

Oregon.


The distribution of Plethodon neomexicanus in New .ecxico.


- 45 -


Figure 8.








- 46 -


Stebbins (1951: 76) believes that specimens from the interior of the

range differ in several respects from those on the coast.

Van Denburgh described this species in 1916, apparently with-

out realizing that the name he proposed, elongatus, was a junior homo-

nym of Salamandra elongata Valenciennes (in Dumeril and Bibron, 1854:

84) (= Flethodon glutinosus). This is a primary homonym due to the

fact that the name elongata is available for Plethodon glutinosus, al-

though the combination Plethodon elongatus has never been used for the

slirr salamander. No subsequent worker has proposed a new name to re-

place the preoccupied name of the Del Norte salamander and it has al-

ways been referred to as Plethodon elongatus. This species has been

known by the name elongatus since 1916 and it is unfortunate that the

name has to be changed, but there is no alternative than to propose a

new name in place of the preoccupied elongatus. The name products

also refers to the elongated condition of this species.


Plethodon neomexicanus Group


Plethodon neomexicanus Stebbins and Riemer

Eurycea multiplicata (Cope). Dunn (1926: 312-4, part).

Plethodon neomexicanus Stebbins and Riemer (1950: 73-80). Stebbins

(1951: 76-9). Stebbins (1954: 58).


Type:- MVZ 49033, an adult male, collected 12 miles west and 4 miles
south of Los Alamos, Sandoval County, New I'xico, at an altitude of

about 8750 feet, by Robert Stebbins, on August 14, 1949.


Diagnosis:- A western Plethodon with 20 trunk vertebrae.






- 47 -


Range:- Known only from the vicinity of the type locality in the Jemez

Mountains of New Mexico, (figure 8).


Description:- This species has not been examined in life and reference

should be made to the original description for an account of the pig-

mentation.

Costal grooves are usually 19, trunk vertebrae 20. Vomerine

teeth range from 4 to 11 in a series in the 7 specimens examined. This

is a large species, the type is over 70 mE. in snout-vent length. Ac-

cording to the data listed in Stebbins and Riemer (1950: 75), sexual

maturity is not reached until approximately 50 m~. in snout-vent length.


This is an elongated salamander, with the largest number of

trunk vertebrae of any of the western forms. It is unusual in that most

specimens have only one phalanx on the fifth toe of the hind limb in-

stead of the usual two phalanges. This is probably an intermediate

stage in the loss of the fifth toe and reduction to the four-toed con-

dition Iknown to have occurred in several unrelated salamander genera.


The dorsal stripe is apparently absent in adults, but present

in juveniles. This form is unique among the western Plethodons in lack-

ing the striped phase in the adult, and also in possessing a large num-

ber of brassy guanophores on the dorsum.



The Eastern Small Plethodons


The Eastern Small Plethodons are characterized by their small

size. Except for P. welleri, they possess a higher number of trunk

vertebrae and are relatively more elongate than the Eastern Large









Plethodons. Four species are included in this group. P. welleri has

a restricted range, and is known only front a few counties in north-

western North Carolina and adjacent Tennessee and Virginia. P. rich-

mondi is a polytypic form with three described races, and probably

others which have not yet been defined. Its range centers in the

Appalachian Plateaus Physiographic Province and it also occurs in im-

mediately adjoining areas of several adjoining Provinces. P. cinereus,

with three subspecies, has the widest range of any Eastern Small Pleth-

odon, occurring over most of eastern United States and southeastern

Canada, except in the region occupied by P. dorsalis. The range of

P. dorsalis is centered in the Interior Low Plateaus Physiographic

Province. P. d. angusticlavius is here regarded as a subspecies of

dorsalis, rather than of cinereus, as originally described by Grobman

(19h4: 302).


The four species form a natural group of closely related forms.

All possess webLed toes, a pigmented poritoneumJ, lowi vomerino tooth

counts, brassy flecks on the dorsum, and white guanophore spots on the

belly and sides. Two species, cinereus and dorsalis, are more similar

to each other than either is to welleri or richmondi. Both cinereus

and dorsalis typically possess at least as much white as black pigment

on their bellies, and both are characterized by the presence of a red-

backed phase in which there is a dorsal band of red or yellow lipophore

pigmentation on the dorsum of the body and tail. In richmondi and

welleri, the white pigment on the belly is limited to small spots and

there is a much greater amount of black pigment than vhite. The red-

bicked phase, present in all of the races of dorsalis and cinereus,


- h8 -







- 49 -


is always absent in richmondi and welleri.


P. r. richmondi is the largest form, and also possesses the

greatest number of trunk vertebrae, but both size and number of ver-

tebrae are slightly reduced in the race popei, and decrease still more

in the subspecies netting. Geographic variation in the number of

trunk vertebrae is most pronounced in this species, with a range of

18 to 24 in all of the subspecies. P. welleri is less elongate and has

fewer vertebrae than other small eastern species. It is unusual in its

possession of ovarian eggs pigmented with melanophores, as well as in

having an extremely dark parietal peritoneum. The brassy flecks on

the dorsum of welleri are concentrated to form large patches, giving it

a unique appearance not found in any other Plethodon. Its nearest rel-

ative is probably P. r. netting, but welleri is closest geographically

to P. r. popei.


In most of their ranges, cinereus and dorsalis are allopatric

forms, but in several areas where their ranges meet, there are records

of the two occurring together in the same locality (see below). In

each of these instances, there is no good evidence for hybridization or

intergradation between the two forms, and for this reason they are

usually considered as distinct species. They differ in several pig-

mentation characters, as well as in the average number of trunk ver-

tebrae and the shape of the mental gland in adult males. P. d~.rsalis

usually has 19 trunk vertebrae (range 18-20). Two races of cinereus

typically have 20 trunk vertebrae (occasionally 19 or 21), while the

third subspecies (polycentratus) usually has 21 or 22 (rarely 23)

trunk vertebrae. In all of the Eastern Small Plethodons except dorsalis,









the mental gland is rather poorly developed and difficult to differen-

tiate (except, perhaps, during the breeding season) from the adjacent

portion of the chin. In dorsalis, on the other hand, it is a distinct

rounded elevated gland, as in the Eastern Large Plethodons. Dunn (1926:

24) suggests that the striped pattern or dorsalis is more primitive than

that of cinereus, since the paired red dorsal spots of several primitive

plethodontids might be expected to pass through a zig-zag striped phase

similar to that of dorsalis before evolving into a uniformly straight-

edged stripe.


In 19L4, Grobman described the narrow-striped populations oc-

curring in southwestern Missouri and northwestern Arkansas as a distinct

subspecies, Flethodon cinereus angusticlavius. This form possesses sev-

eral characteristics that appear to link it more closely with dorsalis

than with cinereus. It usually has 19 trunk vertebrae, as in dorsalis,

while cinereus populations to the northeast and south of the range of

angusticlavius usually have 20 trunk vertebrae. The mental gland is of

the dorsalis type and in some specimens the dorsal stripe has irregular

edges anteriorly, resembling very closely the pattern in dorsalis.

Living specimens of angusticlavius have not been examined, so no accur-

ate information is available on pigmentation characters, but the other

characters strongly suggest that angusticlavius should be linked sub-

specifically with dorsalis rather than cinereus.


The elongation and increased number of trunk vertebrae in P.

r. richmondi, P. r. popei, and P. c. polycentratus would appear to be

a marked specialization of the usual plethodontid type. The forms

welleri and dorsalis would appear to have retained more primitive char-


- 5C -






- 51 -


acteristics, but both have probably changed considerably from their

common ancestor. The relationships of the Eastern Small Plethodons

would appear to be best indicated by the phylogeny outlined in figure

9.

Plethodon welleri Group


Plethodon welleri Walker

Plethodon welleri Walker (1931: 48-51). Walker (1934: 190). Bishop

(1943: 285-7). Grobnan (19h4: 313). Snyder (1946: 174). Hoff-

man and Kleinpeter (1948a: 107). Hoffman (1953: 86-7).


Type:- USNM 84135, an adult male, collected on Grandfather mountain at
an altitude above 5000 feet, near Linville, North Carolina, by W. H.

Weller and Ralph Dury, on August 27, 1930.


Diagnosis:- A dark-bellied, Eastern Small Plethodon with 17 trunk ver-

tebrae and abundant dorsal brassy spotting.


Range:- From Yancey County, North Carolina, northeast in Tennessee

and North Carolina to Mt. Rogers and Vhite Top Mountain, Virginia (fig-

ure 10).


Description:- The dorsal pattern consists of large anastomosing patches

of brassy guanophore spots. These spots appear to consist almost en-

tirely of brassy guanophores, the white type of guanophore pigmentation

present on the sides of welleri and on the dorsum of most plethodons, is

not evident on the back of welleri. The brassy pigment is more concen-

trated than in any other form and the appearance of the dorsal spots is

similar to the large white spots of glutinosus, except for the color,





- 52 -


cinerous serrotus popei
polycentrotus ongusticlovius dorsolis richmondi nettingi



cinereus dorsolls
richmondi















Figure 9. Suggested phylogeny of the Eastern Small Plethodons.


Figure 10. The distribution of Plethodon welleri in North

Tennessee, and Virginia.


Carolina,










which is brassy. Small white spots (0.1-0.2 am.) are scattered over the

belly and are similar to those of P. r. popei. The lateral spots (up to

1 mm. in diameter) are white with a slight amount of brassy flecking.

The costal grooves usually number 16, the trunk vertebrae 17.

Vomerine teeth range from 2 to 7 in a series. In the 39 specimens ex-

amined, maximum known snout-vent length is between 45 and 50 am. Sexual

maturity is reached at about 35 mm. in snout-vent length.


P. welleri is the least elongate of all of the Eastern Small

Plethodons, with the same number of trunk vertebrae (17) as in most of

the Eastern Large Plethodons. In body build it resembles the large

forms more than any other Eastern Small Plethodon. Its small size, ver-

tebral structure, webbed toes, and pigmentation characters would appear

to associate it more closely with the other small eastern forms.


This species has long been believed to be restricted to high

elevations and was recorded only from Flat Top Mountain and Grandfather

Mountain, North Carolina; and Mt. Rogers and ihite Top Mountain, Virgihia.

Recently, Hoffman (1953: 86) collected the species at an elevation of

2500 feet in Johnson County, Tennessee. I visited this locality during

the summer of 1955, and although conditions were very dry, succeeded in

collecting one young specimen, thus confirming Hoffman's record. This

species may well occur in suitable habitats at low altitudes throughout

the region.


Walker (1934: 190) has pointed out that individuals from YVhite

Top Mountain, Virginia, possess more spots on their venters than indivi-

duals from the type locality, Grandfather Mountain, North Carolina.


- 53 -









Specimens from Mt. Rogers (USNM 124h21-9) also possess spotted bellies.

One specimen from Flat Top Mountain, Yancey County, reported by Snyder

(1946: 174) is also described as having a more conspicuous mottling on

the venter. Perhaps the Grandfather Mountain population is unusual in

lacking this characteristic. The low altitude specimen from Tennessee

has a greater amount of dorsal brassy pigment than eight living topo-

types from Grandfather Mountain. I have not examined living specimens

from any of the other localities.


P. welleri is unusual in possessing dark ovarian eggs that

are pigmented with nelanophores. Its closest relative is probably P.

richmondi, which it resembles by the presence of a dark belly. P. r.

nettingi usually has 19 trunk vertebrae, although occasional specimens

possess 18 or 20. Since a small percentage of welleri also have 18

trunk vertebrae (table II), there is slight morphological overlap be-

tween the two forms in this character.


TABLE II

THE NUMBER OF TRUNK VERTFERAE IN GEOGMRPHIC SAMPLES OF PLETHODON WELLERI

Number of trunk vertebrae
Locality
16 17 18

Grandfather Mountain, North Carolina 25

Johnson County, Tennessee 1

White Top Mountain, Virginia 4

Mt. Rogers, Virginia 1 6 2


-S5 -










Plethodon richmondi


In 1938, two new Eastern Small Plethodons, P. richmondi and

P. nettingi, were described by Netting and Mittleman (1938: 297) and

Green (1938: 295). Both are characterized by their dark bellies and

brassy dorsal flecking. Available samples of richmondi from Ohio and

West Virginia possess 20 to 22 costal grooves (21 to 23 trunk vertebrae),

while P. netting has 17 to 19 costal grooves (18 to 20 trunk vertebrae).

They both differ from P. c. cinereus in the number of vertebrae, and they

are distinguished from the entire cinereus group by their dark bellies

and lack of red pigment. Most previous writers are in agreement that

these two forms, richmondi and nettingi, are more closely related to

each other, and to welleri, than they are to cinereus and dorsalis

(Netting and Mittleman, 1938: 292; Green: 1938: 298; Grobman, 1944: 311).


In 1949, Grobman described another flAcked plethodon (P. hul-

dae) which he assigned to this group because of its heavily flecked

dorsum. This form possesses 20 trunk vertebrae, the same as cinereus,

and also has a mottled belly as in cinereus. Rabb (1995: 261-3) and

Muchmore (1955: 170-2) present convincing evidence that huldae, although

possessing brassy dorsal flecks similar to those of richmondi and net-

tingi, is actually based on dark-phase specimens of P. cinereus. Much-

more (1955: 172) goes so far as to state that "since P. c. cinereus

possesses brassy flecks on the dorsum and since P. huldae can no longer

be considered valid, it is certainly desirable that the concept of a

welleri group of Plethodon distinct from a cinereus group be abandoned."

Obviously these forms are all more closely related than any of than are

to the Eastern Large Plethodons or the Western Plethodons, yet the fail-


- 55 -









ure of one suggested character to define the group should have no bear-

ing on the fact that other features are useful in distinguishing it

(dark belly and lack of red dorsal stripe).


Netting and Mittlemau (1938: 292) state that specimens of P.

richmondi from Watauga County, North Carolina, differ from typical

richmondi, but do not give any evidence to support this contention.

Neither Bishop (1943: 239) nor Grobman (19lh: 312) include this locality

in their distribution maps of the range of richmondi, although Grobman

mentions the literature reference to the North Carolina specimens. More

recently, Hoffman and Hubricht (1954: 192) report richmondi from several

localities in southwestern Virginia and northwestern North Carolina, but

do not find any consistent differences between these specimens and topo-

types of richmondi. A study of their material and large series from this

area in the University of Florida Collection have shown, however, that

these southern richmondi are actually quite distinct in the number of

trunk vertebrae from more northern richmondi. This southern form has

recently been described as a new subspecies of richmondi, P. r. popei,

by Highton and Grobman (in press). No other character could be found

to distinguish preserved specimens of popei and richmondi. Living

specimens of P. r. richmondi have not been available during this study,

so it has not been possible to compare the pigmentation characters of

the two forms.


On the basis of the slight overlap in distinguishing charac-

ters, as well as the geographic replacement of the three forms, rich-

mondi, popei, and netting, Highton and Grobman (in press) suggested

that they should all be considered subspecifically related. One


- 56 -










matter, not discussed in their paper, is the apparent sympatric rela-

tionship of netting and richmondi. Although the two forms, to my

knowledge, have not been collected at the same locality, the entire

range of netting, as summarized by Brooks (1948), is surrounded by

records of richmondi.


Thurow (1955) has recently reported netting from Bedford

County, Virginia, a locality about equidistant from the nearest known

netting and popei localities. An examination of these specimens

(CNHM 60512-8) confirms Thurow's suggestion that they belong to this

group. Although they are poorly preserved, their dark bellies indi-

cate that they are not dark-phase cinereus. They differ from both

netting and pope, however, in costal groove count. Five of the seven

specimens have 19 costal grooves, while the other two have 18. The sam-

ple is small, but the probability that they were taken from a popula-

tion of netting similar to that in Randolph and Pocahontas Counties,

West Virginia, which has 19 costal grooves less than 5% of the time, is

very low. Four of the seven Bedford County, Virginia, specimens have

20 trunk vertebrae, two possess 19, and the remaining specimen is too

small to obtain an accurate count from the radiograph, but it probably

also has 20 trunk vertebrae, since it has 19 costal grooves. The pop-

ulation from which these specimens were collected probably represents

an intermediate one between pope and netting. If the usual number of

trunk vertebrae is 20, and this occurs with high frequency, this popu-

lation should probably be given separate subspecific nomenclatorial

status. At present, with only seven poorly preserved specimens avail-

able, too little is known of its variation to described it as new here.

Its importance lies in the fact that an intermediate population, often


- 57 -






- 58 -


possessing 20 trunk vertebrae, is known from a locality between the

ranges of netting (usually with 19 trunk vertebrae) and popei (usually

with 21 trunk vertebrae). This may be considered further evidence that

nettingi and popei are subspecifically related.


In the Valley and Ridge Province of Virginia, there exists a

population of P. r. richmondi which Hoffman and Hubricht (1954: 192)

believe to be different in both color and structural features from

other populations of P. richmondi. They do not state the ways in which

it differs, but an examination of preserved specimens from this region

(Alleghany and Rockingham Counties, Virginia) indicates that these

animals, although possessing a vertebral count similar to richmondi,

differ from it in having a mottled belly. Specimens from these coun-

ties are easily segregated from topotypical richmondi and ppei on the

basis of the character of the belly. In ten specimens of the mottled

bellied form for which vertebral counts are available, six possess 22,

three 23, and one 24 trunk vertebrae. The average is slightly higher

than the usual richmondi number, but does not differ significantly

from it. It is quite likely that examination of living specimens, and

the accumulation of more data on vertebral counts may reveal differ-

ences which will indicate that this population represents a distinct

nominal form. Specimens from other areas in the Valley and Ridge

Province should be studied in order to delimit the range of the mottled-

bellied richmondi.


The distribution of these forms indicates that there is

probably a series of five races, two of them undescribed, each of which

replaces its nearest relative geographically. One extre-" (nottingi)

occurs sympatrically with the two at the other extreme (richmondi and










the undescribed Virginia Valley and Ridge animal). Table III shows the

variation in the vertebral counts of these populations.


TABLE III


GEOGRAPHIC VARIATION IN THE NUMBER OF TRUNK VERTEBRAE IN PLETHODON

RICHMONDI


Number of trunk vertebrae
Sample
18 19 20 21 22 23 24 Mean

P. r. netting 1 22 1 19.0

Bedford County, Virginia 2 4 19.7

P. r. opei 4 51 12 21.1

P. r. richmondi 4 30 9 22.1

Alleghany and Rockingham 6 3 1 22.5
Counties, Virginia




The races of richmondi appear to form a ring of subspecies,

none of which is completely distinct morphologically from the related

adjacent forn (or forms). P. r. netting is not known below 3550 feet

elevation in the Cheat Mountains of West Virginia (Brooks, 1948) and

it thus may be completely isolated ecologically from P. r. richmondi.

There is no information on the upper altitudinal limits of richmondi

in the literature, but it is known from localities throughout most of

West Virginia. A similar case has been reported by Stebbins (1949) for
the western salarrander genus Ensatina, in which the two end forms of an

allopatric series of subspecies occur in immediately adjacent regions


- $9 -





- 60 -


without any evidence of interbreeding. An accurate knowledge of the 0-

cological distribution of the overlapping forms of P. richmondi is lack-

ing. There appears to be little or no overlap between the two terminal

races of Ensatina eschscholtzi, whereas in this case, one nettingg)

definitely occurs within the range of richmondi, for there are records

of the latter to the north of the Cheat Mountains in Pennsylvania, to

the east and weat in West Virginia, and to the south in Virginia.


In these series of populations we probably see the stages

through which the subspecies of richmondi have differentiated. All of

the forms probably have changed somewhat from the original stock, but

the close similarity of all the forms, except for the number of body

segments, would seem to indicate that this change has not been great.

It is difficult to determine v.hich form most closely resembles the an-

cestral condition. P. r. richmondi is unlike most plethodons in its

elongate form and would appear highly specialized. On the other hand,

it wide distribution and the fact that the eastern Valley and Ridge

population has a mottled belly, approaching that of the cinereus group,

might be considered evidence that this form is a more primitive gener-

alized animal. It is possible to reconcile these viewpoints, for it

is often found that a given form is specialized in some ways, yet re-

tains certain other primitive characteristics.


Since P. r. netting is apparently limited to high altitudes

in the Cheat Mountains, it could be interpreted either as a specialized

type adapted to this habitat, or as a relict of a once more widely dis-

tributed spruce forest population. At present, the evidence is not

sufficient to choose between the two alternatives. On the basis of its






- 61 -


fwer trunk vertebrae, netting would appear to be less specialized

than the other subspecies.

Intergradntion between the races of richmondi my occur in

certain regions. P. r. netting may now be completely isolated from

its nearest relative, but the ranges of popei and richondi probably

meet in eastern Kentucky, and intergradation riuht be expected. La-rg

samples from critical areas will be necessary before conclusive evi-

dence is obtained, since intergrades between popei and richfendi, for

example, would be expected to possess about an equal number of specimens
with 21 and 22 trunk vertebrae. Large series of specimens from each

county in and on both sides of the area of intcrgradation would be nne-

cscsry to determine its extent. Such series are not now available

from Kentucky, and hollow symbols representing literature records for

this region on the distribution map (figure 11), my represent popei,

intergradient, or richmondi populations,


Plethodon richnondi richmondi netting and Dittleman

Plethodon richmondi Notting and Mittleman (1938: 287). -:ettinC (1939:

50-1). Dury and Geesing (1940s 31). Bishop (1943: 272-5). Grob-

man (1944: 312). Wood (1945a: L 9). Wood (19LSb: 206-10). Getting
(19t6: 12). Wood (1946: 169). "ood and Duellran (1947: 3). Grob-

man (19b9: 135). -ichmiond (1952: 314). Green .and Walker (1954:

60). Duellman (195h: L0-5). HIoffman and Hubricht (1954k 191-3).
Plethodon richmordi richmoridi Netting and Uittleman. Highton and Grob-

man (in press).

Type:- CM 1189, an adult mle, collected in hitterr Park, Luntington,






- 62 -


Cabell County, West Virginia, at an elevation of 600-700 feet, by Neil

D. Richmond and N. Bayard Green, on October 15, 1938.

Diagnosis:- A dark-bellied Eastern Small Plethodon, usually with 22

trunk vertebrae (range 21-24), which completely lacks red pigment.

Range:- From Centre County, Pennsylvania, south through western Mary-

land, West Virginia, and northwestern Virginia, west to northeastern

Kentucky, and north to southern and eastern Ohio (figure 11).

Description:- This form has not been examined in life and nothing on

the pigmentation can be added to previously published accounts.

The costal grooves usually number 21, the trunk vertebrae 22.

Vomerine teeth range from 3 to 9 in a series. This is the largest

Eastern Small Plethodon, the largest specimen examined, from Alleghany

County, Virginia, is 60 am. in snout-vent length.


Plethodon richmondi popei Highton and Grobman

Plethodon richaondi Netting and Mittleman. Barbour (1953: 85-6).

Hoffman and Hubricht (1954: 191-3).

Plethodon richmondi popei Highton and Grobman (in press).

Iype:- UF 8226, a maturing male, collected at Comers Rock, Grayson-

Wythe County line, Virginia, by Arnold B. Grobman and Marc R. Grobman,

on August 5, 1955.


Diagnosis:- A race of Plethodon richmondi that usually possesses 21

trunk vertebrae (range 20-22).


RanEt- Known from Harlan County, Kentucky; Tazewell, Smyth, Grayson,






- 63 -


Figure 11. The distribution of the subspecies of Plethodon richmondi,

Solid symbols represent localities from vw Lch specimens have

been examined. Hollow symbols, represent literature records.






- 61 -


and ythe Counties, Virginia; and Ashe, Alleghany, and Watauga Counties,

North Carolina (figure 11). This form may have a greater range than

now known. Specimens from adjacent Kentucky and Kest Virginia have not

been examined. They may also belong to this race.

Description:- Living specimens possess both types of guanophore spots

on the dorsum. Small brassy flecks are very abundant and larger white

spots are also present. On the sides and venter there are larger white

or yellow spots.

The costal grooves usually number 20, the trunk vertebrae 21.

Vomerine teeth range from 3 to 8 in a series. The largest specimen ex-

amined is 48 mm. in snout-vent length. Sexual maturity is reached be-

tween 35 and h5 mm. in snout-vent length.


Plethodon richmondi netting Green

Plethodon netting Green (1938: 295-9). Bishop (1943: 266-9). Grobman

(19bh: 313). Brooks (1945: 231). Brooks (1948: 239-44).
Plethodon richmondi nettingi Green. Highton and Grobman (in press).


Type:- CM 10279, an adult male, collected on Barton Knob, near Cheat
Bridge, Randolph County, West Virginia, at an elevation of about L000

feet, by M. Graham Netting, on June 29, 1935.

Diagnosis:- A race of Plethodon richmondi that usually possesses 19 trunk

vertebrae (range 18-20).

Range:- Known from altitudes above 3500 feet in the Cheat Mountains of

Randolph and Pocahontas Counties, West Virginia (figure 11).






- 65 -


Description:- In life, the dorsum of this form is similar in colora-

tion to P. r. popei. The costal grooves usually number 18, the trunk

vertebrae 19. Vomerine teeth range from 3 to 8 in a series. The

largest specimen examined is h5 mm. in snout-vent length. This form

appears to have a smaller average size than the other races of rich-

mondi. Sexual maturity is reached at about 35 mm. in snout-vent length.


Plethodon cinereus Group


Plethodon dorsalis

The range of P. dorsalis centers in the Interior Low Plateaus

Physiographic Province, but does not appear to be restricted to it, for

records are available from several adjacent provinces. This species has

not heretofore been recorded from Georgia, but specimens are now avail-

able from four different localities in that state, three of which are

in the Piedmont Province. The southernmost record is from Upson County

(UIPZ 8557h), only a few miles above the Fall Line. Other records are

for Henry County (UF 8371 (3), UF 8413 (3)); Cobb County (UG 206 (6);

UG 275 (36)); and Dade County (ERA-JTN 12128 (12)). Probably the nat-

ural range of this form includes the western part of the state above

the Fall Line.


The sympatric occurrence of this species with P. cinereus in

Georgia, Tennessee, Indiana, Illinois, and Oklahoma will be discussed

below. The two forms are very similar in appearance, and it is often

difficult to identify individual specimens. For this reason a detailed

study of the two species in the areas of overlap will be necessary be-






- 66 -


fore it can be determined whether or not hybridization occurs.


In Georgia, where there is no overlap in the vertebral counts

of the two species, it is impossible, on other grounds, to identify a

small percentage of the available specimens. There are, however, sev-

eral differences between the two species in Georgia that are apparent

in a large majority of the available specimens. These include the shape

of the mental gland in adult males; the zig-zag stripe of dorsalis, com-

pared to the straight-edged stripe in cinereus; the presence of more red

pigment in front of the eyes on the head of dorsalis; the great reduction

in amount of melanophore pigmentation on the belly of dorsalis; and a

similar reduction in red lipophore pigmentation on the belly of cinereus.

The fact that there are a few exceptions to each of these species char-

acteristics might be considered evidence for hybridization were it not

for the fact that occasional specimens of each species from localities

in which the other form is absent show at least some of the same varia-

tions from the usual pattern. The fact that there is no tendency toward

an increase in the number of trunk vertebrae in Georgia dorsalis popula-

tions is also significant (see table IV). It may be seen from this table

that there is no strong tendency for an increase in the number of verte-

brae in samples of dorsalis that are from areas in which geographic over-

lap with cinereus occurs. We may conclude that there is, as yet, no

good evidence for hybridization between the two species, but that more

study is needed in all of the areas in which the two occur together.


There are no available records of dorsalis in southeastern

Missouri. This region should be explored in order to determine whether

or not dorsalis or angusticlavius are present. Smith (1918: 1) has






- 67 -


recently reported dorsalis from several localities in southeastern

Illinois.


TABLE IV

THE NUMBER OF TRUNK VERTEBRAE IN SAMPLESOF PLET]HDON DORSALIS


Number of trunk vertebrae
Sample
Same 18 19 20


P. d. dorsalis

Georgia* 2 49 2

Alabama 2

Tennessee

Great Smoky Mountains* 34 9

Van Buren County 1 h4 6

Marion County 2

Indiana 1 2

P. d. angusticlavius

Arkansas 23 5

Areas in which cinereus and dorsalis have been taken together at the
same locality.


Dunn (1926: 162) states that the unstriped phase in dorsalis

is confined to adults and that this phase is much lighter than the dark

phase of cinereus. The latter part of his statement appears to be true,

but in a series of 31 living specimens of dorsalis from Van Buren County,

Tennessee (UF 8394), there are five juveniles (16 to 18 mm. snout-vent

length) that are definitely of the dark phase. As in adults, there is






- 68 -


a slight amount of red pigment in the region of the back in which the

stripe is located in red-backed individuals, so that an outline of the

irregular stripe can be seen when held at a certain angle to the light.

The presence of the red pigment, as well as a reduction in the melano-

phore pigmentation does give the lead-backed phase of dorsalis a lighter

appearance than the corresponding phase of cinareus.

Grobman (19i4: 308) presents evidence to show that Baird

should be credited with the authorship of the name dorsalis. Under
Article 21 of the International Rules of Zoolcgical Nomenclature as

of 19h4, he was correct. Since 194h, however, the Rules have been
amended, so that Cope should now be recognized as responsible for this

name.


Plethodon dorsalis dorsalis Cope

Plethodon cinereus dorsalis Cope (1889: 138-9). Blanchard (1926: 368-9).

Bishop (19N3: 236-9). Parker (1948: 22). Chermock (1952: 29).

Plethodon erythronotus (Green). Garman (1894: 38).

Plethodon dorsalis Cope. Stejneger and Barbour (1917: 15). Dunn (1918:

460-2). Dunn (1926: 158-62). Mohr (1937: 4h). Parker (1937: 631
Parker (1939: 75). King (1939: 550-1). Swanson (1939: 687).

Grobman (1944: 308-11). Smith (1.9h8: 1). Sinclair (1950: 50).

Mohr (1952: 59-60). Throw (1955: 62-3). Holman (1955:' 13).

Type;- USNM 3776, collected at Louisville, Jefferson County, Kentucky.

Diagnosis:- An Eastern Small Plethodon with the usual number of trunk

vArtebrae 19 (range 18-20), and a red dorsal stripe with irregular edges






- 69 -


in the red-backed phase.


Range:- From southern Illinois, Indiana, and southeastern Ohio, south

through Kentucky and Tennessee to northern Alabama and northwestern

Georgia (figure 12).


Description:- The dorsum of both color phases of P. d. dorsalis has

small white spots (0.2-0.4 mm.) as well as smaller brassy flecks. The

red-backed phase, in addition to the other pigments, contains a large

amount of red lipophore pigment. All of these chromatophores appear

to be identical with those of P. cinereus. The main difference between

the two species is in the abundance and distribution of the pigments.

The melanophore background in dorsalis is somewhat reduced, giving the

animal an overall lighter appearance than cinereus. The melanophore

pigment on the belly of dorsalis is greatly reduced with a corresponding

increase in the amount of red lipophores, so that the belly appears to

be mottled with red and white rather than black and white, as in ciner-

eus. In dorsalis there is a concentration of red pigment on the head

in front of the eyes. The lateral guanophore pigment is often yellowish

in color. The dorsal red stripe is quite variable. In some specimens

the edges are irregular for the entire length of the body, while in

others, they are irregular only in the anterior half or third of the

body. Tho latter condition is especially common in Georgia specimens,

and occasional specimens from Georgia have straight-edged dorsal stripes

as in cinereus. Specimens with straight-edged dorsal stripes have been

reported also from Indiana (Blanchard, 1926: 369) and Tennessee (Grob-

man, 1944: 309-10).






- 70 -


Figure 12. The distribution of the subspecies of Plethodon dorsalis.






- 71 -


The costal grooves usually number 18, the trunk vertebrae

19. Vomerine teeth range from 3 to 6 in a series. This is a small

species; sexual maturity is probably reached at about 30 mm. in snout-

vent length. The largest specimen examined is U4 mm. in snout-vent

length.

Plethodon dorsalis angusticlavius Grobman

Plethodon cinereus angusticlavius Grobman (19hh: 302). Dundee (19h7:

117). Bragg and Hudson (1951: 89). Bragg (1955: 27-8).


pet- AM~N h0366, an adult male, collected at Mud Cave, near Fairy Cave,
Stone County, Missouri, by B. C. Marshall, on October 1, 1927.


Diagnosis:- A race of Plethodon dorsalis in which the width of the dor-

sal band is usually less than one-third of the width of the body.

Range:- Southwestern Missouri, northwestern Arkansas, and adjacent Okla-

homa, north of the Arkansas River (figure 12).

Description:- Living specimens of this form have not been examined. Of

33 preserved specimens (all from Arkansas), 10 are of the striped phase:

11 show no trace of a stripe, and the remainder show only a faint stripe

on the body, but at the base of the tail, it widens and becomes much

more well-defined. Typically striped angusticlavius also possess a

wider and brighter dorsal stripe at the base of the tail than on the

body. The brightness of the stripe in this region appears to be due to

a reduction in melanophore pigmentation in the dorsal stripe. The dorsal

stripe is often-irregular anteriorly, as in dorsalis.


The costal grooves usually number 18, the trunk vertebrae 19.

Vomerine teeth range from 1 to 8 in a series. The largest specimen






- 72 -


examined is 43 mm. in snout-vent length.



Plethodon cinereus


The red-backed salamander is the most abundant terrestrial

salamandor over most of its range, which includes much of eastern North

America. Two distinct color phases are present, one with a prominent

red or yellow dorsal stripe, the other uniformly dark in appearance.

The frequency of these color phases varies from one locality to another.

In some places, both phases occur in approximately equal numbers, in

others, one type may be rare or absent. The proportional distribution

of these color phases in the various populations has recently been

studied by Thurow, and it is hoped that the results of his work will be

published in the near future. Occasional specimens have been reported

that lack the dark pigment and appear entirely red. One specimen from

New Jersey lacked the red pigment in life and had a colorless dorsal

stripe.


Plethodon dorsalis has been considered a subspecies of P.

cinereus by some workers (Cope, 1889: 138; Blanchard, 1926: 269; Bishop,

1903: 236), and as a distinct species by others (Dunn, 1926: 158; Grob-

man, 1944: 308). Grobmn- points out that although the two forms are

largely allopatric, at some localities, especially in southern Indiana,

the two occur together, Grobman examined the series of dorsalis that

King (1939: 551) reported from the Great Smoky Mountains of Tennessee

and confirmed their identification. Specimens of both species found

together at the same localities are represented in the GSJNP collection.

These localities are White Oak Sinks (elevation 1750 feet) and the Sinks






- 73 -


on the Little River (elevation 1600 feet). On the Tennessee side of the

Great Smoky Mountains, P. cinereus is known from 1600 to 5000 feet and

P. dorsalis from 1200 to 2200 feet.


The two species also occur sympatrically in the Piedmont of

western Georgia. At two localities, 3.8 miles north of McDonough, Henry

County; and 8.5 miles north of Thomaston, Upson County, the two species

have been taken together. At all other localities in western Georgia,

where either P. c. polycentratus or P. d. dorsalis have been collected,

the other form has not been taken.


Bragg (1955:-27) reports that he collected a specimen of P. c.

serratus and a specimen of P. d. angusticlavis at the same locality in

Cherokee County, Oklahoma. Apparently the two species often occur to-

gether where their ranges overlap. Smith (19h8: 1) published several

new records of dorsalis from southeastern Illinois. P. cinereus and

P. dorsalis probably also occur synpatrically in that state. There

appears to be no evidence of intergradation at any of these localities,

confirming the conclusion of Dunn and Grobman that dorsalis and cinereus

have reached the species level of differentiation.


P. cinereus shows little apparent geographic variation over

most of its range, except in the frequency of occurrence of the two

color phases referred to earlier. Only in the Piedmont of Georgia and

in Arkansas and Oklahoma, where the populations of this species have

apparently become isolated from the parent stock, so we find differen-

tiation of the magnitude required for the recognition of subspecies.

The Arkansas and Oklahoma race serratuss) possesses a dorsal band with











serrations at each costal groove. This characteristic is sometimes

slightly developed in some specimens of other populations, but never

reaches the frequency of occurrence or degree of development present in

serratus.


The Georgia Piedmont subspecies (polycentratus) differs from

other populations of cinereus in its increased number of trunk verte-

brae. Data on the number of trunk vertebrae in samples of P. cinereus

are listed in table V. It also differs from most cinereus in possessing

red pignent on the belly between the front limbs. The cinereus popu-

lations in western North Carolina and eastern Tennessee are character-

ized by the complete absence of the dark lead-backed phase. P. c. poly-

centratus differs from these adjacent cinereus populations by the

presence of the lead-backed phase (38% of the type series of polycen-

tratus are dark phase).


Sanders and Smith (1949: 28) report a specimen of Plethodon

cinereus from Fern Lake, near Nacogdoches, Nacogdoches County, Texas.

This specimen (OS 556) lacks the serrate edges in the dorsal stripe

that are present in serratus, the nearest race geographically. It has

19 trunk vertebrae, but this number is not unusual in other populations

of cinereus. Mr. Sanders, who collected the specimen, informs me (in

letter of November 12, 1955) that it was taken with a dip net from a

bunch of fruiting sphagnum moss near the shore line, a most unusual

habitat for this terrestrial species. It may represent an accidental

introduction by man, and the record needs confirmation by the collection

of additional specimens from Texas before this state can be included in


- 74 -






- 75 -


the natural range of this species.



TABLE V

GEOGRAPHIC VARIATION IN THE NUMBER OF TRUNK VEKrEBRAE OF PL;THODON
CINEREUS


Number of trunk vertebrae
Locality
19 20 21 22 23


P. c. cinereus

New Brunswick, Canada 3 li 1

Nova Scotia, Canada 2 2 1

Quebec, Canada 1 1

Indiana 6 2

Missouri 1

New Jersey 7

Virginia (Blue Ridge Province) 7 33 3

Eastern Tennessee and western
North Carolina 5 40 9

P. c. serratus

Arkansas 3 19

P. c. polycentratus

Georgia 21 24 2






- 76 -


Plethodon cinereus cinereus (Green)
Salamandra cinerea Green (1818: 356).

Salamandra erythronota Green (1818: 356). (Type locality, probably

vicinity of Princeton, Now Jersey.)
Plethodon cinereus (Green). Tschudi (1838: 58). Dunn (1926: 163-80).

Sauropsis erythronotus (Green). Fitzinger (18L3: 33).

Plethodon erythronotus (Green). Baird (1850: 285).

Ambystoma erythronotum (Green). Gray (1850: 37).

Salanandra puncticulata Valenciennes in Dumeril and Bibron (1854: 87).
Salamandra agilis Sager (1856: L29). (Type locality, Detroit, Michigan)

Plethodon erythronotus cinereus (Green). Cope (1869: 99).

Plethodon cinereus cinereus (Green). Davis and Rice (1883: 26). Bishop

(1941: 196-219). Bishop (1943: 232-6). Grobman (1944: 300-2).
Plethodon cinercus erythronotus (Green). Cope (1889: 135).

Plethodon huldae Grobman (1949: 136). (Type locality, Hlawksbill Moun-

tain, Madison County, Virginia.)

Type:- Dunn (1926: 165) states that the type is not known to exist. The

type locality is probably in the vicinity of Princeton, New Jersey.

Diagnosis:- A Small Eastern Plethodon with a black and white mottled

belly; usually with 20 trunk vertebrae; and a straight-bordered dorsal

stripe in the red-backed phase.

Range:- Nova Scotia, New Brunswick, southern Quebec and Ontario, Canada;

south through the eastern United States to North Carolina, eastern Ten-

nessee and Kentucky, Ohio, Indiana, Illinois and southeastern iLissouri

(figure 13).






- 77 -


Figure 13. The distribution of the subspecies of Plethodon cinereus.





- 78 -


Description:- The dark, unstriped phase is usually characterized by the

absence of red lipophores, the presence of small white spots on the

dorsum (0.07-0.2 mm in diameter), and numerous ;~..allor brassy flecks on

the head, back, and tail. The belly is mottled with black and yellow

or white guanophores. The lateral guanophore pigment is sir.lar to that

on the belly.


The red-backed phase has sides and belly similar to the dark

phase. The white spots and brassy flecks on the dorsum! are reduced in

the area of the dorsal stripe, but are present on the head and tail.

The color of the lipophores in the dorsal stripe is variable, ranging

from red to yellow.


The costal grooves usually number 19, the trunk vertebrae 20.

Vomerine teeth range from 3 to 9 in a series. The largest specimen ex-

amined is 51 ram. in snout-vent length. Sexual maturity is reached at

about 35 mm. in snout-vent length.


Plethodon cinereus serratus Grobman

Flethodon cinereus serratus Grobman (194: 306-8). Bragg (1952: 23Jh).

Bragg (1955: 27-8).

Type:- CNHM 3946., a female, collected on Rich Mountain, Polk County,

Arkansas, at an elevation of 2500 feet, by Karl P. Schmidt and C. M.

Barber, on March 23, 1938.


Diagnosis:- A race of Plethodon cinereus in which the edges of the

dorsal stripe are serrated at each costal groove.






- 79 -


Range:- West-central uplands of Arkansas and adjacent Oklahoma, south of

the Arkansas idver (figure 13). Also reported by Bragg (1955: 27-8)

from Cherokee County, Oklahoma (north of the Arkansas River).


Description:- The dark phase is apparently rare in this form, only one

of 33 specimens examined lacked the red dorsal band. In red-backed in-

dividuals, the serrations on the edges of the dorsal stripe are present

on the body, but not on the tail. The serrations are due to the fact

that the red pigment in the dorsal stripe extends ventrolaterally toward

the top of each costal furrow. The usual absence of melanophores in this

extension of the dorsal stripe makes the saw-tooth edge of the stripe

conspicuous to the naked eye.


The costal grooves usually number 19, the trunk vertebrae 20.

Vomerine teeth range from 3 to 8 in a series. The largest specimen ex-

amined is 46 mrm. in snout-vent length.


Plethodon cinereus polycentratus Highton and Grobman

Flethodon cinereus polycentratus Highton and Grobman (in press).


Type:- UF 8376, an adult male, collected 2 miles northeast of Palmetto,

Fulton County, Georgia, by Albert H. Highton and Richard Highton, on

February 2, 1954.

Diagnosis:- A race of Plethodon cinereus in which the usual number of

trunk vertebrae is 21 or 22 (rarely 23).


Range:- The Piedmont of western Georgia (figure 13).


Description:- This race is similar in coloration to the typical sub-





- 80 -


species. It differs in possessing a greater number of body segments.

Vomerine teeth range from 3 to 7 in a series. The largest specimen is

112 m. in snout-vent length. Sexual maturity is reached at about 35

mm. in snout-vent length.


The Eastern Large Plethodons


This group, as defined by Grobman (194h: 266) on the basis of

larger size and fewer costal grooves than the other eastern plethodons,

includes six species that inhabit the eastern portion of the United

States. Plethodon glutinosus is the most widely distributed and all

the other species occur within its range. P. yonahlossee occurs in

the Southern Section of the Blue Ridge Province north of the French

Broad River. P. ouachitae, superficially very similar to yonahlossee,

lives in the Ouachita Al'oitains of Arkansas and Oklahoma. The re-

cently described P. caddoensis is known only fro- the Caddo Mountains

of Arkansas. P. jordani includes eight subspecies, all occurring in

the Southern Section of the Blue Ridge Province. The range of P.

wehrlei is centered in the Unglaciated Allegheny Plateau Section of the

of the Appalachian Plateaus Province. (See Grobman (1941) for an analy-

sis of the distribution of these forms.) Two recently described species,

P. dixi and P. Jacksoni are here considered races of wehrlei, because of

their close morphological similarity to wehrlei and the f.ct that they

replace wehrlei geographically.


There has been much speculation on the relationships of the

species within this section of the genus. Dunn (1926: 23) believes

this to be the most primitive group of the genus, with yonahlossee the






- 81 -


most primitive form. Dunn and Grobman (1944: 276) both believe the

relationship between yonahlossee and wehrlei is close. Hairston and

Pope (1948) suggest that yonahlossee is closely related to jordani and

that close similarity between some jordani and glutinosus is merely con-

vergence, but others think that the closest relative of jordani is glutin-

osus. Bishop (1941) even considered one race of jordani (shermani) to
be racially related to glutinosus. On the other hand, Grobman (19h4)

believed that jordani and closely related forms (now all considered

races of jordani) are different enough to warrant the erection of a
separate group, that he called the metcalfi group, distinct from all

the other Eastern Large Plethodons.

Characters studied by previous workers in attempting to de-

termine the morphological similarity and hence the relationships of

these forms, include size, number of vomerine teeth, number of costal

grooves, degree of sexual dimorphism, and pigmentation. In the present

study, most of these characters have been reexamined using large series

of specimens that, for the most part, were not available to previous

workers. Several characters have been found to be extremely variable

and not diagnostic of any one form. For example, Bishop (1941: 18),
Grobman (19t4: 287), and Hairston and Pope (1948: 274) all believed

that the Plethodon jordani group has fewer vomerine teeth than P.

glutinosus. I have counted the vomerine teeth of 269 Florida glutinosus

(figure lW), 125 Virginia glutinosus (the sIae specimens used by Pope

and Pope, 1949, in their study, and our counts essentially agree), 45

glutinosus from the Coastal Plain of Virginia and North Carolina (figure

15), 115 specimens of P. J. jordani (figure 16), 27 specimens of P. J.

shermani, 72 specimens of P. J. netcalfi, 53 specimens of P. J. mela-






- 82 -


ventris, 14 specimens of P. J. rabunensis, 29 specimens of P. J. teya-

halee, and h4 specimens of P. a. unicoi. For a given size, the range

of variation is quite similar for each form, although there is often a

statistically significant difference in the variability and/or the

ontogenetic rate of change in the different species. The data of Popo

and Pope (1951) indicate that P. ouachitae has a similar range of var-

iation in number of teeth as glutinosus and jordani, although the av-

erage number of teeth is slightly higher. The number of teeth in 60

P. wehrlei (figure 17) seems to be less than in the above forms, while

Pope (1950) shows that P. yonahlossee than the above forms. 39 P.

caddoensis, although small in size, have even more teeth than P.

yonahlossee of the sa'e size range (figure 18). Except between adult

wehrlei and yonahlossee, or wehrlei and caddoensis, there is consider

able overlap in the vomerine tooth counts of all the Eastern Large

Plethodons, and even in the case of these, only adults can actually be

distinguished on the basis of this character. It is true that large

series of adults of species that differ greatly in size (e.g. P. g.

glutinosus and P. J. metcalfi) will differ in average number of vo-

merine teeth, but this is mainly a reflection of the difference in size

between the two forms. This character obviously cannot be used as an

aid in identification, since specimens of the two species that are the

same size will have similar womerine tooth counts.


Several other supposed differences mentioned in the litera-

ture are not useful in determining relationships. All, except P.

wehrlei, have a similar number of costal grooves (see table VI). Sex-

ual dimorphism in size is present in several forms that have been

critically studied. The type of lateral guanophore pigmentation varies









- 83 -


S P glutinosus
24 (Pennsulo Florida)
22* *
2 a
2C ** ooooo o -

16 oloo A &, 6 o- d

12 o *


20 2 0 3 0 5 5 5 6


20 25 30 35 40 45 50 55 60
SNOUT-VENT LENGTH (mm)


Figure 1Ji.


Vomerine teeth of 269 peninsula Florida P. glutinosus


plotted against snout-vent length.


P. glutinosus
(Coostol Plain Vo B N.C)


. .
*


20 25 30 35 40 45 50 55 60 65 70
SNOUT-VENT LENGTH (mm)


Figure 15.


Vonerine teeth of 45 P. glutinosus from the Coastal Plain


of Virginia and North Carolina plotted against snout-vent


length.


65 70


"





a









- 86 -


P. jordonl


*
*.. .

.'. . . . .
*. -. .

*..
*
** ** { *-*
* -
~ ~ ~ w *
** *


20 25 30 35 40 45 50 55 60 65

SNO'JT- IhNT LENGTH (mm)








Figure 16. Vomerine teeth of 115 P. J. jordani plotted against snout-


vent length.


P wehrler


.. ..
C
* C


0C
C C **C
C l
C C C
C ** C
C **


20 25 30 35 40 45 50 55 60 65

SNOUT-VENT LENGTH (mm)


Figure 17.


Vomerine teeth of 60 P. w. wehrlei plotted against snout-


vent length.


4 - -

















P. coddoensil


0
@0
*0 0000
.H 0
0@ 4 0
0 04 0 0
0 0


0U i5 0U 35 40 45 50


55 60


SNOUT-VENT LENGTH (mm)




Figure 18. Vomerine teeth of 39 P. caddoensis plotted against snout-


vent length.


Southern Northern


ouochitoe
coddoensis / vc


Figure 19. Suggested phylogeny of the Eastern Large Plethodons.


glutinosus


V __ __ __


- 85 -





- 86 -


somewhat within a species (both individually and geographically), but

there appear to be no consistent differences between species.


There remain, then, only a few characters that can be used

to diagnose or distinguish the Eastern Large Plethodons. Each species

is sympatric with, and often coexists in the same habitat with one or

more of the other species without any evidence of interbreeding. As

far as we know, each is a genetically distinct unit, and there is much

evidence to indicate that different species have different ecological

requirements. Yet preserved specimens that have lost their pigmenta-

tion characters are often extremely difficult to identify because of

the morphological similarity of most of these salamanders.


P. wehrlei is distinguished on the basis of a greater amount

of webbing on the toes, by having an average of one more trunk verte-

bra, and by possessing fewer vomerine teeth than the other species.

P. yonalilossee may be recognized by its distinctive color pattern in

life. It also differs from the others (except caddoensis) in posses-

sing a greater number of vomerine teeth. P. ouachitae has a similar

color pattern to that of yonnhlossee, but it has fewer vomerine teeth.

I have found no character that will consistently separate glutinosus

and jordani. The chin of glutinosus is usually darker than that of

jordani, but certain populations of both species more closely resemble

the other (some Texas glutinosus have very light chins, whereas P. J.

teyahalee and some P. J. rabunensis possess dark chins). The close

morphological similarity between all the forms, except P. wehrlei,

indicate that they are a closely related group of species, with only

rehrlei separated from the others by a number of characteristics.





- 87 -


TABLE VI


THE NUMBER OF COSTAL GROOVES AND VERTEBRAE IN TII EASTERN LARGE

PIETHODONS


Fo No. of costal grooves No. of trunk vertebrae
Form15 16 176 17 18 19
15 16 17 16 17 18 19


P. w. wehrlei

P. w. dixi

P. yonahlossee

P. ouachitae

P. caddoensis

P. glutinosus2

P. J. jordani

P. j. metcalfi

P. j. shermani

P. j. unicoi

P. j. melaventris

P. J. rabunensis

P. 3. teyahalee


3 57


2 48


30 4


12 2

16

35 2


32 319 14


19 212 15


11 68

10 78


8 1


6 6h 3

h 56 1


5 67

1 39


2 24h


1
This is the only form in the table in which the same specimens were used
to count both costal grooves and trunk vertebrae. The data on both char-
acteristics are included only to show the close correspondence between
the two counts. No other specimen of any of the forms listed in the
table was included under both costal grooves and trunk vertebrae.
2
For a geographic breakdown of these counts, see table VIII.






- 88 -


The young of yonahlossee, some wehrlei, and two races of

jordani are known to possess dorsal red spots. These disappear in adult

wehrlei (except P. w. jacksoni) and Aordani, and become incorporated in-

to the dorsal red stripe of yonahlossee. Ho information is yet avail-

able on the very small young of ouachitae, caddoensis, and several of

the races of jordani, but young ouachitae may possess then (see Pope and

Pope, 1951: 1i5). The red dorsal spots appear to be definitely absent

in the young of glutinosus. Red pigment was absent in all of the newly

hatched young I have examined from Florida (Highton, in press), as well

as in very young specimens from many other localities. Dunn (1926: 139)

records a specimen of P. glutinosus from Clayton, Georgia, that had tiny

paired red dorsal spots, but this may well be a specimen of P. .. sher-

mani, known to occur within 10 miles of Clayton. Cope (1889: 141) also

records young specimens of glutinosus from caves in Montgomery County,

Virginia, possessing these spots, but the specimens were probably P. w.

jacksoni.


A reduction in the amount of melanophore pigmentation on the

chin occurs in yonahlossee, ouachitae, caddoensis, wehrlei, most races

of jordani (except teyahalee and some rabunensis) and in some Texas

plutinosus (albagula).


Most species possess a dark belly, but the four northern races

of jordani (jordani, shermani, unicoi, and especially metcalfi) are

light-bellied, as are some southern wehrlei.

P. yonahlossee and P. glutinosus (except for its southeastern

Coastal Plain representatives) attain a larger maximum size than the






- 89 -


other species. P. caddoensis appears to be the smallest Eastern Large

Plethodon. P. J. motcalfi might also be considered a dwarfed form.


Red pigment occurs in adult jacksoni, yonahlossee, ouachitae,

jordani, and shermani. Dorsal guanophores occur in glutinosus, alba-

gula, ouachitae, caddoensis, clensonae, and teyahalee. Lateral guano-

phores are usually present in all Large Eastern Plethodons, except for

four races of jordani (ordani, metcalfi, rielavcntris, and some shemani).


In summary, P. wehrlei seems to be the most distinct of the

Eastern Large Plethodons, while the others seem to be morphologically

very similar to each other, In ry opinion, the hypothetical ancestor

of the group might most reasonably be assumed to have been a moderate-

sized animal with a light chin and dark belly, probably with paired red

spots on the dorsum of the adult, possessing 17 trunk vertebrae, a short

vomerine series, and webbed toes. P. wehrlei is closest to this hypo-

thetical ancestor, although its body has become slightly elongated with

the addition of an extra trunk vertebra. The chin is still light in

all except glutinosus (albagula excepted) and one or two races of jor-

dani. The dark belly has remained in all but the four northern races

of jordani and in southern wehrlei. A larger size has been attained by

yonahlossee and glutinosus, while dwarfing has occurred in northern

jordani (especially metcalfi) and caddoensis. P. yonahlossee has a

much longer vomerine series than the others. The degree of relation-

ship indicated by a study of these characters would seem to indicate a

phylo.-eny as outlined in figure 19.





- 90 -


Plethodon wehrlei Group


Plethodon wehrlei

Plethodon wehrlei inhabits the unglaciated Appalachian Pla-

teaus Province in southwestern New York, western Pennsylvania, extreme

southeastern Ohio, West Virginia, and adjacent Virginia (whore it oc-

curs a short distance outside the Appalachian Plateaus Province).

This species is the most distinct of the Eastern Large Plethodons and

appears to occupy a somewhat isolated position in the group, differing

from the other species in several respects. There is usually more

webbing on the toes of this species, although occasional specimens of

other species, especially P. caddoensis and P. ouachitae, approach P.

wehrlei in this regard. P. wehrlei is the only Eastern Large Plethodon

that ncrmally has 18 trunk vertebrae; all the others usually possess 17.

P. wehrlei possesses fewer vomerine teeth than any other Eastern Large

Plethodon and it is the only species that has melanophore pigmentation

in the peritoneum.


Two close relatives of Plethodon wehrlei from southwestern

Virginia (P. dixi and P. jacksoni) have recently been described as dis-

tinct species by Pope and Fowler (19L9) and Newman (195h). P. dixi

appears to differ from P. wehrlei only in proportions and pigmentation.

It would seem best to regard it as a subspecies of P. wehrloi, since it

replaces the latter geographically. P. jacksoni, based on specimens

from an adjacent county, less than 15 miles from the dixi localities,

essentially differs from P. wehrlei in the retention of the juvenile

dorsal red spots in the adult. Young West Virginia wehrlei often possess

these spots, but they appear to be absent in more northern wehrlei.





- 91 -


Since jacksoni does not otherwise differ from some wehrlei and dixi,

it is also here regarded as a subspecies of P. wehrlei.


Several writers have commented on the geographic variation

in P. wehrlei. Netting (1936: 91) lists ways in which West Virginia

P. wehrlei differ from topotypic Pennsylvania specimens. The former

possess white spotting on the throat and chest, while in Pennsylvania

specimens the white spotting is absent. The lateral white pigment is

also more abundant in West Virginia material.


Dunn (1926: 135) mentions the presence of paired red spots on

the dorsum of a juvenile from Bristol, West Virginia. Brooks (190s:

231) reports that three adults, as well as most of the juveniles, in a

series of 22 specimens from Randolph County, West Virginia, also pos-

sess dorsal red pigment. Bishop (1941: 238) states that none of the

specimens of this species he has examined (presumably all from New York

and Pennsylvania) have shown the slightest trace of red pigment.


Grobman (1944: 287) has pointed out some of the above dif-

ferences between West Virginia and New York and Pennsylvania specimens,

and has also suggested that southern wehrlei may attain a greater size

than northern specimens. In view of these differences in northern and

southern P. w. wehrlei, this form should be further studied for other

evidences of geographic variation. It is possible that the northern

and southern populations should be recognized as distinct subspecies.

In the possession of reduced black pigmentation on the anterior portion

of the belly, southern wehrlei are more similar to dixi and jacksoni

than are northern wehrlei. In the presence of red spots on juveniles,

as well as in some adults, West Virginia specimens are similar to






- 92 -


jacksoni. The increased lateral white pigmentation in southern wehrlei

is also paralleled by jacksoni and dixi.


I have not carefully examined the pigmentation of any of the

races of wehrlei in life, although a casual examination of living speci-

mons from New York, West Virginia, and Virginia (dixi topotypes) was

made before the present study was contemplated. A comparison of the

geographic variation in pigmentation characters within the species, as

well as with other Plethodons is much to be desired. Pope and Fowler

(1949: 1) state that both "bronzy mottling" and ?small light flecks"

are present on the back of dixi. The former disappear rapidly in pre-

servative whereas the latter remain, although fading somewhat. Pope

and Fowler state that they have occasionally observed the "white fleck-

ing" in wehrlei, but never the "bronzy mottling." The white flecking is

probably the same type of spot evident on the specimen figured by Bishop

(194la: fig. 45). The red dorsal spots, present in young West Virginia

wehrlei, were not observed by Pope and Fowler in a large series of para-

typic dixi, including 59 juveniles. The belly of dixi is mottled (pre-

sumably with white guanophores on a melanophore background), whereas

the belly of wehrlei is usually uniformly pigmented with melanophores.

In size, dixi appears to be smaller than the other races. The speci-

mens of dixi that have been examined also appear to have proportionately

narrower heads and slenderer bodies than wehrlei.


Newman (1954: 12) states that the dorsum of jacksoni has "white

flocks" and "silvery mottling," the latter usually disappearing within

twenty-four hours after preservation. I.ottling on the belly is also

present, as in dixi. According to Newman, the dark belly pigmentation




Full Text

PAGE 1

THE RELATIONSHIPS OF THE SALAMANDERS OF THE GENUS PLETHODON By RICHARD HIGHTON A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA June, 1956

PAGE 2

iifipiili J 1262 08552 2885

PAGE 3

tablh; op contents Page INTHOOliCTION . 1 MRHDB0 5> THE RELATION BETWEEN NUMBER OF COSTAL GROOVES AND TRUNK VEitTEBRAE IN PLETHODON 8 PIGMENTATION IN THE GENUS PLETHODON 12 TOO. TO THE SAIAtANDl&b OF 98 GENUS PLBTHODOM lU STSTSttTICS 20 The Western Plethodons 29 Plethodon vandykei Group . . 3h Plethodon vandykei vandykei Van Denburgh .... 35 Plethodon vandykei idahoensis Slater and Slipp . . 37 Flethodon vandykei larselli Burns . . 37 Plethodon vehiculum Group 38 Flethodon d unni Bishop ....... 38 Plethodon vehiculum (Cooper) Ill Plethoaon productus Croup ,. k3 Plethodon productus , new nans 1*3 Plethodon neomexicanus Group • U6 Plethodon neowexican us Stebbins and Riemer .... U6 The Eastern Small Plethodons U7 Plethodon wclleri Group 5l Flethodon welleri Valker 5l Plethodon richraondi r ichmondi Netting and Mittleran 61 Plethodon richraondi popei 1 ighton and Grobman . . 62 Plethodon richmondi netting! Green .. 6U

PAGE 4

ii Plethodon cinereus Group . 65 Plethodon dors alls dorsalla Cope ........ 68 Plethodon dorsalis angusticlavius Grobman .... 71 Plethodon cinereus cinereus (Green) . 76 Plethodon cinereus serratug Grohman 78 Plethodon cinereus polycentratus Highton and Grobman 79 The Eastern Large Plethodons 80 Plethodon wehrlei Group 90 Plethodon wehrlei welirlei Fowler and Dunn .... 93 Plethodon wehrlei dixl Pope and Fowler ..... 96 Plethodon wehrlei jacksoni Newman ........ 96 Plethodon yonahlossee Group 97 Plethodon yonahlossee Dunn ........... 100 Plethodon ouachitae Dunn and Keinse ....... 103 Plethodon caddoensis Pope and Pope ....... 105 Plethodon glutinosus Group . 106 Plethodon jordani jordani Blatchley 129 Plethodon jordani metcalfj Brimley 132 Plethodon jordani shermani Stejneger 13k Plethodon jordani unicoi, new subspecies .... 135 Plethodon jordani raelaventris Pope and Hairston . 136 Plethodon jordani rabunensis Pope and Hairston . 138 Plethodon jordani teyahalee Hairston 1U0 Plethodon jordani clamsonae Brimley lUl Plethodon glutinosus glutinosus (Green) 193 Plethodon glutinosus albagula Grobman . 196 SUM3ART 197 LITERATURE CITED 198

PAGE 5

iii LIST OF FIGURES Figure Page 1. Lateral and dorsal views of the trunk vertebrae of representatives of the three major groups of the genus Plethodon 23 2. Suggested phylogeny of the major subdivisions of the genus Plethodon , 26 3. Suggested phylogeny of the Western Plethodons 33 U, The distribution of the subspecies of Plethodon vandykei in Washington, Oregon, and Idaho. . 36 5. The distribution of Plethodon dunni in Oregon and southwestern Washington, . . Uo 6. The distribution of Plethodon vehiculum in Oregon, Washington, and British Columbia . U2 7. The distribution of Plethodon productus in California and Oregon US 8. The distribution of Plethodon neomexicanus in New Mexico h.5 9. Suggested phylogeny of the Eastern Small Plethodons ... 5>2 10. The distribution of Plethodon welleri in North Carolina, Tennessee, and Virginia. 52 11. The distribution of the subspecies of Plethodon richmondi 63 12. The distribution of the subspecies of Plethodon dorsalis . ....... 70 13. The distribution of the subspecies of Plethodon cinereus 77 Hi. Vomerine teeth of ?6° peninsula Florida P. glutinosus plotted against snoutvent length. .7 83 15. Vomerine teeth of \& P. glutinosus from the Coastal Plain of Virginia and North Carolina plotted against snoutvent length. ... ... 83 16. Vomerine teeth of ll£ P. j. jordani plotted against snout-vent length. . ."" 8U

PAGE 6

It Figure Page 17. Vomerine teeth of 60 P. w. wehrlei plotted against snout-vent length 8ii 18. Vomerine teeth of 39 P. caddoensis plotted against snout-vent length 85> 19. Suggested phylogeny of the Eastern Large Plethodons ... 85 20. The distribution of the s\ibspecies of Plethodon v/ehrlei . 9$ 21. The distribution of the Plethodon yonahlossee Group . . . 102 22. The distribution of the subspecies of Plethodon jordani . 112 23. The distribution of the dark-chinned ( glutinosus ) and light-chinned ( albagula ) populations of P. glutinosus in central Texas . . 189 2U. The distribution of Plethodon glutinosus in eastern United States 195

PAGE 7

INTRODUCTION The North American renus Flethodon presently has more recognised forms than any other genus of salamanders. It is the type genus of the lungless family Plethodontidae, reviewed so ably by Dunn in 1926. At present, this is the most successful salamander family; over half of the living species of the Order Caudata are plethodontids . North America is the center of the distribution of the family, but one genus, Hydromantes , also occurs in Europe, and members of several genera enter the Neotropical region. Salamanders of this family occupy habitats ranging from strictly aquatic cavernicoles ( lyphlomolge and haideotriton ) and mountain stream dwellers ( Leurognathus ) to others like i'lathodon , which are completely terrestrial, even to the ex-bent of laying their eggs on land. Dunn believed that Plethodon is the most primitive genus in the attached-tongue branch of the family, and that Hemidactyiium , fesatina , Batrachoseps , and Aneides are more specialised deriv?tives of a Plethodonlike ancestor. In 1926, only eleven forms of Plethodon were known to Dunn. In 19U3> Bishop listed 17 species (one has since been removed from the genus) and two subspecies (both are now considered different species). In 19UU, Grobman reviewed the distribution and relationships of the eastern section of the genus, including some sixteen species and subspecies. There are now twentyfive forms recognized in eastern North America, with seven more in the western United States and Canada. Grobman (19U1a* 266) divided the eastern forms into two groups, the Large Plethodons and the Small Plethodons. He suggested that these

PAGE 8

2 groups might actually represent distinct renera or subgenera, but he reserved Judgement in this matter until the relationship of the western species with the eastern forms could be determined. One of the purposes of the present study is to determine these relationships by a comparative study of the morphology of all the species of the genus Plethodon. Of the twenty-five genera in the family Plethodontidae, only two others show as great or greater disjunctions in their distributions as that found in Plethodon , These are hydromantes , with two forms in Europe and three species in California, and Aneides , with four species in western North America and one in the Appalachian Mountains of the eastern United States. In the past two decades, several new plethodons have been discovered in both eastern and western North America. A review of the relationships of these forms offers a valuable opportunity to study the evolution of this important group of North American salamanders. The importance of exa'iining plethodons in life has been emphasised by most recent workers, "wany species, particularly the Large Eastern forms, are extremely difficult to identify after years of preservation. In few other vertebrates are differential characters between species so rare. For this reason, a special attempt was made to obtain living specimens of each form. Twenty-five of the thirty-two forms recognized herein have been studied nlive. The list of persons who have contributed specimens to this project is long and I wish to express to each of them my sincerest thanks and appreciation for their valuable aid: Charles G. Adams,

PAGE 9

3 Ifclter Auffenberg, Herbert Barden, Albert P. Blair, E. E. Brown, Jerry and Esther Brown, Douglas U. Burns, Archie Carr, A. H. Chaney, Roger Conant, John '.V. Crenshaw, J. C. Dickinson, John Dolan, llerndon Dowling, Philip C. Dumas, Helen T. Gaige, M. Ruth Gilreath, Arnold B. Grobraan, T. P. Haines, Keith L. Hansen, Robert Hellraan, Albert H. nighton, Thelraa Howell, Leslie tfubricht, Robert Humphries, Richard M. Johnson, James Kezer, J. D. Kilby, James Knepton, H. D. Leake, Edwin H. McConkey, John S. Mecham, Sherman A. i&nton, .ilfred T. Heill, Howard T. Odum, Larry H. Ogren, David Pettus, John Quinby, George Rabb, Neil D. Richmond, .William Riemer, Bernard Roseman, Doug Rossrnan, Albert Schwartz, ilcbert S. Simmons, Ralph Sinclair, William Sloan, Peter Smith, Bette Starner, Charles J. Stine, Valter Stone, Virginia Switzer, Sam R. Telford, G. M. Thorp, Gordon Thurow, Don Tinkle, Arnold and Rusty Van Pelt, Charles P. talker, "illiam 'itt, and harry Yeatman, Most of the material given me by these persons has been deposited in the University of Florida collection. This collection has formed the nucleus of the material on which this study is based. Some additional specimens have been borrowed from other collections, but I did not attempt to follow the course of the conventional monographer and examine every specimen available in museums, simply for the sake of completeness. The material borrowed from other museums includes the following: Western Plethodons fron the University of California j Texas P. glutinosus from Bryce C. Brown, Ottys Sanders, John S. Mecham, and the Strecker lluseumj P. d. angusticlavius , F. glutinosus , P. ouachitae , and P. caddoensis from the University of Arkansas; P. c. serratus , P. w. wehrlei , and P. w. dixi from the

PAGE 10

-uChicago Natural History Museums P. glutinosus, P. w. wehrlei , and P. richmondi from the Carnegie Ifuseura; P. cinereus from the ?Juseura of Comparative Zoology; P. richmondi from the United States National Museum; P. glutinosus and P. ^. rabunensis from the Charleston J'useum; P. dorsalis from the University of Georgia; P. richmondi and P. glutinosus from the Cincinnati Society of Natural History; P. dorsalis from the Ross Allenilfred T. Neill collection; and the entire Plethodon collection of the Great Smoky '.fountains National Park. For the loan of material in their care, I wish to thank the following museum officials: Neil D. I&chraond, Carnegie Museum (CM); Albert Schwartz, Charleston Museum (ChM); Robert Inger, Chicago Natural History Museum (CNHM); Ralph Dury, Cincinnati Society of Natural History (CSNH); Arthur Stupka, Great Smoky Mountains National Park (GSHNP); Arthur Loveridge, Museum of Comparative Zoology (MCZ); .ilfred T. Neill, Ross Allen Reptile Institute ( ,r .aA-V,TN); Bryce C. Brown, Strecker !?useun (SM); Herndon Bowling, University of Arkansas (UA); Bernard Martof, University of Georgia (UG); William tLemer, University of Florida (UF); Charles F. Walker, University of Michigan, Museum of Zoology (UMMZ); and Doris M. Cochran, United States National Museum (U3NM). The following individuals have generously let me borrow material in their private collections: Bryce C. Brown (BCB); John S. Mecham (JSM); and Ottys Sanders (OS). In addition I have greatly benefitted from stimulating discussion of some of the problems which have arisen during the course of this study with the following persons: Drs. Robert Bader, Pierce Brodkorb, Archie Carr, John W. Crenshaw, J. C. Dickinson, Richard A.

PAGE 11

-5Edwards, Coleman J. Goin, Howard T. Odum, villiam Riemer, H. K. Wallace, and Messrs. 'alter Auffenberg, Robert Hellman, Sichard M. Johnson, and . ilf red T. Neill. itywife, Anne, besides being patient and sympathetic while undergoing considerable personal inconvenience during many long hours of field work in the various hot, wet, cold, and mosquito-infested places where these animals live, has also materially contributed to the project by typing, filing, proofreading, and especially by taking practically all the color notes on geographic variation of the P. glutinosus group. The Chairman of my Graduate Supervisory Conmittee, Dr. Arnold B. Grobmnn, has been a constant source of intellectual stimulation, encouragement and helpful advice and criticism. METHODS The importance of studying living plethodons, because of the usual loss and alteration of their pigments in preservatives, has already been mentioned. It is almost impossible to study geographic variation in pigmentation without considerable field work, and it was initially hoped that this could be done with all the forms in the genus. Unfortunately, several of the forms have not been seen alive and so our knowledge of their geographic variation has not been augmented. However, I have been able to do a considerable amount of field work with some of the Eastern Large Plethodons, and the result has been a considerable amount of new information concerning them. I believe that when similar studies of other species are made, they will be found to vary as much as glutinosus and jordani in pigmentation characters .

PAGE 12

6 A perusal of the literature on this genus will disclose that most workers have given considerable weight to variation in the number of costal grooves. A careful study of the literature indicates, however, that there is not agreement in the reports of number of costal grooves in many species. The reason for this was apparent to the writer when I failed to obtain consistent counts on the same specimens counted at different times. A method for accurately determining the true variation in body segmentation would be a necessary preliminary to the use of this character in studying relationships. Since the costal grooves show a direct correlation with the number of vertebrae, a method of counting the number of trunk vertebrae by the use of X-ray photographs was devised. As an outgrowth of this study, a method for counting costal grooves which will accurately reflect the body segmentation of salamanders of this genus will be reported below. I'-any other characters have been used in the study of the systematics of this genus. As far as possible, these have been restudied using much new material that has recently become available. Several nomenclatorial changes will be suggested based on an attempt to best indicate the biological relationships of these organisms. The results of this study have clearly indicated that there are three major natural subdivisions of the genus Plethodon . These would probably be considered subgenera in most animal groups, but herpetologists have rarely used subgenera in their classification. Although the use of subgenera would be helpful in this case, common names will be used for these groups in conformity with current herpetological practices. They will be referred to as the "Vestern Pleth-

PAGE 13

7 odons," "Eastern Small Flethodons," and "Eastern Large Plethodons." Each of these subdivisions of the genus is further divided into species groups consisting of species which are more closely related to each other than they are to any other similar group. In a few cases, species which are morphologically quite distinct from any other species in their section of the genus have been segregated into a species group consisting of a single species. For each taxon, information on the morphology, variation, and distribution is given, followed by a synonomy, data on the type specimen, diagnosis of the form, and a description of the pigmentation, segmentation, number of vomerine teeth, and size. Detailed descriptions of several hundred specimens were made during the course of this study. For most forms, similar data are available in Bishop (19U3) or Stebbins (195D or in the original description. It is unnessary to repeat these here, since the concern is variation of populations, not detailed descriptions of individual specimens. Dunn (1926) presents fairly complete synonomies for each of the forms known at that time. Therefore the large number of references on the distribution and habits of the more common forms are not included here. An attempt has been made to include the first reference to each combination of names, as well as most of the papers dealing with the systematics of each form. Check lists have usually been omitted, except when new combinations of names or other new information is presented. Consistency has not been a goal, and some papers dealing with habits or distribution have been included in the synonomies of little known species, while similar papers on well known forms

PAGE 14

8 that do not contribute new information have been omitted. The locality map3 have been prepared mainly as an aid to the discussion of the distribution of the various species groups. They are based on literature records as well as specimens examined by the writer. Maps shoving the distribution of many of these forms are available elsewhere, but there have been many changes in the taxonomy since the publication of some of these, and it seems desirable to include them here in spite of the fact that there is some repetition. New information on the geographic distribution of several of the forms is included on some of the maps. Seventeen subspecies are recognized in this paper. A n examination of the distribution maps indicates that several of the polytypic species exhibit terminal raciation and most of these races are isolated from their nearest relatives by areas of uninhabited country. This is true of the races of P. vandykei , P. dorsalis, P. cinereus , and to some extent, P. glutlnosus . The subspecies of P. wehrlei are poorly defined, and need more study. There do not appear to be natural barriers between some of the subspecies of P. richmondi and P. jordani , and there is evidence that intermediate populations are present between some of the races. A more detailed consideration of raciation in both of these species is given below. THE RELATION BETWEEN NUMBER OF COSTAL GROOVES AND TRUNK VERTEBRAE IN PLETHODO N Radiographs of series of specimens of all the species in this genus have indicated that the number of trunk vertebrae in a

PAGE 15

9 single population is remarkably constant. (Since there is little differentiation of the salamander body vertebrae into regions, Francis (193U) suggests that all precaudal vertebrae, except the atlas and the sacrum, be called trunk vertebrae .) There is never a variation of more than three trunk vertebrae In a single population and with few exceptions, the intermediate figure has a very high frequency of occurrence. The value of this character in studying variation becomes apparent when it is found that a difference of a single vertebra between two populations is easily detectable and has been used successfully as a key character to separate a high proportion of specimens of two contrasted forms. a method of counting costal grooves that would accurately reflect the number of trunk vertebrae would be much more effective in the taxonoTdc study of this genus. At present, their use is rather limited, since they are rarely counted consistantly by different workers. Pope (19^0: 102) correctly summarizes the existing situation by stating that "variation of a magnitude of one or two grooves will be meaningless unless the same worker has made all the counts and done so *ith great care." The method described here has been found to correctly correspond with the number of trunk vertebrae with over 90i accuracy. The atlas is not related to any costal grooves. The first trunk vertebra is located just anterior to the gular fold and its rib does not appear to be associated with the gular fold or with any of the costal grooves. The rib of the second trunk vertebra is located in the myoseptum of the first costal groove. The external position of this groove varies somewhat, but is usually close to the front

PAGE 16

10 limb insertion. If the first costal groove is poorly defined or absent, as often occurs when it is located directly over the front limb insertion, it should be counted, as the space obviously corresponds to a vertebra whether the groove is visible or not. The individual variation in the position of the first costal groove is probably due to the variation in the position of the pectoral girdle relative to the vertebrae. The girdle may be located opposite the second trunk vertebra, between the second and third trunk vertebrae, or opposite the third trunk vertebra. The ribs of the third trunk vertebra extend posteriorly so that the second costal groove is always posterior to the front limbs. There is a one-to-one relation between the remaining tody vertebrae and the costal grooves, except in the case of rare aberrations (one vertebra possessing two or more pairs of ribs, or one vertebra possessing a rib on only one side, in which case a corresponding costal groove is present only on that side). The last pre-sacral vertebra (usually the only trunk vertebra which does not possess ribs) is represented by the groove just anterior to the insertion of the hind limbs. Often this groove joins ventrally the one just preceeding it so that on the lower sides there is only onegroove which is forked dorsally. Both grooves should be counted, since they correspond to two separate vertebrae. The groove present over the hind limb insertion (sometimes weaklydeveloped or absent) corresponds to the sacral vertebra, and each groove on the tail corresponds to a caudal vertebra. There is much more individual variation in the position of the sacral vertebra in relation to the pelvic girdle than there is in the relation of the shoulder girdle to the second and third trunk vertebrae.

PAGE 17

11 The sacral vertebra may be located so far anterior to the pelvic girdle that the first caudal vertebra is opposite the hind limbs. It may also be slightly anterior, opposite, or slightly posterior to the pelvic girdle. Occasionally when it is posterior to the girdle, the costal groove corresponding to the last trunk vertebra is located over the hind limb. This is the major source of error in attempting to accurately correlate the number of costal grooves with the number of trunk vertebrae. Fortunately, however, fewer than 10;? of the specimens examined were in this category. Using this method, costal groove counts were made on 8£> specimens of several species of the genus Flethodon (including specimens possessing 1$ to 22 trunk vertebrae), and 93% of the counts accurately represented the number of trunk vertebrae obtained from X-ray photographs. Since the first trunk vertebra is not associated with a costal groove, the number of costal grooves is always one less than the number of trunk vertebrae. It would therefore appear that this method can be used advantageously to obtain an accurate estimate of the number of trunk vertebrae of all the species of the genus Hethodon . Occasionally the sacral rib may attach to one vertebra on one side of the animal and to the following vertebra on the other side. Rarely, there are two sacral ribs issuing from two successive vertebrae on the same side of the animal. The suggested method of counting costal grooves is of no value in detecting such abnormalities.

PAGE 18

12 PIGMENTATION IN THE GENUS PLEIHODQN Pigmentation in living specimens was studied with the aid of a dissecting microscope. Although there is great variety in the coloration of the animals included in this genus, the actual different types of pigments are few. No histological or biochemical studies of the pigments have been made, thus similarity in the appearance of the pigmentation is the basis for the above statement. Three main types were identified and are called melanophores, guanophores, and lipophores, following Ctebbins (1951). Melanophores are present in all the forms in the genus. They are responsible for the dark ground color of these salamanders. The other pigments generally occur in gaps in the raelanophore ground color. Guanophores are responsible for the white and brassy spots, present on many of the species. There appears to be little difference in the structure of the different colored guanophores, but mainly a difference in the amount and color of light reflected from the spots. The iridescence characteristic of animals with "brassy" "dorsal spots appears to be centered in small round crystalline structures found at intervals along the pseudopodia of the guanophores. The number of these crystals seeiis to determine the amount of the iridescence that has been variously described by different workers as "brassy flecking," metallic golden spotting," "golden blotches," "bronsy mottling," and "frosting." This type of guanophore pigmentation is present in the iris of most of the Eastern Small Plethodons and the >'/estern Flethodons, and is present on the dorsum of many of the species. It is responsible for the brassy dorsal flecks of glutinosua , ouachitae , dixl , clemsonae , popei , netting!,

PAGE 19

13 cinoreus, poly cent ratus , d orsalis , welle ri, and vehiculum . In welleri , these guanophores are concentrated to form dense clusters. In glutinosus, they are usually associated with other white guanophores. In the other forms they are scattered over the back and are not clumped into spots. Brassy flecks are rarely found other than in the iris or on the dorsum of salamanders of this genus. Guanophores that lack the brassy iridescence are common on the belly and sides of many species, and are also present on the dorsum, occasionally occurring there with the brassy type. They have a much g renter tendency to be clumped together to form larger spots than do the brassy flecks, but are occasionally found singly. In glutinosus , for example, it is often possible to see, on the same animal, every type of intermediate between the brassy type and those which lack the brassy iridescence. The white guanophores are characteristically found on the ! orsum of glutinosus, ouachitae , cinereus , polycentratus , popei , dorsalis , and vehiculura. They are present on the sides of almost all of the eastern species (except Tietcalfi -ind melaventris ) , and on the bellies of all the Eastern Small "lethodons. Often these lateral and ventral spots have a yellowish color, but this ±b not due to the presence of brassy flecks. Lipophores do not have the structure of guanophores in that they lack the pseudopodia which can easily be observed in the guanophores. The red color of jordani , shernani , wehrlei , yonahlossee , ouachitae , cinereus , polycentmtus , serratus , dorsalis , and vehiculum

PAGE 20

-Ikappears to be duo to the same type of lipophore pigment. The dorsal band of yonahlossee is a darker color because of the additional presence of raelanophore pigment. Lipophores may also be yellow in color, as in some cinereus, vehiculum, and dunni . The variety of colors present in the genus seems to be due entirely to variation in the abundance of these pigments or various combinations of the three. In some forms, the lipophores or guanophores or both are lacking. There is also variation in the concentration of melanophores . These pigmentation characters may vary somewhat within a species, both individually and geographically, but are fairly uniform in most forms, enabling a person fa Miliar with living specimens to easily identify most salamanders by the color pattern alone. The phylogenetic significance of the distribution of these pigments in the various plethodons is discussed below in the accounts of the various forms. KEI TO THE SALAMANDERS OF THE GENUS PLETHODON The key is based mainly on the average number of trunk vertebrae occurring in each of the forms. Before using this key, it is suggested that the section on the method of counting costal grooves that accurately reflects the number of trunk vertebrae be road. A small percentage of specimens of each form may not be correctly identified on this basis, but a small series will usually key out correctly, l&nges are included and locality data may proove more helpful in identifying preserved specimens than pigmentation characters. 1 a Costal grooves usually lli 2 b Costal grooves 13> or more h

PAGE 21

-152 a Ventral color reddish (IJultnomah County, Oregon, and Skamania County, .iashington) £• 2larseI ^ b Ventral color not reddish 3 a ' Yellow or orange dorsal stripe contrasts sharply with the lateral black ground color; the proximal segments of the limbs dark in color (Kootenai County, Idaho) P. t. idahoensis b Ground color light, not contrasting sharply with the dorsal stripe; yellow lipophore pigment similar to that found on the dorsum present on the proximal segment of the limbs (western Washington) £• J« vandy^ei U a Costal grooves usually l£ (western Oregon and southwestern Washington) Z* ^£2* b Costal grooves 16 or more £ a Costal grooves usually 16 b Costal grooves 17 or more 6 a Belly mottled with yellow or red, white and black; size small, usually under £0 mm. snoutvent length; often a red, tan, or yellow dorsal stripe (southwestern British Columbia, including Vancouver Island, western Washington and western Oregon) £• v*l«fl»! b Belly usually dark, at least posteriorly, or with scattered small white ventral spots; size usually larger (except welleri and caddoensls ); dorsal stripe usually absent (except yonahlossee and ouachitae ) (eastern United States) 7 7 a Red pigment present on back, legs, or rfieeks 8 b No red pigment present on animal u

PAGE 22

168 a Red pigment confined to sides of head or legs in adults . 9 b Red pigment largely restricted to dorsum 10 9 a Red pigment most abundant on sides of head (Great Smoky Mountains of Tennessee and North Carolina) . . P. jj. jordani b Red pigment most abundant on legs (Nantahala (fountains, North Carolina) Z* A* shermani 10 a White pigment lacking in dorsal stripe (Blue Ridge Mountains of southwestern Virginia, northeastern Tennessee, and northwestern North Carolina) P. yonahlossee b Abundant white pigment occurring within the dorsal stripe (Ouachita /mountains of Arkansas and Oklahoma) . P. ouachitae 11 a Sise small, adults not over !>0 nm. snoutvent length; back with large coalescing iridescent brassy spots, usually covering about half the area of the dorsum (Blue Ridge Province of southwestern Virginia, northeastern Tennessee and northwestern North Carolina) P. welleri b Back without dorsal brassy spots, or if present, they are small in size and cover leas than one quarter of the area of the dorsum 12 12 a Body entirely black, guanophores and lipophores absent . . 13 b Dorsum and/or sides with guanophore spotting lU 13 a Belly much lighter than dorsum (mountains of western North Carolina and adjacent Tennessee and Virginia) P. J. me teal fi b Belly almost as dark as dorsum (southwestern North Carolina) ....... P. ^. melaventris

PAGE 23

17 lU a Back with large conspicuous white or brassy dorsal spots . 1$ b Back without large conspicuous guanophore spotting, or if present, the spots are of very small size 17 13> a Melanophore pigmentation on chin greatly reduced, compared to belly 16 b Melanophore pigmentation on chin similar to that on belly (eastern United States from southern New York to northern Florida, west to eastern Louisiana, Illinois, Missouri, eastern Oklahoma, and the Balcones Escarpment in Texas) P. g. t;lutino3tts 16 a Dorsum with white spots only (Balcones Escarpment in east central Texas) P. £. albagula b Dorsum with large white spots and smaller brassy flecks (Caddo Mountains of Arkansas) ......... P. caddoensis 17 a Dorsum with very small white or brassy spots 18 b Dorsum black, without any guanophore spotting . 19 18 a Dorsum covered with tiny brassy flecks (vicinity of Jocassee, South Carolina) . P. i« clemsonae b Dorsum with very tiny scattered white spots (Tusquitee and Snowbird Mountains of western North Carolina) P. jj. teyahalee 19 a Belly much lighter than ground color of back (Unicoi Mountains of western North Carolina ami eastern Tennessee) £*i* uni-coi b Belly nearly as dark as dorsal ground color (mountains of northeastern Ceorgia £• i* rabunensis

PAGE 24

18 20 a Costal grooves usually 17 21 b Costal grooves 18 or more .. 23 21 a Large red spots on dorsum of adult (southwestern Virginia) P. w. jacks oni b Red spots absent fron dorsum of adult 22 22 a Dorsum with numerous white and brassy spots (Roanoke County, Virginia) £• 2« dixi b Dorsum with very small white or brassy spots (Cattaraugus County, Hew York, south through western Pennsylvania, adjacent Ohio, and West Virginia) P. w. wehrlei 23 a Costal grooves usually 18 2k b Costal grooves 19 or more 27 2U a Size large, up to 66 mm. snoutvent length (northwestern California and southwestern Oregon) P. productus b Size small, up to \\% ram. in snout-vent length (eastern United States) 25 ?$ a Belly mottled with red, yellow, white, and black pigment . 26 b Belly black, with small white spots (Cheat Mountains of West Virginia) p. r. netting! ?6 a Dorsal stripe very narrow, less than 1/3 the width of the body (southwestern Missouri, northwestern Arkansas, and adjacent Oklahoma) P.. d. angusticlavius b Dorsal stripe much wider than 1/2 the width of the body, edges of stripe very irregular (southern Illinois, Indiana, and southern Ohio, south through Kentucky and Tennessee to northern Alabama and northwestern Georgia 1 ! . P. d. dorsalis

PAGE 25

19 27 a Costcl grooves usually 19 28 b Costal grooves 20 or more 30 28 a Fifth t. c on rind foot usually v&th one phalanx (Jemez f New exico) P. neomexicanus b Fifth toe on hind font with two phalanges (eastern North 29 a Dorsal red stripe with straight edges (southern Canada, south to North Carolina, eastern Kentucky, southern Illinois, and eastern issouri) P. c. cinereus b Dorsal red stripe with serrations corresponding to each costal groove (western Arkansas and eastern Oklahoma) P. c. serratus 30 a Bed pirment present on rides and dorsum (northwestern Georgia. P. c. p olycentratus b Ho red pigment on animal 31 31 a Costal grooves usually 21 or more (western Pennsylvania, southern Ohio, eastern Kentucky, south to northwestern Virginia) P. r. richmondi b Costal grooves usually 20 (southwestern Virginia, northwestern North Carolina, and adjacent Kentucky) . P. r. popei

PAGE 26

20 SYSTEMATICA Flethodon Tschudi Plethodon Tschudi (1838* 58). Genotype (by original designation): Sala^ndra glutinosa Green. Phatnomatorhina Bibron in Bona] -arte (1839). (Substitute name) Sauropsis Fitzinger (I8h3: 33). (non Sauropsis Agassis, 1832, Jahrb. Ittn., 3 ! 1^2). Genotype (by original designation): Salamandra erythrono ta Green. Diagnosis ;Plethodontidae with tongue attached in front; premaxillae separate? teeth on posterior portion of maxillae; tail not constricted at base; five toes on the hind feet; no palmar tubercles; terminal phalanges normal; and no aquatic larval stage. According to Dunn (19?6), the closest relatives of Plethodon are Batrachoseps , Ensatina , Aneides , and r.emidactyliura . All four of these genera differ from Plethodon in several fundemental characteristics. I^atrachoseps and H emidactylium possess only four toes on the hind feet. Knsatina and Hemidactylium possess a basal constriction of the tail. Ensatina has palmar tubercles. The premaxillae are fused in Aneides and Batrachoseps (except B« wri^hti ) . Hemidactylium has an aquatic larval stage. Aneides lacks teeth on the posterior portion of the maxilla and has expanded terminal phalanges. Dunn (1926: 22) believed that Plethodon is the most primitive genus in this group of genera and that the others, with the possible exception of Kn sat in a, are more specialized than Plethodon . The genus Plethodon is the largest in number of species, and there is more divergence v.ithin the genus than in the other related genera. This might be considered nvidence for, but not necessarily proof of, a greater age for this genus.

PAGE 27

21 Qrobman (l°UUs 66) suggests that the relationship between the Eastern id Small Plethodons is not close and lists size and costal groove differences which distinguish them. Actually there is some over! en tie Eastern Large and Small Plethodons in these characters, tut there are other important differences between the two groups. to mental gland is much better developed in male Eastern Large Plethodons than in the -..astern Small Plethodons. The worm-like body and shorter lo,:s of the Eastern Small Plethodons is characteristic. The Easte-ohodons have fewer teeth and there is a great deal of variation in the number of trunk vertebrae within the group (range 16-2U), kereas the .astern Large "lethodons she./ very little variation (range 16-19). There is more v/ebbing on the toes of the small forms than in the large species. The Eastern Large 1 lethodons have an unpigmented parietal oritoneum, vjhile in the s^ali species the peritoneum is pigmented ith melanophores. The differences in the structure of the vertebrae are discussed below. One species, P. wehrlei , is usually included with the Eastern Large Methodon? on the basis of size, but in several ways it is intermediate between the two groups. It has more costal grooves and fewer tepth than other i>astern Large Plethodons. Its toes are vebbed as in the E astern Small Plethodons and the peritoneum has a few melanophores. ere it not for the intermediate character of this species, the two groups should probably be recognized as distinct genera, but the somewhat inter a ±L ate nature of wehrlei would seem to indicate that they have not tocome sufficiently distinct to justify such an action. The two grouor: could best be regarded as subgenera, but to conform with current . :ctise, the common names, Eastern Large Plethodons and Eastern

PAGE 28

22 Small Plethodons, will be used in this paper. Although P. wchrlei possesses more characters that would link it with the Eastern Small Plethodons than do any of the other Eastern Large Plethodons, it is probably more closely related to members of the latter group. Therefore, it is included as a separate species group under the Eastern Large Plethodons. Its intermediate position is important, however, in linking the two groups. During studies on the osteology of this genus, certain differences in the vertebrae of the two eastern sections of the genus have been noted. The height of the vertebrae of the Eastern Small Plethodons is proportionately less and the vertebrae usually lack the neural spines that are present on those of the large eastern species (see figure 1). P. wehrlei is not intermediate in this regard, but closely resembles the other Eastern Large Plethodons. The vertebrae of all the eastern species have been examined except for caddoensis and richmondi . To investigate the relationship of the western forms with those in eastern North America, each of the above differential characters was studied in all five of the western species. As with the eastern forms, the size varies within the group, but four of the five western species are as large as most of the Eastern Large Plethodons. The number of trunk vertebrae varies considerably in the western forms (range 15-20). The mental gland is poorly developed in all the Western Plethodons. The body form is variable, ranging from short and stout in vandykel to very elongate in neomexicanus . The number of vomerine teeth is low in the western forms. The toes of two species (dunni and vehi-

PAGE 29

23 Figure 1. Literal and dorsal views of the trunk vertebrae of representatives of the three major groups of the genus Plethodon. A. P. glutino sus . B. P. welleri C. P. dunni

PAGE 30

2UcuLum) are ver Bltj btly nebbed, while the other ohree species have webbed toes. -itoneum of all of the western Plethodons is pigmented with lei nophorea, thm vertebrae of the western forms are quite different from both eastern types (P. neomexicanus has not been examined), are proportionately longer and have longer transverse processes than any of the eastern species. As a group, the riestem Plethodons are more variable, and in at least on<* character, the structure of the vertebrne, they are very distinct from both eastern sections of the l. It would appear that tho Western and Eastern Plethodons have benn separated for a long time, anu that both have undergone considerable speciation during this period. In several ways, the Kaetern Small Plethodons resemble the Western Plethodons nore closely than do the eastern i^arge Plethodons. The red or yellow dorsal stripe is more common in both groups than it is in the rtnstern Large Plethodons. Both have a great amount of variation in the number of trunk vertebrae an i degree of elongation of the body, both have low vomerine teeth counts and a pigmented peritoneum. Both hove a less well developed mental gland than the Kastern Large lothodons (except for dorsalis). It v ould appear that the Basterr ?thodons are the more specialized forms, with a larger number of vomerine teeth, loss of pigmentation in the peritoneum, and increase in the -evelopment of the mental gland. The fact that P. wehrlei possesses characteristics of both groups would indicate that both were derived from a common ancestor. This common ancestor was prob-bly '.ore like the Eastern Small Plethodons and the Western Plethodons in ost of its characteristics, since it would be unlikely that these two groups would have independently converged toward each other

PAGE 31

.25in so many ways from an ancestor that was similar to the Eastern Large Plethodons aa suggested by Dunn. The relationships of the three groups would appear to be best indicated by a phylogeny similar to that shown in figure 2. The relationship of the Western Plethodons with the other plethodontid genera in western North America needs further study. It may be that the Western Plethodons are more closely related to Batrachosepjs, Ensa tina, or western A neides than they are to any of the Eastern Plethodons. The western plethodontid salamanders may have been isolated from their eastern relatives for a long period of time. The fact that two groups now included in the genus Plethodon , one in eastem North America and the other in the western part of the continent, have both retained many primitive characteristics, does not necessarily mean that they are still genetically related to one another. The fact that each of these groups (the Western and Eastern Plethodons) has more species than any other plethodontid genus in its region, as well as the the fi tct that these species are so diverse, would suppot the view that they have been separated for a long period of time. A review of this entire problem is needed, but it would involve an investigation of the characteristics of the genera Aneides , Batrachoseps , Ensatina , and liemidactylium , and is beyond the scope of this study. Such an inquiry should be completed before erecting a new genus for the Western Plethodons , but this arrangement seems to be indicated by the present incomplete information. The systematic arrangement of the genus Plethodon suggested in this paper is as follow3t

PAGE 32

26 Eastern Large Plethodons Eastern Small Plethodons Western Plethodons Figure 2. Suggested phylogeny of the major subdivisions of the genus Plethodon.

PAGE 33

27'.Yestern Plethodons Plethodon vandykei Group Plethodon vandykei vandykei Van Denburgh Plethodon vandykei ldahoensis Slater and Slipp Pletho ; on vandykei larsclli uurns Plethodon vehiculum Group Flethodon dunni Bishop Pletho Ion vehiculum (Cooper) Plethodon productus Group Plethodon productus new name Plethodon neomexic^nus Group Plethodon neomexJcanu e Stebbins and Riemer Eastern Small Plotho^ons Plethodon seller! Group Plethodon welleri alker Plethodon richmondi richnondi Netting and Ifittle ian Pletho; ion richmondi popei iii^hton and Grobman Plethodon riclunond i nettingi Green Plethodon cinereus Group Plethodon dorsalis dorsalis Cope Plethor'.on dorsalis angusticlaviu3 Grobman Pletho?on cinereus cinereus (Green) Plethodon cinereus serratus Grobman Plethodon cinereus polycentratus Highton and Grobman

PAGE 34

28 Eastern Large Plethodons Plethodon wehrlei Group Plethodon wehrlei wehrlei Fowler and Dunn Plethodon wehrlei dlxi Pope and Fowler Plethodon wehrlei jacksoni Newman Plethodon yonahlossee Group Plethodon yonahlossee Dunn Plethodon ouachitae Dunn and Heinz e Plethodor caddoensis Pope and Pope Plethodon glutinosus Group Plethodon jordani Jordan! Blatchley Plethodor. jordani metcalfi Briraley Pi ethodon jo rdani shermani Ste jneger Plethodon jordani unlcoi new subspecies II ethodon jordani melaventris Pope and Hairston Plethodon jordani rabunensis Pope and Iiariston Plethodon jordani teyahalee hairston Plethodon jordani cle nsonaa Brimley Plethodon glutinosus glutinosus (Greta) Plethodon ^;lutin osus albagula Grobraan

PAGE 35

29 The Western Plethodons The Western Plethodons include five species, none of which, except for dunni and vehiculum , appear to have close affinities. P. vandykei , with three presently recognized subspecies, vandykei , larselll , and idahoensis , occurs in coastal Washington, northwestern Oregon, and northern Idaho. P. dunni is known from southwestern Washington and coastal Oregon. P. vehiculum ranges on the coast from central Oregon to southern British Columbia, including Vancouver Island. P. productus (formerly P. elongatus ) occurs in a small area in northwestern California and southwestern Oregon. P. neonexicanug is known only from the Jemez fountains of New Mexico. Stebbins (I9f>l) has recently surraarized the knowledge of the amphibians of western North America, and has given detailed descriptions of all five species in life. Little new information on individual or geographic variation can be offered here since I have had no field experience with these animals and only four forms ( idahoensis , dunni , vehiculum , and productus ) have been examined in life. Reference should be made to Stebbins* book for further information on these forms. Radiographs of a series of each of the forms (except the recently described P. v. larselli ) were taken and the number of trunk vertebrae in each form is shown in table I. The usual number of trunk vertebrae in each species of Western Plethodon is different, and none of the known forms possess 18 trunk vertebrae as the modal number. Except for vehiculum , all of the Western Plethodons are fairly larp;e in size. All but neomexicanus possess a stripe phase in the adult, and all but dunni and vehicului have webbed toes. The vomerine tooth

PAGE 36

-30TABI£ I THE NUMBER OF TRUNK VEitTEBRAE IN OSTEIN FORMS OF THE GKNUS PLETHODOW Number of trunk vertebrae Form 15 16 17 18 19 20 P. . vandykei

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31 counts of all of the western forms are low compared to Eastern Large Plethodons of similar size; vehiculu m and productus have especially low vomerine counts. P. vandykei is unusual in possessing a parotid gland and in having the lowest number of vertebrae in the genus. P. neomexicanus is unusual in being the only species in the genus that has a reduced fifth toe on the hind limbs. PI et hod on dunni and Plethodon vehiculurri appear to be rather closely related. P. dunni attains a larger size than P. vehiculum and has more vomerine teeth. P. vehiculum usually has one more trunk vertebra than dunni . P. dunni has more aquatic habits than vehiculum . The color of the dorsal lipophores in the striped phase of the two forms is different. The stripe of dunni is usually greenish yellow, while that of vehiculum varies from reddish tan to yellow. The toes are very slightly webbed in both and most other structural features are similar. The other Western Plethodons possess a considerable amount of webbing on their toes. The amount of webbing on the toes does not appear to be correlated with aquatic tendencies, since the two most aquatic forms, dunni and vandykei , represent extremes in the absence and presence of webbing in the western forms. The forms vehiculum and productus , both with more terrestrial tendencies, also differ greatly in amount of webbing between the toes. Moreover, in the eastern United States, some of the forms have webbed toes, while others do not, yet all are terrestrial. There is no doubt that dunni and vehiculum should be regarded

PAGE 38

32 as distinct species, since the two occur sympatrically through most of the range of dunni without any evidence of interbreeding. A study of this relationship and the possible differences in the habitat niches of the two forms, suggest e 1 by Stebbins (19J>1* 65), is much to be desired. The forms, vandykei , productus , and neomexic&nus all seen to differ from e*>ch other as much or more than species groups in the Eastern Flethodons. The fact that they are so distinct morphologically makes it difficult to determine their relationships. In all probability, these forms have been differentiating for a long period of time. There has probably been a considerable amount of extinction of humid forest dwelling wastern fiorth American amphibians due to climatic changes which have occurred during or since the Tertiary. The present discontinuous distribution of this genus in the area, with only two species recorded in the Rocky fountains, separated by almost a thousand miles, supports this view. Without doubt there are probably other undiscovered forms of this genus in the western United States, .'iany salamanders of the region are difficult to collect, except during extremely favorable conditions, and some may have been overlooked in spite of intensive collecting. There remain 'nany ?reas in which there has been very little collecting, and these may be profitably searched for members of this genus. Lowe (1955s ?50), for example, believes that plethodontid salamanders will be found in the higher mountains of Arizona because conditions there are similar to those where they have been taken in other areas. The relationships of these salamanders would seem to be best indicated by the phylogeny shown in figure 3«

PAGE 39

33 VQnd )' ,
PAGE 40

3U Plethodon vandykei Group Plethodon vandykei This species is known from the Coast Range, Olympic fountains, and Cascade ijfountains of western ashington; Coeur d'Alene Lake, in northern Idaho; and from Larch :Jountain, Multnomah County, Oregon. The Idaho population was described as a distinct species ( idahoensis ) by Slater and Slipp (I9h0i 38), but has been considered a subspecies of vandykei by Lowe (19^0: 93) and more recent workers. Although the Idaho and Washington populations of this species are usually believed to be isolated from one another, Savage (195? '• 183) suggests that they may be connected by a corridor of high humid country along the international boundary between Washington and British Columbia. The two forms, idahoensis and vandykei , apparently differ mainly in color characters. In 195U, Burns discovered Plethodon vandykei in Multnomah County, Oregon. Two specimens from Larch Mount sin were unusual in possessing red bellies. A series of eleven specimens collected on the north side of the Columbia River, at Archer Falls, Skaiaania County, Washington, also agreed In coloration with the Oregon specimens and this red-bellied form was described as the third subspecies of vandykei , P. v. larselli . Stebbins (I95>ls 80-1) states that there are two color phases of P. v. vandykei . The light phase has the color of the belly and sides very similar to that of the dorsal stripe, while the dark phase has a darker lateral and ventral pigmentation. This species is usually found in very damp situations and thus replaces P. dunni ecolofically, as veil as geographically. It has re-

PAGE 41

35cently bean shown that their ranges do overlap slightly in Multnomah County, Oregon (Burns, 195U: 85) and Pacific County, Washington (Storm, 1955: 6h-5). Plethodon vandykei vandykei Van Denburgh Plethodon vandykei ?«B Denburgh (19o6: 61). Dunn (1926* 151-3). Slater (1933: liU). Bishop (19U3-* 275-8). Storm (1955: 6U-5). Plethodon vandykei vandykei Van Denburgh. Lowe (1950: 93). Stebbins (1951: 80-U). Stebbins (195U: 56-7). Sinter (1955: 132-3). Type :CAS 6910 (now destroyed), collected at Paradise Valley, Mount Ranier National Park, ashington, by E. C. Van Dyke, in July, 1905. Diagnosis ;A. testem Plethodon with 15 trunk vertebrae. It differs from idahoensis in having a adder dorsal stripe, li.hter ground color, and by presence of yellow pigment like that found on the dorsum on the proximal segment of the limbs. tiange tKnown from western Washington, from Clallam County south to Pacific County and east to Pierce County (see figure U). Description :This subspecies has not been seen in life and only four preserved specimens were exauned. Stebbins (1951: f 0-1) gives detailed color notes on this form. Costal grooves usually number 1U and the trunk vertebrae 15. In the available specimens, vonerine teeth range from 7 to 10 in a series. The largest individual is 56 mm. in snoutvent length.

PAGE 42

-36P. vandykei O vandykei A lorselli / idahoensis Figure U. The distribution of the sutrpecies of Plethodon vandykei in Washington, Oregon, and Idaho.

PAGE 43

37 Plethodon vandykei idahoensis Sinter and Slipp Plethodon idahoensis Slater and Slipp (19k0: 38). Slater (19U.J 81, 85, 103). Bishop (19U3J 259-61). Hilton (19U8: 120). Plethodon vandykei idahoensis Slater and Slipp. Lowe (1950s 93) • Stebbins (1951: HO-U). Stebbins (l95ii: 56-7). Type:U3NK 11050li, an adult male, collected at the northeast corner of Coeur d'A.lene Lake, Kootenai County, Idaho, at an elevation of 2160 feet, by James R. Slater, on September 13, 1939. Diagnosi s:A race of Plethodon vandykei in which the yellow or orange dorsal stripe is narrower than in P. v. vnndykei and contrasts sharply with the lateral black ground color, and the proximal segments of the limbs are dark in color. Range ?Known only from the type locality in northern Idaho (figure U). Description ;The back has a dorsal orange or yellow lipophore stripe. The rest of the sni-al Is pigmented with melanophores, except for raelanophore gaps on the chin where there are yellow lipophores. There are also brassy guanophores in the iris, and a few scattered white guanophores on the belly and sides. The costal grooves usually number l)j, the trunk vertebrae 15. Vomerine teeth range from 5 to 12 in a series. Of 15 specimens examined, the largest is 56 mm. in snout-vent length. Plethodon vandykei larselli Lurns Plethodon vandykei larselli Burns (195h» 83-7).

PAGE 44

38 Type :USNM 13U129, an adult male, collected on the north slope of Larch Mountain, three milos from the summit on the ultnomah Falls Trail, Multnomah County, Oregon, on ?tay 2U, 1953, by Douglas M. Burns. Diagnosis tA race of Plsthodon vandykei in which the ventral color is cardinal red to reddish orange. Ran^e iKnovm only from the type locality and from Archer Falls, Skamania County, Washington (figure U). Description :Specimens of this subspecies have not been examined and reference should be made to the original description for details of structure and coloration. Plethodon vehiculum Group Plethodon dunni Bishop Plethodon dunni Bishop (193U: 169). Jewett (1936: 71). Fitch (1936i 637-8). Graf, Jewett, and Gordon (1939: 102). Gordon (1939: 55-6), Slater (1939: 15U). Bishop (19h3: 2U2-6). Stebbins (1951: 68-72). Stebbins (195U: 5U-5). Storm (1955« 61*-5). Dumas (1955: 65). Slater (1955: 132). Type :USNM 95196, an adult female, collected just outside the city limits of Portland, Clackamas County, Oregon, by Stanley G. Jewett, Jr., on January 13, 193i». Diagnosis :A Western i'lethodon with 16 trunk vertebrae. Range :Curry County, in southeastern Oregon, north to Pacific County,

PAGE 45

39 Washington, east to the western slope of the Cascade ikjuntains (figure Description :The dorsal stripe is made up of yellowish green lipophores. This pigment also occurs in abundance on the sides, but is reduced on the belly. On the lovrer sides and belly there are a few yellowish guanophores. There are brassy guanophores in the iris. The costal grooves usually number 15, the trunk vertebrae 16. The medial end of the vomerine series projects posteriorly toward the parasplencid patches, so that the two vomerine series form a V. Vomerine teeth range from U to 13 in a series. The largest specimen examined is 65 nee, in st> out-vent length. P. dunni is closely related to P. vehiculum , but differs in size, coloration, number of vo*nerine teeth and number of body segments. It occurs from southwestern Washington south through coastal Oregon. This species occurs sympatrically with P. v andykei in southwestern Washington, with P. productus in southwestern Oregon, and with P. vehiculum throughout most of its range. P. dunni is apparently the most aquatic Plethodon . Stebbins (1951: 70) states that it is almost invariably found in places that are saturated with water, and that it will often take to the water in an attempt to escape capture. Most individuals possess a dorsal stripe that is greenish yellow in color, but Stebbins (195>1* 69) reports that nelanistic specimens lacking the dorsal stripe have been found in Benton County, Oregon.

PAGE 46

Uo Figure 5. The distribution of Plethodon dunni in Oregon and southwestern > ashing >m.

PAGE 47

UlPlethodon vehiculum (Cooper) Arobystoma vehiculum Cooper (1660: pi. 31, fig. U). Plethodon intermedius Baird (in Cope, 1867: ?09). Cope (1869* 100). Strauch (1870: 72). Boulenger (1882: 57). Cope (1883: ?3). Garman (163U: 36). Cope (1889: Hi5-7). Cox (1907s 52). Van Denburgh (1916: 218-9). Fowler and Dunn (1917* ?5). Hardy (1926: ?2). Kermode (1926: 35). Dunn (1926: 15U-6). Slevin (1928: 52-5). Logier (1932: 317-8). Slater (1933s UU). Svihla (1933 s 39). Slater ( 1931a s llO-l). Slevin (l93Us U6). Cowan (1937s 18). Plethodon vehiculus (Cooper), bishop (193U$ 171). Jewett (1936: 71). Watney (1938: 89). Slater (1939s 15U). Graf, Jewett, and Gordon (1939s 10-1). Brown and Slater (1939s 9). Plethodon vehiculum (Cooper). Slater (19k0: li3). Slater and Brown (19U1: 75-7). Bishop (19it3» 278-81). Stebbins (1951 « 8b-7). Stebbins ( 1951.1 : 57-9). Logier And Toner (1955: 17). Slater (1955s 133-h). Type :Apparently no longer in existance. The type locality is Astoria, Oregon. Diagnosis :A Western Flethodon with 17 trunk vertebrae. Range sFrom Coos County, in southwestern Oregon, north to southwestern British Columbia, including Vancouver Island (figure 6). Description :Lipophores in the region of the dorsal stripe may be absent, producing a uniformly colored phase, or present, resulting in a

PAGE 48

-U2 Figure 6. The distribution of Plethodon vehiculum in Oregon, 'Washington, and British Columbia.

PAGE 49

-U3striped phase. The color of the dorsal stripe is quite variable, varying from light yellow, through yellow, orange, and red to brown. The lipophores are absent from the sides, but are present on the belly. The sides are black with only a fe?/ small white guanophore spots. These are also present on the belly, and along with the melanophore and lipophore pipmentation give it a mottled appearance. Small white guanophore spots similar to those on the sides, as well as smaller brassy flecks, are present on the dorsum of the dark phase individuals. Brassy guanophores are present in the iris. Costal grooves usually number 16, trunk vertebrae 17. Vomerine teeth range from 3 to 7 in a series and the two series converge posteriorly to form a V as in dunni . The largest specimen h9 ex-' '-dined is 5l mm. in snout-vent length. This species is the smallest Western Plethodon and has the largest range. It is the most abundant Plethodon over most of its range. P. vehiculum is superficially quite similar in appearance to the eastern P. cin er eus . Dark and striped phases are present and the belly is mottled in both species. The similarities between the two are probably due to convergence or parallel evolution because in all other characters, P. vehiculum is morphologically more similar to the other Western Plethodons. Plethodon productus Group Plethodon p roductus , new name Plethodon elongatus Van Denburgh (1916: 216—8). ( non ) Salaraandra elongata Valenciennes in Dumeril and Bibron (18£U« 81;). Grlnnell and

PAGE 50

-UUCamp (1917* 13U). Storer (192$: 21, 10U). Dunn (19?6: 156-8). Slevin (1926: 55-7). Slevin (1931*: U6-53). Wood (193U: 191). Fitch (1936: 636). Wood (1939: 110). Gordon (1939: 13, X>, 56). Bishop (19UJ: 21*6-9). Hilton (I9I46: U5). Stebbins and Reynolds (19h7: Ul-2). Stebbins (1951: 72-6). Stebbins (1951: 55-6). Type :CAS 29096, collected at Requa, Del Horte County, California, by J. ft. Slevin, May 22-26, 1911. Diagnosi s:A 1 estern Plethodon with 19 trunk vertebrae. RangerSouthwestern Oregon and northwestern California (figure 7). Descriptio n*The belly of this species is very dark with a few sacttered white guanophore spots. The orange to reddish brown dorsal lipophore stripe is usually brighter in juveniles than in adults, where it is often reduced or absent. Often it is divided into a right an-' left dorsolateral stripe by the presence of medial melanophore pigment. The costal grooves usually number 18, the trunk vertebrae 19. Vomerine teeth ranfe from U to 7 in a series. This is a lar^e species, the largest of 18 specimens exandned is 66 mm. in snout-vent length. This is the most elongnte of the estern Plethodons in the Pacific Northwest. Its greater number of body segments distinguishes it from the species vandykei , dunn i, and vehiculum , and its webbed toes also distinguish it from the last two. It is probably more closely related to P. neomexLcanus than the other -Vestcm Plethodons. P. productus has a rather li-nited distribution. It is known only from Trinity, Humboldt, and Del Norte (Jounties, C-lifornia; and Curry County, Oregon.

PAGE 51

us >
PAGE 54

U6 Stebbins (1951* 76) believes that specimens from the interior of the range differ in several respects from those on the coast. Van Denburgh described this species in 1916, apparently without realizing that the name he propose*', elongatus , was a junior homonym of Salamandra elongata Valenciennes (in Dumeril and bibron, 185U: 8U) ( = Hethodon glutinosus ) . This is a primary homonym due to the fact that the name elo ngata is available for Plethodon glutinosus , although the combination Plethodon elongatus has never been used for the slimy salamander. No subsequent worker has proposed a new name to replace the preoccupied name of the Del Norte salamander and it has always been referred to as Plethodon elongatus . This species has been known by the name elongatus since 1916 and it is unfortunate that the name has to be changed, but there is no alternative than to propose a new name in place of the preoccupied elongatus. The name productus also refers to the elongated condition of this species. Plethodon neomexicanus Group Plethodon neomexicanus Stebbins and Riemer -urycea multiplicata (Cope). Dunn (1926: J12-U, part). Plethodon neomexicanus Stebbins and PJLemer (1950: 73-60). Stebbins (1951: 76-9). otebbins (195U* 58). Type :MVZ U9033, an adult male, collected 1? miles west and h miles south of Los Alamos, Sandoval County, New .' exico, at an altitude of about 8750 feet, by Robert Stebbins, on August lb, 19U9. Diagnosis :A estern Plethodon with 20 trunk vertebrae.

PAGE 55

».7 • i Range rKnown only from the vicinity of the type locality in the Jemez Mountains of Mew Mnxico, (figure 8). Description :This species has not been examined in life and reference should be made to the original description for an account of the pigmentation . Costal grooves are usually 19, trunk vertebrae 20. Vomerine teeth range from h to 11 in a series in the 7 specimens examined. This is a large species, the type is over 70 mm. in snoutvent length. According to the data listed in Stebbins and Riemer (1950: 7$), sexual maturity is not reached until approximately £0 mm. in snoutvent length. This is an elongated salamander, with the largest number of trunk vertebrae of any of the western forms. It is unusual in that most specimens have only one phalanx on the fifth toe of the hind limb instead of the usual two phalanges. This is probably an intermediate stage in the loss of the fifth toe and reduction to the four-toed condition haw to have occurred in several unrelated salamander genera. The dorsal stripe is apparently absent in adults, but present in juveniles. This form is unique among the estern Plethodons in lacking the striped phase in the adult, and also in possessing a large number of brassy guanophores on the dorsum. The Eastern Small Plethodons The Rastem Small Plethodons are characterized by their small size. Except for P. welleri , they possess a higher number of trunk vertebrae and are relatively more elongate than the Eastern Large

PAGE 56

U8 Flethodons. Four species are included in this group. P. weileri has a restricted ran f 'e, and is knovn only from a few counties in northwestern North Carolina and adjacent Tennessee and Virginia. P. richmondi is a polytypic form with three described races, and probably others which have not yet been defined. Its range centers in the Appalachian Plateaus i hysior;raphic lrovince Bad it also occurs in immediately adjoining areas of several adjoining Provinces. P. clnere us, with three subspecies, has the widest range of any Eastern Small Plethodon, occurring over most of eastern United States and southeastern Canada, except in tha region occupied by P. dors alia . The ranpe of _. dorsalis is centered in the Interior Low Plateaus Physiographic I rovince. P. d. angusticl^yjus is here regarded as a subspecies of dorsalis , rather than of cine reus, as originally described by urobman (19W*: 302). The four species forvi natural group of closely related forms. All possess webi.ed toes, a pigmented peritoneum, lot; vomerine tooth counts, brasry flecl:s on the dorsum, and white guanophore spots on the belly and siies. . . o species, cine reus and dors r. li s, are more similar to each other than either is to weileri or richmo ndi . hoth cinereus and dorsalis typically possess at least as much white as black pigment on their bellies, and both are characterized by the presence of a redbacked phase in which there is l dorsal band of red or yollow lipophore pigmentation on the dorsum of the body and tail. In richmondi and velleri , the white pirment on the belly is limited to small spots and there ij much greater amount of blac 1 ,pigment than white. The redb-jcVod phase, present in all of the races of dorsalis and clnereus ,

PAGE 57

-U9is always absent in richmondi and welleri . P. r. r ichmondi is the largest form, and also possesses the greatest number of trunk vertebrae, but both size and number of vertebrae are slightly reduced in the race popei , and decrease still more in the subspecies netting! . Geographic variation in the number of trunk vertebrae is most pronounced in this species, with a range of 18 to 2li in all of the subspecies. P. welleri is less elongate and has fewer vertebrae than other small eastern species. It is unusual in its possession of ovarian eggs pigmented with melanophores , as well as in having an extremely dark parietal peritoneum. The brassy flecks on the dorsum of welleri are concentrated to form large patches, giving it a unique appearance not found in any other Plethodon . Its nearest relative is probably P. r. nettingi , but welleri is closest geographically to P. r. pop el . In most of their ranges, cinereu s and dorsalis are allopatric forms, but in several areas where their ranges meet, there are records of the two occurring together in the same locality (see below). In each of these instances, there is no good evidence for hybridization or intergradation between the two forms, and for this reason they are usually considered as distinct species. They differ in several pigmentation characters, as well as in the average number of trunk vertebrae and the shape of the mental gland in adult rnalee. P. dorsali s usually has 19 trunk vertebrae (range 18-20). Two races of cinereus typically have 20 trunk vertebrae (occasionally 19 or 21), while the third subspecies ( polycentratus ) usually has 21 or 22 (rarely 23) trunk vertebrae. In all of the Eastern Small Plethodons except dorsalis,

PAGE 58

*>the mental gland is rather poorly developed and difficult to differentiate (except, perhaps, during the breeding season) from the adjacent portion of the chin. In dorsalis, on the other hand, it is a distinct rounded elevated gland, as in the Eastern Large Plethodons. Dunn (1926% ?U) suggests that the striped pattern or dorsalis is more primitive than that of cinereus, since the paired red dorsal spots of several primitive plethodontids might be expected to pass through a zig-zag striped phase similar to that of dorsalis before evolving into a uniformly straightedged stripe. In 19hU, Grobman described the narrowest riped populations occurring in southwestern Missouri and northwestern Arkansas as a distinct subspecies, Flethodon cinereus angxtsticlavius . This form possesses several characteristics that appear to link it more closely with dorsalis than with cinereus . It usually has 19 trunk vertebrae, as in dorsalis , while cinereus populations to the northeast and south of the range of angusticlavius usually have 20 trunk vertebrae. The mental gland is of the dorsalis type and in some specimens the dorsal stripe has irregular edges anteriorly, resembling very closely the pattern in dorsalis . Living specimens of angusticlavius have not been examined, so no accurate information is available on pigmentation characters, but the other characters strongly suggest that angusticlavius should be linked subspecifically with dorsali s rather than cinereus . The elongation and increased number of trunk vertebrae in P. r. richmondi, P. r. popei, and P. c. polycentratus would .appear to be a marked specialisation of the usual plethodontid type. The forms welleri and dorsalis would appear to have retained more primitive char-

PAGE 59

51acteristics, but both have probably changed considerably from their common ancestor. The relationships of the Eastern Small Plethodons would appear to be best indicated by the phylogeny outlined in figure 9. Plethodon welleri Group Plethodon wellori alker Plethodon welleri ..alker (1931? U8-51). Walker (193U: 190). Bishop (19U3s 285-7). Grobman (I9hk: 313). Snyder (19H6» 17U). Hoffman and Kleinpeter (19U8a: 107). Hoffman (19^3* 86-7). Type :USNM 8lil35, an adult male, collected on Grandfather fountain, at an altitude above $000 feet, near Linville, North Carolina, by IS. H. Weller and Ralph Dury, on August ?7, 1930. Diagnosis ;k dark-bellied, Eastern Small Plethodon with 17 trunk vertebrae and abundant dorsal brassy spotting. Range ?From Yancey County, iiorth Carolina, northeast in Tennessee and North Carolina to Mt. Rogers and ihite Top fountain, Virginia (figure 10). Description ;The dorsal pattern consists of large anastomosing patches of brassy guanophore spots. These spots appear to consist almost entirely of brassy guanophores, the white type of guanophore pigmentation present on the sides of welleri and on the dorsum of most plethodons, is not evident on the back of welleri . The brassy pigment is more concentrated than in any other form and the appearance of the dorsal spots is similar to the large white spots of glutlnosus , except for the color,

PAGE 60

*2cinereus serrotus popei polycentrafus / / ongusticlovius dorsalis richmondi / nettingi wellen richmondi Figure 9. Suggested phylogeny of the Eastern Small Plethodons. Figure 10. The distribution of Plethodon welleri in North Carolina, Tennessee, and Virginia.

PAGE 61

53which is brassy. Small white spots (0.1-0.2 mm.) are scattered over the belly and are similar to those of P. r. popei . The lateral spots (up to 1 ran. in diameter) are white with a slight amount of brassy flecking. The costal grooves usually number 16, the trunk vertebrae 17. Vomerine teeth range from 2 to 7 in a series. In the 39 specinens examined, maximum known snoutvent length is between U5 and $0 mm. Sexual maturity is reached at about 35 »m. in snoutvent length. P. welleri is the least elongate of all of the Eastern Small Plethodons, with the same number of trunk vertebrae (17) as in most of the Eastern Large Plethodons. In body build it resembles the large forms more than any other Eastern Sm?ll Plethodon. Its small size, vertebral structure, webbed toes, and pigmentation characters would appear to associate it more closely with the other small eastern forms. This species has long been believed to be restricted to high elevations and was recorded only from Flat Top fountain and Grandfather Mountain, riorth Carolina; and itb. Rogers and hite Top Mountain, Virginia. Recently, Hoffman (1953' 86) collected the species at an elevation of 2500 feet in Johnson County, Tennessee. I visited this locality during the summer of 1955, and although conditions were very dry, succeeded in collecting one young specimen, thus confirming Hoffman's record. This species may well occur in suitable habitats at low altitudes throughout the region. V/alker (193U» 190) has pointed out that individuals from hite Top Mountain, Virginia, possess more spots on their venters than individuals from the type locality, Grandfather Mountain, Worth Carolina.

PAGE 62

-& Specimens from », u^ers (USNK 12lih?l-9) ~lso possess spotted bellies. One specimen from Flat Top fountain, Yancey County, reported by Snyder (19U6: 17U) is also described as having a more conspicuous mottling on the venter. Perhaps the Grandfather Mountain population is unusual in lacking this characteristic. The low altitude specinen from Tennessee has a greater amount of dorsal brassy pigment than eight living topotypes from Grandfather fountain. I have not examined living specimens from any of the other localities. P. welleri is unusual in possessing dark ovarian eggs that are pigmented with nelanophores . Its closest relative is probably P. richraondi , Tfhich it resembles by the presence of a dark belly. F. r. netting! usually has 19 trunk vertebrae, although occasional specimens possess 18 or 20. Since a small percentage <>f welleri also have 18 trunk vertebrae (table II), there is slight morphological overlap between the two forms in this character. TABLE II THE NUMBER OF TRUNK VERTEBRAE IN GEOGftlPtilC SAMPLES OF PLETHDDON WELLERI Number of trunk vertebrae Locality 16 17 16 Grandfather {fountain, North Carolina 25 Johnson County, Tennessee 1 "hite Top Mountain, Virginia U Mt. Rogers, Virginia 16 2

PAGE 63

55Plethodon richmondi In 1938, two new Eastern Small Plethodons, P. rlchmondl and P. netting! , -were described by Netting and Vittleman (1938 1 297) and Green (1938: 295). Both are characterized by their dark bellies and brassy dorsal flecking. Available samples of richmondi from Ohio and ftest Virginia possess 20 to 22 costal grooves (21 to 23 trunk vertebrae), while P. netting! has 17 to 19 costal grooves (18 to 20 trunk vertebrae). They both differ from P. c. cinereus in the number of vertebrae, and they are distinguished from the entire cinereus group by their dark bellies and lack of red pigment. Most previous writers are in agreement that these two forms, richmondi and netting! , are more closely related to each other, and to welleri , than they are to cinereus and dorsalis (Netting and Mittleman, 1938 s 292 j Creeni 1938s 298j Grobman, 19UU*. 311). In 19U9, Grobman described another fl«cked plethodon (P. huldae) which he assigned to this group because of its heavily flecked dorsum. This form possesses 20 trunk vertebrae, the same as cinereus , and also has a mottled belly as in cinereus . Rabb (1955* 261-3) and rftichmore (1955: 170-2) present convincing evidence that huld ae, although possessing brassy dorsal flecks similar to those of richmondi and nettingi, is actually based on dark-phase specimens of P. cinereu s . tfachmore (1955: 172) goes so far as to state that "since P. c. cinereus possesses brassy flecks on the dorsum and since F. huldae can no longer be considered valid, it is certainly desirable that the concept of a welleri group of Plethodo n distinct fron a cinereus group be abandoned." Obviously these forms are all more closely related than any of than are to the Eastern Large Plethodons or the Western Plethodons, yet the fail-

PAGE 64

-56ure of one suggested character to define the group should have no bearing on the fact that other features are useful in distinguishing it (dark belly and lack of red dorsal stripe). Netting and s»ittleman (1938: 292) state that spacinens of P. richmondi frora ttatauga County, North Carolina, differ from typical richmondi , but do not give any evidence to support this contention. Neither Bishop (19U3' 239) nor Grobm&n (lSol*: 312) include this locality in their distribution myps of the range of richmondi , although Grobman mentions the literature reference to the North Carolina specimens. Afore recently, fcoffman and Kubricht (195U$ 192) report richmondi frora several localities in southwestern Virginia and northwestern liorth Carolina, but do not find any consistant differences between these specimens and topotypes of richmondi . A study of their material and large series frora this area in the University of Florida Collection have shown, however, that these southern richmondi are actually quite distinct in the number of trunk vertebrae from more northern richmondi . This southern form has recently been described as a new subspecies of richmondi , P. r. popei , by Highton and Grobman (in press). No other character could be found to distinguish preserved specimens of popei and richmondi . Living specimens of P. r. richmondi have not been available during this study, so it has not been possible to compare the pigmentation characters of the two forms. On the basis of the slight overlap in ..dstinguishing characters, as well as the geographic replacement of the three forms, richmondi , popei , and netting! , highton and Grobman (in press) suggested that they should all be considered subspecifically related. One

PAGE 65

-57matter, not discussed in their paper, is the apparent sympatric relationship of netting! and richmondi , Although the two forms, to my knowledge, have not been collected at the same locality, the entire range of nettingi , as summarized by Brooks (I9u8), is surrounded by records of richmondi . Thurow (1955) has recently reported netting! from Bedford County, Virginia, a locality about equidistant from the nearest known nettingi and popel localities. An examination of these specimens (CNHM 6o5l2-8 ) confirms Thurow' s suggestion that they belong to this group. Although they are poorly preserved, their dark bellies indicate that they are not dark-phase cinereus . They differ from both nettingi and popei , however, in costal groove count. Five of the seven specimens have 19 costal grooves, while the other two have 18. The sample is small, but the probability that they were taken from a population of nettingi similar to that in Randolph and Pocahontas Counties, West Virginia, which has 19 costal grooves less than 5£ of the time, is very low. Four of the seven Bedford County, Virginia, specimens have 20 trunk vertebrae, two possess 19, and the remaining specimen is too small to obtain an accurate count from the radiograph, but it probably also has 20 trunk vertebrae, since it has 19 costal grooves. The population from which these specimens were collected probably represents an intermediate one between popei and nettingi . If the usual number of trunk vertebrae is 20, and this occurs with high frequency, this population should probably be given separate subspecific nomenclatorial status. At present, with only seven poorly preserved specimens available, too little is known of its variation to described it as new here. Its importance lies in the fact that an intermediate population, often

PAGE 66

-repossessing 20 trunk vertebrae, is known from a locality between the ranges of netting! (usually with 19 trunk vertebrae) and popei (usually with 21 trunk vertebrae). This may be considered further evidence that netting! and popei are subspecifically related. In the Valley and Ridge Province of Virginia, there exists a population of P. r. richmondi which Hoffman and hubricht (19SU* 192) believe to be different in both color and structural features from other populations of P. richmondi . They do not state the ways in which it differs, but an examination of preserved specimens from this region (Alleghany and Rockingham Counties, Virginia) indicates that these animals, although possessing a vertebral cour.t similar to richmondi , differ from it in having a mottled belly. Specimens from these counties are easily segregated from topotypical richmondi and popei on the basis of the character of the belly. In ten specimens of the mottled bellied form for which vertebral counts are available, six possess 22, three 23, and one 2h trunk vertebrae. The average is slightly higher than the usual richmondi number, but does not differ significantly from it. It is quite likely that examination of living specimens, and the accumulation of more data on vertebral counts may reveal differences which will indicate that this population represents a distinct nominal form. Specimens from other areas in the Valley and Ridge Province should be studied in order to delimit the ran^e of the mottledbellied richmondi . The distribution of these forms indicates that there is probably a series of five races, two of them undescribed, each of which replaces its nearest relative geographically. One ext^(nettingi) occurs sympatrically with the two at the other extreme ('rLc)imondi and

PAGE 67

»the undescribed Virginia Valley and Ridge animal). Table III shows the variation in the vertebral counts of these populations. TABLE III GEOGRAPHIC VARIATION IN THE NUMBER OF TRUNK VERTEBRAE IN PLETHODON RICHMONDI Sample Number of trunk vertebrae 18 19 20 21 22 23 21* Mean P. r. netting!

PAGE 68

60 without any evidence of interbreeding. An accurate knowledge of the •coloeical distribution of the overlapping forms of P. richraondi is lacking. There appears to be little or no overlap between the two terminal races of Ensatin a eschscholtzi , whereas in this case, one ( netting! ) definitely occurs within the range of richmondi , for there are records of the latter to the north of the Cheat ^fountains in Pennsylvania, to the east and west in West Virginia, and to the south in Virginia. In these series of populations we probably see the stages through which the subspecies of richmondi have differentiated. All of the forms probably have changed somewhat from the original stock, but the close similarity of all the forms, except for the number of body segments, would seem to indicate that this change has not been great. It is difficult to determine which form most closely resembles the ancestral condition. P. r. richraond i is unlike most plethodons in its elongate form and would appear highly specialized. On the other hand, it wide distribution and the fact that the eastern Valley and Hidge population has a mottled belly, approaching that of the cinere\i3 group, ndcht be considered evidence that this form is a more primitive generalized animal. It is possible to reconcile these viewpoints, for it is often found that a given form is specialized in some ways, yet retains certain other primitive characteristics. Since P. r. netting! is apparently limited to high altitudes in the Cheat Mountains, it oould be interpreted either as a specialized type adapted to this habitat, or as a relict of a once more widely distributed spruce forest population. At present, the evidence is not sufficient to choose between the two alternatives. On the basis of its

PAGE 69

61 fewer trunk vertebrae, n etting ! would appear to be less specialised than the other subspecies* Intergradation between the races of richmondi msy occur in certain regions. P. r. netting! insy now be completely isolated from its nearest relative, but the ranges of p opei I >nd richmondi probably meet in eastern Kentucky, an:; intergradation raight be ejected. Large samples frora critical areas will be necessary before conclusive evidence is obtained, since intererades between pope! and richmondi , for exanple, would be expected to possess about an equal number of specimens with 21 and 22 trunk vertebrae, I-arge series of specimens from each county in and on both sides of the area of intergradation would be neecos sary la determine its extent. Such series are not now available from Kentucky, and hollow symbols representing litemture records for this region on the distribution nap (figure 11), may represent popei , intergradient, or richmondi populations. Plethodon r i chmondi richmondi Netting and tSittleman Plethodon richmon d Netting and Wittleman (1938* 267). ^ettinj(1939* $0-1). l>uxy and Geseing (19U0* 31). Bishop (19U.,: 72-$). Grobaen (19UU* 312). Wood (19U5a« 19). Wood (19h5b» 206-10). Netting (191*6 1 12). ood (19h6: 169). Stood and Duellnan (19U7* 3). Grobman (3Jfcfl 135). ftLchsnond (1952? 31fc). Green and Walker (195U* 60). rjuellna* (195U* ti0-5). Hoffman and Hubricht (1951.1 191-3). Plethodon richmondi richmonil Netting and Uittleraan . Bightcn and Grobman (in press). 1ype »CM liil89, an adult male, collected in Hitter Park, Luntington,

PAGE 70

62 Cabell County, .Vest Virginia, at an elevation of 600-700 feet, by Neil D. Richmond and N. Bayard Green, on October 15, 1938. Diagnosis »A dark-bellied Eastern Small Plethodon, usually with 22 trunk vertebrae (range ?l-2ii), which completely lacks red pigment. Range :From Centre County, Pennsylvania, south through western Maryland, '."est Virginia, and northwestern Virginia, west to northeastern Kentucky, and north to southern and eastern Ohio (figure 11). Description ;This form has not been examined in life and nothing on the pigmentation can be added to previously published accounts. The costal grooves usually number 21, the trunk vertebrae 22. Vomerine teeth range from 3 to 9 in a series. This is the largest Eastern Small Plethodon, the largest specimen examined, from Alleghany County, Virginia, is 60 ram. in snoutvent length. Plethodon richmond i popei Kighton and Grobman Plethodon richmondi Ivetting and I/ittleraan. Barbour (1953* 65-6). Hoffman and Hubricht (1951* 191-3). Plethodon richmondi popei Highton and Grobman (in press). TypeiUF 8226, a maturing :nale, collected at Comers Rock, GraysonWythe County line, Virginia, by Arnold B. Grobman and iarc R. Grobman, on August 5, 1955. Diagnosis *A race of Plethodon richmondi that usually possesses 21 trunk vertebrae (range 20-22). Range *Known from Harlan County, Kentucky; Tazewell, Smyth, Grayson,

PAGE 71

-63Figure 11. The distribution of the subspecies of Plethodon rlchraondi . Solid symbols represent localities from w* Lch specimens have been examined* Hollow symbols. represent literature records.

PAGE 72

-Aland Wythe Counties, Virginia; and Ashe, Alleghany, and 'atauga Counties, North Carolina (figure 11). This form may have a greater range than now known. Specimens from adjacent Kentucky and »>est Virginia have not been examined. They may also belong to this race. Description :Living specimens possess both types of guanophore spots on the dorsum. Small brassy flecks are very abundant and larger white spots are also present. On the sides and venter there are larger white or yellow spots. The costal grooves usually number 20, the trunk vertebrae 21. Vomerine teeth range from 3 to 8 in a series. The largest specimen examined is UR mm. in snoutvent length. Sexual maturity is reached between 35 and b5 mm. in snout-vent length. Plethodon richmondi netting! Green Plethodon netting! Green (193B; 295-9). Bishop (19U3' 266-9). Grobraan (19iiii: 313). Brooks (19U5: 231). Brooks (19U8: 239-UU). Plethodon richmondi nettingi Green. Highton and Grobman (in press). Type :CM 10279, an adult male, collected on Barton Knob, near Cheat Bridge, Randolph County, West Virginia, at an elevation of about hOOO feet, by M. Graham Netting, on June 29, 1935. Diagnosis :A race of Plethodon richmondi that usually possesses 19 trunk vertebrae (range 18-20). Range :Known from altitudes above 3500 feet in the Cheat Mountains of Randolph and Pocahontas Counties, West Virginia (figure 11).

PAGE 73

65Description :In life, the dorsum of this form is similar in coloration to P. r. popei . The costal grooves usually number 18, the trunk vertebrae 19. Vomerine teeth range from 3 to 8 in a series. The largest specimen examined is h$ mm. in snout-vent length. This form appears to have a smaller average size than the other races of rich mondi . ^exual maturity is reached at about 35 mm. in snout-vent length. Plethodon cinereus Group Plethodon dors alls The range of P. dorsalis centers in the Interior Low Plateaus Physiographic Province, but does not appear to be restricted to it, for records are available from several adjacent provinces. This species has not heretofore been recorded from Georgia, but specimens are now available from four different localities in that state, three of which are in the Piedmont Province. The southernmost record is from Upson County (UMMZ 8557U), only a few miles above the Fall Line. Other records are for Henry County (UF 8371 (3), UF 8U13 (3))j Cobb County (UG 206 (6)j UG 275 (36))| and Dade County (ER/UWTN 12128 (1?)). Probably the natural range of this form includes the western part of the state above the Fall Line. The sympatric occurrence of this species with P. c inereus in Georgia, Tennessee, Indiana, Illinois, and Oklahoma will be discussed below. The two forms are very similar in appearance, and it is often difficult to identify individual specimens . For this reason a detailed study of the two species in the areas of overlap will be necessary be-

PAGE 74

66 fore it can be determined whether or not hybridization occurs. In Georgia, where there is no overlap in the vertebral counts of the two species, it is impossible, on other grounds, to identify a small percentage of the available specimens. There are, however, several differences between the two species in Georgia that are apparent in a large majority of the available specimens. These include the shape of the mental gland in adult males; the zig-zag stripe of dorsal is , compared to the straight-edged stripe in cinereusj the presence of more red pigment in front of the eyes on the head of dorsalis ; the great reduction in amount of melanophore pigmentation on the belly of dorsalis ; and a similar reduction in red lipophore pigmentation on the belly of cinereus . The fact that there are a few exceptions to each of these species characteristics might be considered evidence for hybridization were it not for the fact that occasional specimens of each species from localities in which the other form is absent show at least some of the same variations from the usual pattern. The fact that there is no tendency toward an increase in the number of trunk vertebrae in Georgia dorsalis populations is also significant (see table IV). It may be seen from this table that there is no strong tendency for an increase in the number of vertebrae in samples of dorsalis that are from areas in which geographic overlap with cinereu s occurs. We may conclude that there is, as yet, no good evidence for hybridization between the two species, but that more study is needed in all of the areas in which the two occur together. There are no available records of dorsalis in southeastern Missouri. This region should be explored in order to determine whether or not dorsalis or ancusticlavius are present. Smith (19U8: 1) has

PAGE 75

-67recently reported dorsalis from several localities in southeastern Illinois. TABLE IV THE NUMBER OF TRUNK VERTEBRAE IN SAMPLES OF PLETHODON DORSALIS

PAGE 76

68 a slight amount of red pigment in the region of tne tack in whicn the stripe is located in red-backed individuals, so that an outline of the irregular stripe can be seen when held at a certain angle to the light. Phe presence of the red pigment, as well as a reduction in the cielanophore pigmentation does give the lead-backed phase of dorsalis a lighter appearance than the corresponding phase of c ina reus. urobman (19U+: 308) presents evidence to show that Baird should be credited with the authorship of the name dorsalis . Under Article 21 of the International Rules of Zoological Nomenclature as of 19Uh, * e was correct. Since 19UU, however, the Rules have been amended, so that Cope should now be recognized as responsible for this name. Plethodon dorsalis dorsalis Cope Plethodon cinereus dorsalis Cope (1889: 13ft-9). Blanchard {1926: 368-9). bishop (19h3: 236-9). Parker (19U8: 22). Chermock (1952: 29). Plethodon erythronotus (Green). Jarman (189U» 38). Plethod on dorsalis Cope. Stejneger and Barbour (1917: 15). Dunn (1918: 1*60-2). Dunn (1926: 158-62). Mohr (1937*UO) . Parker (1937: 63 X Parser (1939: 7?). King (1939: 550-1). Swansori (1939: 687). Grobman (19UU: 308-11). S-4th (19liP: L). Sinclair (1950: 50). Mohr (l<>52: 59-60). Thnrow (1955: 62-3). Holman (1955-* 1U3). Type;USNM 3776, collected at Louisville, Jefferson County, Kentucky. Diagnosis :An Astern Small Plethodon with the usual number of trunk vertebrae 19 (range 18-20), and a red dorsal stripe with irregular edges

PAGE 77

-69 in the red-backed phase. Range tFrom southern Illinois, Indisna, and southeastern Ohio, south through Kentucky and Tennessee to northern Alabama and northwestern Georgia (firure 12) . Description :The dorsum of both color phases of P. d. dorsal is has small white spots (0.2-0 .U mm.) as well as smaller brassy flecks. The red-backed phase, in addition to the other pigments, contains a large amount of red lipophore pigment. All of these chromatophores appear to be identical with those of P. cinereus . The main difference between tha two species is in the abundance and distribution of the pigments. The melanophore background in dorsalis is somewhat reduced, giving the animal an overall lighter appearance than cinereus . The melanophore pigment on the belly of dorsalis is greatly reduced with a corresponding increase in the amount of red lipophores, so that the belly appears to be mottled with red and white rather than black and white, as in cinereus . In dorsalis there is a concentration of red pigment on the head in front of the eyes. The lateral guanophore pigment is often yellowish in color. The dorsal red stripe is quite variable. In some specimens the edges are irregular for the entire length of the body, while in others, they are irregular only in the anterior half or third of the body. The latter condition is especially common in Georgia specimens, and occasional specimens from Georgia have straightedged dorsal stripes as in cinereus . Specimens with straightedged dorsal stripes have been reported also from Indiana (Blanchard, 1926: 369) and Tennessee (Grobraan, 19UU» 309-10).

PAGE 78

-70P dorsoiis O dorsoiis A ongusticlovius Figure 12. The distribution of the subspecies of Flethodon dorsalis ,

PAGE 79

71 The costal grooves usually number IP, the trunk vertebrae 19. Vomerine teeth range from 3 to 6 in a series. This is a small species; sexual maturity is probably reached at about 30 mm. in snoutvent length. The largest specimen examined is Uh mm. in snout-vent length. Plethodon dors alls angusticlavius Urobman Plethodon cinereu s angusticlavius Grobman (19hht 302). Dundee (19U7* 117). Bragg and Hudson (1951: 69). bragg (1955: 27-8). Type *A.MNH U0366, an adult male, collected at Mud Cave, near Fairy Cave, Stone County, Missouri, by B. C. Marshall, on October 1, 1927. lUagnosis:A race of Plethodon dorsalis in which the width of the dorsal band is usually less than one-third of the width of the body. itenge ;Southwestern Missouri, northwestern Arkansas, and adjacent Oklahoma, north of the Arkansas River (figure 12). Description ;Living specimens of this form have not been examined. Of 33 preserved specimens (all from Arkansas), 10 are of the striped phase. 11 show no trace of a stripe, and the remainder show only a faint stripe on the body, but at the base of the tail, it widens and becomes much more well-defined. Typically striped angusticlavius also possess a wider and brighter dorsal stripe at the base of the tail than on the body. The brightness of the stripe in this region appears to be due to a reduction in melanophore pigmentation in the dorsal stripe. The dorsal stripe is often irregular anteriorly, as in dorsalis . The costal grooves usually number 18, the trunk vertebrae 19. Vomerine teeth range from h to 8 in a series. The largest specimen

PAGE 80

72 examined is h3 nan. in snout-vent length. Pjetholon cinereus The red-backed salamander is the most abundant terrestrial salamander over most of its range, which includes much of eastern Korth America. Two distinct color phases are present, one with a prominent red or yellow dorsal stripe, the other uniformly dark in appearance. The frequency of these color phases varies from one locality to another. In some places, both phases occur in approximately equal numbers, in others, one type may be rare or absent. The proportional distribution of these color phases in the various populations has recently been studied by Thurow, and it is hoped that the results of his work will be published in the near future. Occasional specimens have been reported that lack the dark pigment and appear entirely red. One specimen from New Jersey lacked the red pigment in life and had a colorless dorsal stripe . Plethodon dorsalls has been considered a subspecies of P. cinereus by some workers (Cope, 1669: 138; Blanchard, 1926: 269; Bishop, 19l»3 : 236), and as a distinct species by others (Dunn, 1926: 158; Grobman, 19UI;: 306). Grobman points out that although the two forms are largely allop."tric, at some localities, especially in southern Indiana, the two occur together, Grobman examined the series of dorsalls that King (1939: 55D reported from the Great Smoky Mountains of Tennessee and confirmed their identification. Specimens of both species found together at the same localities are represented in the GSMNP collection. These localities are White Oak Sinks (elevation 17!?0 feet) and the Sinks

PAGE 81

73 on the Little Kiver (elevation 1600 feet). On the Tennessee side of the Great Smoky Mountains, P. cinereus is known from 1600 to 5000 feet and P. dorsal is frora 1200 to 2200 feet. The two species also occur sympatrically in the Piedmont of western Georgia. At two localities, 3.8 miles north of ticDonough, ilenry County; and 8.5 ndles north of Thomaston, Upson County, the two species have been taken together. At all other localities in western Georgia, where either P. c. polycentratus or P. d. dorsali s have been collected, the other form has not been taken. Eragg (19E>5* 27) reports that he collected a specimen of P. c. serratus and a specimen of P. d. angusticlavis at the same locality in Cherokee County, Oklahoma. Apparently the two species often occur together where their ranges overlap. Smith (19lt8: 1) published several new records of dorsal i s from southeastern Illinois. P. cinereus ?nd P. dorsalis probably also occur sympatrically in that state. There appears to be no evidence of intergradation at any of these localities, confirming the conclusion of Dunn and Grobman that dorsalis and cinereus have reached the species level of differentiation. P. cinereus shows little apparent geographic variation over most of it 8 range, except in the frequency of occurrence of the two color phases referrec to earlier. Only in the Piedmont of Georgia and in Arkansas and Oklahoma, where the populations of this species have apparently become isolated from the parent stock, so we find differentiation of the magnitude required for the recognition of subspecies. The Arkansas and Oklahoma race (serratus) possesses a dorsal band with

PAGE 82

-7k3errations at each costal groove. This characteristic is sometimes slightly developed in some specimens of other populations, but never reaches the frequency of occurrence or degree of development present in serratus . The Georgia Piedmont subspecies ( polyc ent rat us ) differs from other populations of cinereus in its increased number of trunk vertebrae. Data on the number of trunk vertebrae in sables of P. cinereus are listed in table V. It also differs from most ci nere us in possessing red pigment on the belly between the front limbs. The cinereus populations in western North Carolina and eastern Tennessee are characterized by the complete absence of the dark lead-backed phase, P. c. polycent ratus differs from these adjacent cinereus populations by the presence of the lead-backed phase (38% of the type series of polycent ratus are dark phase). Sanders and Smith (19U9* 28) report a specimen of Plethodon cinereus from Fern Lake, near Nacogdoches, Nacogdoches County, Texas. This specimen (OS $£6) lacks the serrate edges in the dorsal stripe that are present in serratus , the nearest race geographically. It has 19 trunk vertebrae, but this number is not unusual in other populations of cinereus . Mr. Sanders, who collected the specimen, informs me (in letter of November 12, 1955) that it was taken with a dip net from a bunch of fruiting sphagnum moss near the shore line, a most unusual habitat for this terrestrial species. It may represent an accidental introduction by man, and the record needs confirmation by the collection of additional specimens from Texas before this state can be included in

PAGE 83

-7*the natural range of this species. TABLE V OEOQRAPHIC VARIATION IN THE NUltfBER OF TRUNK VEKTEBHAE OF PLETiiODON 6XHSBB0I Locality Eastern Tennessee and western North Carolina P. c. serratus Arkansas P. c. polycentratus Georgia Number of trunk vertebrae 19 20 21 22 23 P. c. cinereus

PAGE 84

-76Plethodon cinereus cinereus (CJroen) Sajamandra cinerea Green (1818: 356). Salanandra erythronota Preen (1818: 356). (Type locality, probably vicinity of Princeton, New Jersey.) Plethodon cinereus (Green). Tschudi (183R: 58). Dunn (1926: 163-80). Sauropsis erythronotus (Green). Fitzinger (I8h3: 33). Plethodon erythronotus (Green). Baird (1P50: ?85). Ambystoma erythronotum (Green). Gray (lf>50: 37). Salanandra puncticulata Valenciennes in Dumeril and bibron (1R5U: 87) . Salamandra agllis Sager (1858: U29). (Type locality, Detroit, Michigan) Plethodon erythronotus cinereus (Green). Cope (1869: 99). Plethodon cine reus cinereus (Green). Davis and Rice (18>'3: 26). Bishop (19U1: 196219). Bishop (19U3: 232-6). Grobman (19UU: 300-2). Plethodon cinereus e rythronotu s (Green). Cope (1PF9: 135). Plethodon h uldae Grobman (19U9: 136). (Type locality, hawksbiU Itountain, Vtadison County, Virginia.) Type :Dunn (1926: 165) states that the type is not known to exist. The type locality is probably in the vicinity of Princeton, New jersey. Diagnosis :A Snail Eastern Plethodon with a blacV and white rattled belly; usually with 20 trunk vertebrae; and a straight-bordered aorsal stripe in the red-backed phase. Range :Nova Scotia, New Brunswick, southern Quebec and Ontario, Canada; south tli rough the eastern United States to North Carolina, eastern Tennessee and Kentucky, Ohio, Indiana, Illinois and southeastern ...issouri (figure 13).

PAGE 85

77 Figure 13. The distribution of the subspecies of Hethodan einereua ,

PAGE 86

78 Description iThe dark, unstriped phase is usually characterized by the absence of red lipophores, the presence of anall dhite spots on the dorsum (0.07-0.2 mm in diameter), and numerous $' .aller brassy flocks on the head, back, and tail. The belly is mottled with bl&ck and yellow or white guanophores . The lateral guanophore pigment is sinil?r to that on the beU.y, The red-backed phase has sides and belly si:nilar to the dark phase. The white spots and brassy flecks on the dorsum are reduced in the area of the dorsal stripe, but are present on the head and tail. The color of the lipophores in the dorsal stripe is variable, ranging from red to yellow. The costal grooves usually number 19, the trunk vertebrae 20. Vomerine teeth range from 3 to 9 in a series. The larpest specimen examined in 5l ram. in snoutvent length. Sexual maturity is reached at about 35 ran. in snout-vent length. Plethudcn cinereus 3erratus Grobman Flethoj.on cinereus s err at us Grobman (1S J UU* 306-8). Bragg (19$2: ?Ub). Bragg (19#» 27-8). Type:CNKM 39U6U, a female, collected on Rich fountain, Polk County, Arkansas, at an elevation of ?5>00 feet, by Karl P. Schmidt and C. M. Barber, on ^arch 23, 1938. Diagnosis :A race of rT Lethodon cinereus in which the edges of the dorsal stripe are serrated at each costal groove.

PAGE 87

79 Range :Aest-central uplands of Arkansas and adjacent Oklahoma, south of the Arkansas niver (figure 13). Also reported by Bragg (195$: 27-8) from Cherokee County, Oklahoma (north of the Arkansas fiiver). Description :The dark phase is apparently rare in this form, only one of 33 specimens examined lacked the red dorsal band, in reoV-backed individuals, the serrations on the edges of the dorsal stripe are present on the body, but not on the tail. The serrations are due to the fact that the red pisrment in the dorsal stripe extends ventrolateral^ toward the top of each costal furrow. The usual absence of melanophores in this extension of the dorsal stripe makes the saw-tooth edge of the stripe conspicuous to the naked eye. The costal grooves usually number 19 » the trunk vertebrae 20. Vomerine teeth ranpe from 3 to R in a series. The largest specimen examined is U6 mm. in snoutvent length. Plethodon cinereus polycentratus Highton and Grobman rlethodoti cinereus polycentratus Highton and Crooner, (in press). Type :UF 6376, an adult male, collected ? miles northeast of Palmetto, Fulton County, Georgia, by Albert H. Highton and Richard Highton, on February 2, 195U. Diagnosis :A race of P lethodon cinereus in which the usual number of trunk vertebrae is 21 or 22 (rarely 23). liange:The Piedmont of western Georgia (fipure 13). Description :This race is similar in coloration to the typical sub-

PAGE 88

80 species. It differs in possessing a greater number of body segments. Vomerine teeth range fron 3 to 7 in a series. The largest specimen is h2 mm. in sn cutvent length. Sexual rrfiturity is reached at about 3?> mm. in snoutvent length. The Eastern Large Plethodons This group, as defined by Grobman (I9I1I4S 266) on the basis of larger size and fewer costal grooves than the other eastern plethodons, includes six species that inhabit the eastern portion of the United tes. Flethodon glutinosus is the most widely distributed and all the other spt^cies occur within its range. F. yonahlossee occurs in the Southern Section of the Blue Ridge Frovince north of the French Broad River. P. ouachitae , superficially very similar to yonahlossee , lives in the Ouachita Vountains of Arkansas and Oklahoma. The recently described P. caddoenais is known only froii the Caddo Mountains of Arkansas. P. jnrda ni includes eight subspecies, all occurring in the Southern Section of the Blue Ridge Province. The range of P. wehrlei is centered in the "Jnglaciated Allegheny Plateau Section of the of the Appalachian Plateaus Province. (See Orobman (l°UU) for an analysis of the distribution of these forme.) Two recently described species, P. dixi and P. ja cksoni are here considered races of wehrle i, because of their close morphological similarity to wehrlei and the fast that they replace wehrlei geographically. There has bean much speculation on the relationships of the species within this section of the genus. Dunn (19?6» 23) believes this to be the most primitive group of the genus, with yonahlossee the

PAGE 89

81 most primitive form. Dunn and Grobman (19hh> 276) both believe the relationship between yonahlossee and wehrlei is close. Hairston and Pope (19U8) stiggest that yonahlossee is closely related to jordani and that close similarity between some jordani and glutinosus is merely convergence, but others think that the closest relative of jordani is glutinosus . Bishop (19lil) even considered one race of jordani ( shermani ) to be racially related to glutinosus . On the other hand, Grobman (I9UI4) believed that jordani and closely related forms (now all considered r-), 115 specimens of P. ^. jordani (figure 16), 27 specimens of P. j. shermani , 72 specimens of P. ^. netcalfi, !?3 specimens of P. ^. mela-

PAGE 90

82 ventris , U* specimens of P. ^. rabunensis , 29 specimens of P. £. teyahalee, and hh speci ens of P. ^. unicoi . For a given size, the range of variation is quite similar for each form, although there is often a statistically significant difference in the variability and/or the ontogenetic rate of change in the different species. The data of Pope and Pope (1951) indicate that P. ouachitae has a similar range of variation in number of teeth as glutinosus and jordani, although the average number of teeth is slightly higher. The number of teeth in 60 P. wehrlei (figure 17) seems to be less than in the above forms, while Pope (195>0) shows that P. yonahlossee than the above forms. 39 P. caddoensis , although small in size, have even more teeth than P. yonahlossee of the same size ranee (figure 18). Except between adult wehrlei and yonahlossee , or wehrlei and c addoensis , there is considerable overlap in the vomerine tooth counts of all the Eastern Large Plethodons, and even in the case of tJ ese, only adults can actually be distinguished on the basis of this character. It is true that large series of adults of species that differ greatly in size (e.g. P. g. glutinosus and P. ^. metcalf i ) will differ in average number of vomerine teeth, but this is mainly a reflection of the difference in size between the two forms. This character obviously cannot be used as an aid in Identification, since specimens of the two species that are the same size will have similar vomerine tooth counts. Several other suppose' 1 differences mentioned in the literature are not useful in determining relationships. All, except P. vrehrlei, have a similar number of costal grooves (see table VI). Sexual dimorphism in size is present in several forms that have been critically studied. The type of lateral guanophore pigmentation varies

PAGE 91

-8326 24 2 2 2C IB 16 14 12 10 8 6 4 2 P (Pen glutinosus Insula Florida) • • • • • • • • i m i j p n •— fe«y*« » • •• » S SBS y 1 1 < tf ^
PAGE 92

6U24 22 X 20 P jordont ••••«•• • • • •B • • • ••• *•• • • •• # • ••••* •• *• 20 25 30 35 40 45 50 55 60 SNO'JTVtNT LENGTH (mm; Figure 16. Vomerine teeth of 11$ P. ^. jordani plotted against snoutvent length.

PAGE 93

-8*-

PAGE 94

86 somewhat within a species (both individually and geographically), but there appear to be no consistant differences between species. There remain, then, only a few characters that can be used to diagnose or distinguish the Eastern Large Plethodons. Each species is sympatric with, and often coexists in the same habitat with one or more of the other species without any evidence of interbreeding. As far as we know, each is a genetically distinct unit, and there is much evidence to indicate that different species have different ecological reauirements. Yet preserved specimens that have lost their pigmentation characters are often extremely difficult to identify because of the morphological similarity of most of these salamanders. P. wehrlei is distinguished on the basis of a greater amount of webbing on the toes, by having an average of one more trunk vertebra, and by possessing fewer vomerine teeth than the other species. P. yonahlossee may be recognized by its distinctive color pattern in life. It also differs from the others (except caddoensis ) in possessing a greater number of vomerine teeth. P. ouachitae has a similar color pattern to that of yonahlossee , but it has fewer vomerine teeth. I have found no character that will consistantly separate glutinosus an jordani . The chin of glutinosus is usually darker than that of jordanl , but certain populations of both species more closely resemble the other (some Trxas glutinosus have very light chins, whereas P. J. teyahalee and some P. ^. rabunensis p ossess dark chins). The close morphological similarity between all the forms, except P. wehrlei , indicate that they are a closely related group of species, with only wehrlei separated from the others by a number of characteristics.

PAGE 95

87 TABLE VI THS NUMBER OF COSTAL GROOVES AND VERTEBRAE IS THE EASTERN LARGE PLETHODONS Form No. of costal grooves No. of trunk vertebrae 15 16 17 16 17 18 19 P. w. wehriei

PAGE 96

88 The young of yonahlossee , some wehrlei, and tv/o r?tces of jordani are known to possess dorsal red spots. These disappear in adult wehrlei (except P. w. jacksoni ) and jordani , and become incorporated into the dorsal red stripe of yonahlossee . No information is yet available on the very small young of ouachitae, caddoensis , and several of the races of jordani , but young ouachitae may possess then (see Pope and Pope, 1951* Hi5). The red dorsal spots appear to be definitely absent in the young of glutinosus . fled pigment was absent in all of the newly hatched young I have examined from Florida (Highton, in press), as well as in very young specimens from many other localities. Dunn (1926: 139) records a specimen of P. glutinosus from Clayton, Georgia, that had tiny paired red dorsal spots, but this may well be a specimen of P. £. sherraani, known to occur within 10 miles of Clayton. Cope (1869* lUl) also records young specimens of glut in onus frou csves in Montgomery County, Virginia, possessing these spots, but the specimens were probably P. w. jacksoni . k reduction in the amount of melanophore pigmentation on the chin occurs in yonahlossee , ouachitae , caddoensis , wehrlei , most races of jordani (except teyahalee nnd some rabunensis ) and in some Texas glutinosus ( albagula ) . Most species possess a dark belly, but the four northern races of Jordan! ( jordani , shermani , unicoi , and especially metcalfi ) are light-bellied, as are some southern wehrlei . P. yonahlossee and 1 . glutinosus (except for its southeastern Coastal Plain representatives) attain a larger maximum size than the

PAGE 97

69 other species. P. caddoensis appears to be the smallest Eastern Large Plethodon. P. £. notcalfi might also be considered a dwarfed form. Ked pigment occurs in adult jacks oni , yonahlossee , ouachitae , Jordan! , and ghermani . Dorsal guanophores occur in glutinosus , alb agula, ouachitae, caddoensis , clems onae , and teyahalee . Lateral guanophores are usually present in all Large Eastern Plethodons, except for four races of jordani ( jordani , metcalfi , melaventris , and some shemiani ) , In summary, P. wehrlei seems to be the most distinct of the Eastern Large Plethodons, tvhile the others seem to be morphologically very si Hilar to each other, In my opinion, the hypothetical ancestor of the group might most reasonably be assumed to have been a moderatesi' ed animal with a light chin and dark belly, probably with psired red spots on the dorsum of the adult, possessing 17 trunk vertebrae, a short vomerine series, and webi»ed toes. P. wehrlei is closest to this hypothetical ancestor, although its body has become slightly elongated with the addition of an extra trunk vertebra. The chin is still light in all except glutinosus ( albagula excepted) and one or two races of jordani . The dark belly has remained in all but the four northern races of jordani and in southern wehrlei . A larger size has been attained by yonahlossee and glutinosus , while dwarfing has occurred in northern jordani (especially metcalf i ) and caddoensis . P. yonahlossee has a much longer vomerine series than the others. The degree of relationship indicated by a study of these characters would seem to indicate a phyloreny as outlined in figure 19.

PAGE 98

90 Plethodon wehrlei Group Plethodon wehrlei Plethodon wehrlei inhabits the unglaciated Appalachian Plateaus Province in southwestern New York, western Pennsylvania, extreme southeastern Ohio, West Virginia, and adjacent Virginia (where it occurs a short distance outside the Appalachian Plateaus Province) . This species is the most distinct of the Eastern Large Plethodons and appears to occupy a somewhat isolated position in the group, differing from the other species in several respects. There is usually more webbing on the toes of this species, although occasional specimens of other species, especially P. caddoensis and P. ouachitae , approach P. wehrlei in this regard. P. wehrlei is the only Eastern Large Plethodon that normally has 18 trunk vertebraej all the others usually possess 17. P. wehrlei possesses fewer vomerine teeth than any other Eastern Large Plethodon and it is the only species that has melanophore pigmentation in the oeritoneum. Two close relatives of Plethodon wehrlei from southwestern Virginia (P. dixi and P. jacks oni ) have recently been described as distinct species by Pope and Fowler (19U9) and Newman (195U). P. dixi appears to differ from P. wehrlei only in proportions and pigmentation. It would seem best to regard it as a subspecies of P. wehrlei , since it replaces the latter geographically. P. jac'^sonl , based on specimens from an adjacent county, less than 13> miles from the dixi localities, essentially differs from P. wehrlei in the retention of the juvenile dorsal red spots in the adult. Young Vest Virginia wehrlei often possess these spots, but they appear to be absent in more northern wehrlei.

PAGE 99

91 Since jacksoni does not otherwise differ from some wehrlei and dixl , it is also here regarded as a subspecies of P. wehrlei . Several writers have commented on the geographic variation in P. wehrlei . Netting (1936: 91) lists ways in which West Virginia P. wehrlei differ from topotypic Pennsylvania specimens. The former possess white spotting on the throat and chest, while in Pennsylvania specimens the white spotting is absent. The lateral white pigment is also more abundant in West Virginia material. Dunn (19?6: 135) mentions the presence of paired red spots on the dorsum of a juvenile from Bristol, tfest Virginia. Brooks (19li5* 231) reports that three adults, as well as most of the juveniles, in a series of 22 specimens from Randolph County, West Virginia, also possess dorsal red pigment. Bishop (19Ulx 238) states that none of the specimens of this species he has examined (presumably all from New York and Pennsylvania) have shown the slightest trace of red pigment. Grobman (19UU* 287) has pointed out some of the above differences between West Virginia and New York and Pennsylvania specimens, and has also suggested that southern wehrlei may attain a greater sise than northern specimens. In view of these differences in northern and southern P. w. wehrlei , this form should be further studied for other evidences of geographic variation. It is possible that the northern and southern populations should be recognized as distinct subspecies . In the possession of reduced black pigmentation on the anterior portion of the belly, southern wehrlei ore more similar to dixi and jacksoni than are northern wehrlei . In the presence of red spots on juveniles, as well as in some actults, West Virginia specimens are similar to

PAGE 100

92 jacksoni . The increased lateral white pigmentation in southern wehrlei is also paralleled by jacksoni and dirt . I have not carefully examined the pigmentation of any of the races of wehrlei in life, although a casual examination of living specimens from New York, West Virginia, and Virginia ( dixi topotypes) was made before the present study was contemplated. A comparison of the geographic variation in pigmentation characters within the species, as well as with other Plethodons is much to be desired. Pope and Fowler (19U9: 1) state that both "bronzy mottling" and "small light flecks" are present on the back of dixi. The former disappear rapidly in preservatives whereas the latter remain, although fading somewhat. Pope and Fowler state that they have occasionally observed the "white flecking" in wehrlei , but never the "bronzy mottling. 11 The white flecking is probably the same type of spot evident on the specimen figured by Bishop (19Ula: fig. U5). The red dorsal spots, present in young West Virginia wehrlei , were not observed by Pope and Fowler in a large series of paratypic dixi, including 59 juveniles. The belly of dixi is mottled (presumably with white guanophores on a raelanophore background), whereas the belly of wehrlei is usually uniformly pigmented with melanophores. In size, dixi appears to be smaller than the other races. The specimens of dixi that have been examined also appear to have proportionately narrower heads and slenderer bodies than wehrlei . Newman (195U* 12) states that the dorsum of jacksoni has "white flecks" and "silvery mottling," the latter usually disappearing wi-thin twenty-four hours after preservation. 7'ottling on the belly is also present, as in dixi . According to Newman, the dark belly pigmentation

PAGE 101

93 fades in preservatives. Since some adult wehrlei from West Virginia have been reported with red spots on the dorsum, it may be found that these wehrlei and jacks oni are very similar, if not the same form. If this were the case, the name jacksoni would be available for southern wehrlei, should the latter be shown to be a distinct subspecies (see above). A. search should be made for P. w. dixi in southwestern Virginia, for it is very unlikely that it is restricted to the vicinity of two caves only two miles apart. Populations from other localities will be important in determining the amount of variation in the form, as well as its relationships with jacksoni and wehrlei . Plethodon wehrlei wehrlei Fowler and Dunn Plethodon wehrlei Fowler and Dunn (191?: 23-U). Dunn (1926: 133-6). Bishop (1927t 117-8). Hassler (193?: 95-6). Walker (1933: 2?U). Netting (1936: 89-93). Hetting (1936: 28-30). Bishop (I9ljla: 232-9). Lachner (19U2: 263). Bishop (191*3: 281-h). Grobman (19UU: 285-7). Brooks (19U$: 231). Netting (19U6: 12). Netting, Green, and Richmond (19H6: l!?7-6o). Brooks (19U8: 2UU). Grobman (19U9: 136). Newman (195U* 13). lype :ANSP 19123, collected at Two Lick Hills, Indiana County, Pennsylvania, in Sept ember, 1911, by R. W. Wehrle. Diagnosis :An Eastern Large Plethodon with webbing between the toes on the hind foot often extending to the joint between the first and second phalanges; 18 trunk vertebrae; and a lower number of vomerine teeth (U11 in a single series) than in other Eastern Large Plethodons. Dorsal

PAGE 102

-9Urod pigment is usually absent in adults, and white pigment is usuallyabsent from the belly, except on the chin and between the anterior limbs. Range ;Prom Allegany State Park, Cattaraugus County, New York, south in the Allegheny Plateau Section of the Appalachian Plateaus Province through western Pennsylvania and West Virginia, except for the southwestern portion of the latter state (figure 20). Also recorded from saonroe and Washington Counties, Ohio, by Walker (1933* 22U), and from the Valley and Ridge Province of Highland County, Virginia, by Netting, Green, and Richmond (19U6: 157). A specimen from the Blue Ridge Physiographic Province of Nelson County, Virginia, is questionably referred to this species by Grobman (1°U9: 136). Jescription tThe pigmentation of this form has not been studied in life. Several writers have mentioned that this species is brown or bluish in appearance, probably an indication of a reduction in the intensity of melanophore pigmentation, as often occurs in the Eastern Small Plethodons. (The distinction between two types of guanophores on the dorsum given in the original descriptions of dixi and jacks oni are also reminiscent of the Eastern Small Plethodons.) Lateral guanophore spots are present on the sides, as well as on the chin of southern wehrlei. The young and occasional adults from rtest Virginia possess paired red dorsal spots (Brooks, 19U5* 231) • The costal grooves usually number 17> the trunk vertebrae, 18. Vomerine teeth range from U to 11 in a series. This form is moderate in size, the largest specimens are about 70 mm. in snoutvent length.

PAGE 103

-95Figure 20. The distribution of the subspecies of Flethodon wehrlei,

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-96Plethodon wehrlei dlxi Pope and Fowler Plethodon dixi Pope and Fowler (19U9s 1-U). Fowler (1951: 1U7-8). Type :CNHM 56510, a male, collected at Dixie Caverns, Roanoke County, Virginia, on July 11, 1°U8, at an altitude of 1170 feet, by John W. Funkhouser, Sarah K. Pope, Clifford H. Pope, Hallowell Pope, and Whitney Pope. Diagnosis :Similar to P. w. wehrlel , except for slightly smaller 3ize, slenderer head and body, the presence of white guanophore mottling on the belly, and small brassy flecks on the dorsum. ^angeiKnown only from Dixie Caverns, Hew Dixie Caverns (about UUO yards from Dixie Caverns), and Blankenship Cave (about 2.U miles east of Dixie Caverns), in western Roanoke County, Virginia (figure 20). Description ;For a description of the type series in life, see i'ope and Fowler (19U9). The costal grooves usually number 17, the trunk vertebrae, 18. Vomerine teeth range from U to 9 in a series. The largest specimen Measured by Pope and Fowler was 57 mm. in snoutvent length. Plethodon wehrlel Jacks oni Newman Plethodon jacksoni Newman (195U» 9-lU). Type :An adult male, collected at Trillium Vale, about 1 mile east of Blacksburg, Montgomery County, Virginia, at an elevation of 2100 feet, on February 11, 1950, by Walter B. Newman.

PAGE 105

97Dlagnosis :Similar to P. w. wehrlel , except that the juvenile coloration, paried red dorsal spots, is retained in the adult. Range tKnown frora the vicinity of Blacksburg, Montgomery County, Virginia, and Smith Mountain Gorge, Pittsylvania County, Virginia (figure 20). The latter record extends the ranfe of Plcthodon wehrlei into the Piedmont Physiographic Province, Pescriptiom Specimens of this form have not been examined and no information other than that published in the original description is available. Plethodon yonahlossee uroup SSost previous workers have believed that this group is very closely related to P. wehrlei and it has even been suggested (Grobraan, 19hhi ?76) that yonahlossee and wehrlei may eventually be shown to be subspecifically related. For reasons given above, it is suggested here that the yonahlossee group is actually more closely related to the g lutinos u s group than either is to the wehrlei group . The three species included in the yonahlossee group, yonahlossee , ouachltae , and caddoensis , appear to differ from the glutinosus group mainly in pigmentation characters. There are no other known consistant differences between the two groups, although there is variation in size and other characters within each group. The disjunction in the ranges of the species of this group suggests that the prototype was once widely distributed in the eastern United States, and that due to subsequent environmental changes, the

PAGE 106

96 three populations have become isolated. In external appearance, at least, yonahlossee is quite similar to ouachitae . P. caddoensis , occupying an area adjacent to ouachitae , appears quite distinct from both ouacrdtae and yonahlossee » This would seem to indicate that the two species, caddoensis and ouachitae , have been undergoing differentiation west of the i&ssissippi River for quite some time. Pope and Pope (19f>l) have suggested that there may be two other undescribed foras of this group in Oklahoma and Arkansas. P. yonahlossee is most abundant at intermediate altitudes in the soutliern Appalachians, being rare or absent in the northern-type spruce-fir forests as well as at low altitudes below 2f>00 feet (Pope, 1951: 81). In Arkansas and Oklahoma, P. ouachitae is known frora 1700 to 2800 feet on Rich Mountain, while P. caddoensis has been taken at lower elevations, from 9!>0 to 1200 feet, both P. ouachitae and P. yonahlossee occur in abundance at localities where P. glutinosus is also present, so it would appear that the two are able to coexist with P. glutinosus under certain conditions. It would therefore seem most likely that factors other than competition with plutinosus are responsible for the absence of the P. yonahlossee group at lower altitude* in the eastern United States, lossibly the higher temperatures existing at lower elevations are important factors in restricting the present distribution of these forms, but so little is known about the life hiettery, physiology, and ecological requirements of these animals that it is useless to speculate further on these matters. In several ways the yonalilossoe group appears somewhat closer to P. wehrlei than do the members of the glutinosus group. For example,

PAGE 107

-99some oaddoensis and ouachitae possess toes which are slightly webbed, approaching the condition in w ehrlei . The presence of red dorsal spots in the young of yonahlossee (and possibly ouachitae ) are similar to those in the young of southern wehrlei (but these are also presont in the young of two members of the glutinosus group, jordani and shcrraani ). On the other hand, all three species in the j'-onahlossee group have an average of more vomerine teeth than both wehrlei and the glutinosus group, and all three resemble the glutinosus group in possessing 17 trunk vertebrae, as opposed to the 18 of wehrlei . It is apparent that no one species in the yonahlossee group is closer to wehrlei than any of the ethers. Thus, it would seem that, as a group, their relationship to wehrlei is quite remote. The knowledge of the three species in this group has recently been summarized and augmented by two excellent papers by Fope (1950) and Pope and Pope (19£l). Little new inforraation on variation, ecology, and life history, other than that already discussed, can be offered here. Cirrobman (19M*: 278) has discussed the relationship between o uachitae and yonahlossee and suggested the need for further morphological as well as experimental studies to determine whether the two should be considered separate species, different subspecies, or undifferentiated populations of the same species. The work of the Popes has demonstrated that these forms differ in size at maturity, maximum size, coloration, and in the number of vomerine teeth, although there is some overlap between the two in the last two mentioned charactors. Consequently, they are certainly not undifferentiated populations of

PAGE 108

100 the same species. Whether to consider them different species or subspecies is more difficult to determine. "ttie differences between them appear, however, to be as great or greater than those between other distinct species in the genus (e.g. jordani and glutinosus j cinereus and dorsalis j cr dunni and vehiculum ) . It would seem best to continue to regard thera as separate species. Pope and Pope (1951: li.9) suggest that the relationship between caddoensis and ouaci sitae might be subspecific, but the differences in vomerine teeth, size, and color pattern would appear to indicate that they, too, have reached the species level of differentiation. The three might appropriately be called a superspecies O-ftyr, 1931s 2). Plethodon yonahlossee Dunn Plethodon glutinosus (Green). Briraley (191?' 137-8) Plethodon yonahloss ee Dunn (1917? 598-603). Dunn (1950* 130-1). Breder and Breder (1923: 15). Dunn (1926* 129-33). Bailey (1937: 2-3). Gray (1939: 106). Bishop (l«>b3: 287-90). Grobman (19lb: 287). Wood (19U7: 273-U). Hairston and Pope (19U8: 276-7). Hairs ton (19U9: 53-6). Pope (1950: 79-105). Newman (195U: 13). Type :AUNH U63U, collected near the Yonahlossee iioad, about l£ miles from iAnville, Avery County, North Carolina, at an altitude of Ii200 feet, on August 16, 1916, by E. a. Dunn and I a S. Uevin. Diagnosis :A large species, with paired dorsal red spots in the young, and a dorsal chestnut-colored stripe in the adult. Guanophores lacking in the region of the dorsal stripe, but concentrated on the sides to form a white or light gray lateral band.

PAGE 109

101 Range JNortheast of the French Broad River in the Blue Ridge Province of North Carolina, Tennessee, and southeastern Virginia (figure 21). Pope (195>0: 82) mentioned that this species had never been found east of the New Hiver in Virginia, but more recently, Newman (19£U* 13) reported two specimens from Kibler Park, Patrick County, Virginia. It may eventually be found farther north in the Blue Ridge Province in Virginia (Floyd and Franklin Counties). Description *The belly is black with few to many small white spots. The throat is usually light. The dorsal chestnut-colored stripe is made up of both red and black pigment. The red pigment appears to be somewhat similar to the red pigment of jordani and shermani , but the additional presence of black pigment gives the animal its characteristic chestnut-colored dorsum, rather than the bright red of the other forms. The black pigment is concentrated around the mucous glands in the skin. The young possess paired dorsal red spots and these often remain evident after several years of preservation, unlike the chestnut band of the adults. The reason for this appears to be the absence of melanophores in the red spots of the young, so that when the red pigment dissolves in the preserving fluid, a light melanophore-free "spot" remains. In the adult, the melanophores occur over the entire back, so when the red pigment disappears, the animal appears black. Occasional specimens have a reduced amount of red pigment on the dorsum in life, and one specimen reported by Pope (1950s 97) lacked the dorsal red pigment completely. The lateral guanophore pigmentation is so concentrated that in most individuals it forms a light gray or white lateral band.

PAGE 110

102 Figure 21. The distribution of the Plethodon yonahlossee Group,

PAGE 111

103 The costal grooves usually number 16, the trunk vertebrae 17. Vomerine teeth range from 5 to 22 In a series, a greater number than in any other Plethodon . The largest specimen examined by Pope (1950) mas a female, 85 mm. in snout-vent length. Sexual maturity is apparently reached between 55 and 65 mm. in snoutvent length. Plethodon ouachitae Dunn and Heinze Plethodon ouachitae Dunn and Heinze (1933? 121-2). Bishop (19U3 : 26972). Qrobnan (19UU: 285). Pope and Pope (1951* 129-52). Type :USNM 92U8U, an adult male, collected on the north side of Rich Mountain, Polk County, Arkansas, on May 30, 1933, by Albert A. Heinze and Dorothy A. Boyer. Diagnosis :A moderate-sized species with a dorsal chestnut-colored stripe as in yonahlossee , but with numerous white spots and brassy flecks on the back, that are not present in yonahlossee. The lateral band is often yellow in color, whereas in yonahlossee it is usually white or very light tiray. Range :Known only from Rich Mountain in Polk County, Arkansas, and LeFlore County, Oklahoma (figure 21). Also reported by Dundee (19U7* 118) from McCurtain County, Oklahoma, but Pope and Pope (1951* 13U) present arguments that the specimen on which this record is based is not ouachitae , but an undescribed form. Description :The variation in over a hundred specimens of this species has been studied by Pope and Pope (1951). I have examined only three living specimens of this species, so reference is made to their paper

PAGE 112

lou for the frequency of the occurrence of pigmentation characters. The belly is dark. The intensity of melanism on the chin is variable, ranging from very dark in a small percentage of specimens to little evident or lacking in 27%. 93% of the Popes' specimens possessed yellow or white spots on the chin and over half possessed these spots on the belly. Lateral white or yellow pigment is so abundant on the sides that in 85% of the specimens it forms a continuous band. The back has four different types of chromatophores . Black pigment is present on the dorsum and is concentrated around the glands in the skin, as in yonahlossee . Between these nlands there is a fairly uniform amount of red pigment. In addition there are two kinds of guanophores scattered over the dorsum. There are small white spots (the "white speckling" of Pope and Pope) and smaller brassy flecks (their "frosting" ) . The white spotting was lacking in 2% of their specimens and the brassy flecks were lacking in 19%. There was considerable variation in the abundance of both types of guanophores. There is also a great deal of variation in the arnount of red on the back. In 21% of their specimens they state that the back was nearly all black. After preservation, a 26 mm. juvenile exhibited two rows of faintly pink spots on the back, indicating that this form may have the juvenile red spots present in several other Raotem large Plethodons. The costal grooves usually number 16, the trunk vertebrae 17. Vomerine teeth range from U to 18 in a series. The largest specimen,

PAGE 113

105a female, is 67 mm. in snout-vent length. Sexual maturity is apparently reached between US and 55 mm. snoutvent length. Plethodon caddoensis Pope and Pope Plethodon ouachitae Dunn and Heinae. Grobman (19UU« 28$, part). Plethodon caddoensis Pope and p ope (1951: 1U8). Type :CNHM 61959, a female collected at an altitude of 1200 feet on Polk Creek Mountain, Montgomery County, Arkansas, by Sarah H. Pope, on May 9, 1950. Diagnosis :A small species with melanophore pigmentation almost entirely absent on the chin and belly anterior to the fore-legs. Dorsal spots are larger and more numerous than in P. glutinosus . It possesses an average of more vomerine teeth for its size range than any other Plethodon . The toes are often slightly webbed at the base. Range :Known only from the Caddo Mountains of southern Jtontgomery County, Arkansas (figure 21). Description :This species has not been examined in life and little more can be offered than that already discussed. Pope and Pope (195D give a description of the holotype and two paratypes in life. The costal grooves usually number 16, the trunk vertebrae 17. Vomerine teeth range from 7 to Hi in a series. This is a small species, the largest specimen of 39 examined is 52 mm. in snout-vent length. Sexual maturity is reached at about U0 mm. snoutvent length. Pope and Pope believed that all three of their specimens were immature, although two of these were U5 and U7 mm. in snout-vent length.

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106 Plethodon glutinosus Group Plethodon jordanl In 1901, Blatchley described a red-cheeked Plethodon from the Great Smoky Mountains as Plethodon Jordan! . During the next three decades, three additional related forms were described by Stejneger (1906) and Brialey (1912, 192?) as separate species, although each was obviously closely related to P. jordanl and inhabited an adjacent region. Both Dunn (1926: lh6; l5l) and Orobman (I9bk: 289) commented on the close relationship of these forms and suggested that intergrades between some of the adjacent forms might eventually be found. Brimley also recognized their close relationship and mentioned that "they hardly seem distinct enough to be full species" (1928: 23). The studies of Hairston and Pope (hairston and Pope, 19U8; Pope and Hairston, 19U8; hairston, 1950) added three new forms to the group and demonstrated that intergradation occurs between several adjacent i'oras. In 1950, hairston recognized seven subspecies of P. jordani , and another supposed member of this group (P. kentucjd) was described by attleman as a distinct species in 195l. It was later listed as a subspecies of Plethodon jordani by Schmidt in 1953, but Clay, Case, and Cunningham (1955) have shown that the specimens on which it was based are actually P. glutinosus. In spite of the fact that much has been learned about the systematic s of this group during recent years, there still remain numerous gaps in the knowledge of the distribution and relationships of many of the forms, some of which have heretofore been recorded only from a single type locality. During the summer of 1955, I had an op-

PAGE 115

107portunity to spend several weeks collecting in the southern Appalachians and a special effort was made to obtain salamanders of this group from several critical areas. In addition, a large number of plethodons of this species in the University of Florida collection, assembled since l
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108 Jordan! , melaventris , clemsonae , rabunensls , and teyahalee were regarded as dark-bellied races, while metcalfi and shermani were decribed as being light-bellied. No one has offered an explanation for the discontinuous distribution of the light-bellied populations. The new material that has been collected in the last five years indicates, however, that several other populations of Plethodon Jordan! are lightbellied. These occur in the northern Cowee '.fountains (a range lying between the Mantahala Mountains inhabited by P. j. shermani and the Balsam Mountains inhabited by P. £. metcalfi ); in the Unicoi Mountains (this population will be described as a new subspecies, P. £« unicoi , below); and in the Great Smoky Mountains. In order to study the Intensity of ventral melanophore pigmentation in an objective manner, the freshly preserved specimens collected during the summer of 1955 were compared with a standard color guide (Villalobos and Villalobos, 19U7) and the degree of darkness of the belly recorded. Since immature specimens of all the forms have much lighter bellies than adults, only the variation in sexually mature individuals was studied. Table VH shows the data on this character obtained from specimens taken in various parts of the range of Plethodon Jordan! . It is obvious that there is considerable variation in the darkness of the bellies of specimens in each population. However, with the exception of P« ^. jordani , the forms considered to be dark-bellied by previous workers, do average darker than those which were considered to be light-bellied. P. j. jordani should actually be considered a light-bellied subspecies, since the average belly pigmentation is well within the range of the other light-bellied races ( shermani and metcalfi ) The distinction between lightand dark-bellied populations is not ab-

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109 o as a Pi O EH B M as m _ « w ft. s 5^ g£ p: ^ fe 5 5 a o s d * _ ^S o m» as o 5q £2 ^ *-•' P* M •» M M M 59 EH a. El 3 S 9 o> vo co %r» oo • • • • . Q H 00 CO COS cv o o en ^ * * • O O CM ©\ «H ft CM •H
PAGE 118

110 vO

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Ill solute, but is only an average difference that can be detected by examining series of freshly preserved specimens. Nine living adult specimens from Grandfather Mountain ( metcalfj ) are almost as dark as preserved melaventris (range, 5>J mean, 7*0), and 21 living adult melaventris are slightly darker (range, 2-7$ mean U.7). Again, an average difference exists, but living specimens cannot be compared with preserved specimens, and probably additional fading may be expected after several years of preservation. Different preservatives probably give rise to other variations in amount of fading. A series of ten living specimens of metcalfj from Grandfather Mountain and ten living melaventris from Highlands were both separated into two groups of five each and one was kept in light and the other in dark conditions for eight days. At the end of this period, the four groups did not appear to have changed detectably in the intensity of their melanophore pigmentation. In a similar experiment, Grobman (195>0) found that the belly pigmentation of lightand dark-bellied races of De smog na thus fuscus was subject to changedin intensity when exposed to different light conditions. The distribution of the lightand dark-bellied populations of Plethodon jordani can now be reexamined in the light of these new data. If populations averaging 7.5 or under are considered dark-bellied and those over 7.5 light-bellied, it can be seen from figure 22 that the light-bellien populations ( jordani , metcalfj , shermani, and unicoi ), represented by hollow symbols, are not isolated from one another by intervening dark-bellied populations, as was formerly believed. If populations that average 8.0 and over, or 8.5 and over are considered light-bellied, and the remainder dark-bellied, the results would be

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112 Figure 22. The distribution of the subspecies of Plethocion jordani , Solid symbols represent dark-bellied races and hollow symbols represent light-bellied subspecies.

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113 similar. If a lighter dividing line were used, then some races here considered light-bellied would include both types. The dark-bellied populations ( inelaventris , rabunensis , clemsonae , and teyahalee ), indicated by solid symbols in figure 22, apparently occur in the southern portion of the range of Plethodon jordani . Most previous workers were convinced that the subspecies of ilethodon jordani were either isolated in the present mountain ranges or in the valleys between them during former periods of different climatic conditions (see Kairston, 19$0» for a discussion of this problem). The present distribution of the races of Plethodon Jordan! does not necessarily support either of these views. P. j. met calf i , for exa-Tple, occurs in several different mountain ranges (Black, Balsam, Cowee) that are separated by low uninhabited river valleys. It intergrades with P. j,. jordani at high altitudes in the northern Balsaa Wountains. On the other hand, it is separated from P. ^. shermani by the valley of the Little Tennessee River. P. ^. sher.iani occurs on both sides of the Nantahala River, although a few miles west of the river it is replaced by P. ^, teyahalee . P. £, melaventris and P. ^. rabunensis intergrade in the vicinity of Rabun Bald, the highest elevation between the ranges of the two races. It is apparent that some races are separated by river valleys, while others are separated by mountain ranges. Intergradation usually occurs in the mountain ranges where two forms come together, but cannot occur in some of the deep valleys that are presently ecologically unsuitable to members of this species. River valleys and high mountain ranges may thus both serve as barriers between pairs of different subspecies, and both appear to be factors important in the present distribution of the subspecies of

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-naP. jordani . The first differentiation that occured in P. jordani was probably a separation of the protojordani stock into a northern lightbellied form and a southern dark-bellied form. It is useless, at this time, to speculate when or how this occurred, since we have no idea of the geologic age of the species concerned. Later, each of these forms developed four restricted groups that can be recognized today as separate subspecies During the Pleistocene, climatic changes were frequent, and it is quite possible that at different times both river valleys and mountain ranges acted alternately as routes of dispersal and barriers to migration. The differences between the two major groups of races are not limited to belly pigmentation alone. The northern races are smaller and show a more marked sexual dimorphism in the swelling of the snout in males. Dorsal guanophores are absent in the northern light-bellied forms, although present in several populations of the southern group (clemsonae and teyahalee ) . Lateral guanophores are present in both groups, but are most abundant in the southern populations, especially rabunensis . Red lipophores appear in shermani and jordani and occasionally in other populations, especially those adjacent to sherman i. The northern races are restricted to higher altitudes, whereas the southern dark-bellied races occur at low altitudes (under 3^00 feet) as well. The southern forms are morphologically much more similar to glutlnosus , and the two species often coexist in the same habitat, while the northern, light-bellied forms are usually altitudinally separated from P. glutlnosus (Hairston, 1951).

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-11$The subspecies Jordan! and shermani are obviously closely related. Both forms have young that have similar red dorsal spots (Bailey, 1937s £-6; ^ood, 19U7a, 19h7b). Both have fairly light bellies and both possess red pigment as adults. H&irston (1950* 270) suggests that intergradation nay have taken place between the two during the Pleistocene when the Little Tennessee River Valley might have been inhabited by the two. The Cheoah Mountains, which rise to over £000 feet in elevation, lie directly between the Smokies and the Nantahalas and an intermediate population may be present there at this time. This area should certainly be searched for it probably supports a member of the jordan i group. One might expect s hermani , t eyahal ee, jordani , intermediates between any of these, or perhaps, a new form. Hairston (1900) reported intergrades between jordanl and metcalfj , m etcalfj and melaventris , and rabunensis and shermani. He is probably correct in suggesting that jordani and shermani are very closely related, rather than being circuitously connected through the subspecies metcalfj , melaventris , and rabunensis . Hairston (1950) reports intergrades between jordani and metcalfj from Hyatt Ridge, a ridge that runs south from the vicinity of Mt. Guyot in the main Smoky Mountain chain. I have examined some of his specimens (GSMMP Pm 20-29) and agree with the interpretation that they are intermediate between the two forms. A series of 10 specimens from Spruce fountain (QSMNP Pj 96; UF Bl50), 7 miles east of Hyatt Ridge in the direction of the range of metcalfj , are typical jordani (except for one specimen that has a reduction of rerl pigment on one cheek). Four miles to the south of Spruce Mountain, at Ileintooga Ridge, 15 specimens (UF 8121, 8256) all completely lack red pig-

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-116ent on the cheeks. At Chiltoskle Ridge, about half way between these two localities, six specimens (UF 81U7) are interoediate between jortteni and metcalf i . Of these, one specimen has no red on the cheeks, another has normal jordani red cheeks, and the remaining four all have a very reduced amount of red on the cheeks. The zone of intergradatlon between the two forms on Balsam .'fountain therefore appears to be very narrow (i.e. less than four miles), ring (1939s 553) reports taking the two form3 less than a mile apart without any evidence of intergradation. He does not give the exact location of his collections, but 1$ specimens from Hyatt Ridge in the GSHNP collection (Prn 1-13, 3500' j Pm lli-l$, U080') all appear to lack any trace of red on the cheeks. Two specimens (GSMNP Pj 81-82) taken at U5o > feet on Hyatt Ridge, are fairly typical jordani . Unfortunately, no further locality data are available for any of these specimens, but they indicate that both fairly typical jordani and metcalfi , as well as intergrades between the two (see above), occur on Hyatt Ridge, the same situation that occurs on Balsam .fountain, the next ridge to the east. King (1939* 55l) reports that only two of several hundred specimens of jordani from the Smokies that he examined lacked red cheeks. There is, of course, considerable variation in the amount of red that occurs on the cheeks of jordani at the type locality in the center of its range where the incidence of rei pigmentation reaches its greatest development. Some specimens have red extending across the gular region, shoulders, air even onto the fore limbs. Others have the red restricted to a part of toe cheek only (usually the upper half) . In examining the specimens of jordani in the GSMMP collection, I found only two that lacked any trace of red on the cheeks. One, PJ 22, is from Mt. Sterling

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117 at the extreme east end of the Park, and the other, Pj k$» is one of two specimens with the same number. There is no specimen labelled Pj $hj so apparently an error has been made in tagging the speciiaens. Since PJ h$ and Pj $h are iron different localities (hughes Hidge, and Long Hungry Hidge, respectively) some jj miles apart, it seems best to disregard this record. Two series collected by the writer during the summer of 19$St at Gregory Bald (UF 8125) and at Mt. Sterling (UF 8llil), at opposite ends of the Park, both show a great reduction in the amount of red on the cheeks. None of the $2 specimens from Gregory Bald completely lack red cheek pigment, but 18 (6?£) have less red on the cheeks than 138 of 1U1 topotypes examined from Indian Gap (UF 809U, 8H1I, 825U). The series from Gregory Bald may be tending slightly toward the new form, £&coi, in that several specimens have small white spots on the sides of the head and body. Of 1U specimens from tft. Sterling, two entirely lack red cheek pigment, six have it greatly reduced, and six fall within the usual range of topotypes of jordani . The specimens fro.T sit. Sterling may be approaching me teal fi , which is known from two localities on the Cataloochee Divide (GSMNP Pm 16, Sugar Tree Lick; and UF 81UU, Cove Creek Gap). Four specimens from a&x Patch akmntain (UF 8135)* on the east side of the Pigeon River, are typical metcalfi . Specimens in the GSMNP collection from many localities in the central portion of the Smokies are all similar to topotypical jordanl . Large series from sinple localities are not available, except from the type locality, so it is impossible to analyze populations from different areas in order to determine, to any further extent, whether or not

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118 any trends in the reduction of red pigment on the cheeks and intensity of ventral melanophore development are present. P. ^. cietcalfi has the most entensive range of any of the races of 1. Jordan! . It occurs from the Cowee and Balsam iSoun tains in southwestern North Carolina to southwestern Virginia. The northern-most record for netcalfi in Virginia is Burkes Garden (Hoffman and Kleinpeter, 19U8), a locality in the Valley and Ridge Physiographic Province. This race seems to be fairly uniform throughout its range. The fact that an isolated population of netcalfi occurs on Cowee Bald in the Cowee mountains seams to indicate that there was a former connection across the present low Tuckaseigee River valley with metcalfi populations in the Balsam "fountains. The Cowee fountains range to the southeast into aalaventris territory and thus do not connect directly with any other mountain range known to be inhabited by metcalfi. The young of metcalfi have not been described. A special effort was made to obtain such specimens in order to ascertain whether or not they possessed the red dorsal spots of juvenile shermani and jordani . only three young of the year were found, all from the vicinity of lit. Pisgah, in southern Haywood County, iv'orth Carolina. None had any red pigment and two, both 19 ram. in snout-vent length showed no trace of guanophores either. The other specimen, 20 mm. in snoutvent length, had numerous guanophores scattered over the back ana sides, unlike any adult metcalfi examined. Hairston (19$0) presents evidence for intergradation between metcalfi and melavantris in the southern Balsam Mountains, he compared specimens from both the northern and southern slopes of the balsams in

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119 the vicinity of Beech Gap, sad found an apparent increase in darkness of the belly from north to south. Most light-bellied populations include specimens that are as dark as some melaventris or rabunensia (see table VII), but sables from the top of the Balsam T'ountainsj near Beech Gap (UP 8109), near Wagon Road Gap (UF 8108), and Mt. Pisgah (UP 8lU5>) average only slightly darker than other samples of ractcalfj , Intergradation between these two fonts probably takes place to the south or southwest of the top of the Balsam Mountains in this area, not on the crest of the mountains. I have not examined freshly preserved or living material from any localities between Beech Gap and highlands. Until such material is available from these areas, I would hesitate to assign subspecific identifications to the populations that exist in this area, as well as to those reported from Burke, Henderson, and Polk Counties, North Carolina, and from Greenville and Pickens Counties, South Carolina, by Grobman (19Uli), Hoffman and Rubricht (195U) and Schwartz (1953* 156? 19J>U), although these workers assignel specimens from these counties to the southern dark-bellied form ( melaventris ) . These localities are tentatively indicated on the distribution map of these forms as me la ventris (figure 22). The great difficulty in distinguishing preserved P. giutinosus and the southern dark-bellied races of P. jordani casts some doubt on the correct identification of some of the records of melaventris cited in the literature. At high elevations in the Unicoi Mountains, at altitudes above U000 feet, in the vicinity of Haw I'Jiob and Johns Knob, a light-bellied population of P. jordnni was discovered by Dr. Arnold B. Grobman in 19$). In the same year, Hairston described P. ^. teyahalee , based on seven specimens from Teyahalee I'ald in the nearby Snowbird Mountains.

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1?0 The comparison of living topotypes of teyahalee with the form from the Imicoi Mountains reveals that the latter population is not teyahalee , but an undescribed form. It is distinguished from teyahalee by its lighter belly, the absence of snail red spots on the legs and the usual absence of tiny white spots on the back. Its light belly also distinguishes it from rabunensls , melaventris , and clenso nae. The absence of red pigment distinguishes it from Jordan i and shermani , and the presence of lateral white or yellow spots distlnjmishes it from metcalfi . As noted above, specimens of jordani from Gregory Bald, in the extreme west end of the Great Smoky Moun tains, show some tendency toward unicoi characters, and may represent intermediates between the two forms. P« ^. shermani , the red-legged form from the ftantahala mountains, occupies an area extending from Tellico Gap, in the northern portion of these mountains, south to the vicinity of the Georgia state line, Bailey (1937* 5) records this race from the Tusquitee fountains, indicating a geographic overlap of shermani and teyahalee (see below). Hairston (1900) reports several series of dark-bellied shermani from the periphery of its range. One of these localities is Tellico Gap, in the northern Nantahala ^fountains. Two fresh specimens (one of them immature) from this locality (UF 6178) also have very dark bellies. As postulated by Hairston, since the dark-bellied forms occur in adjacent areas, at least to the south ( rabunensis ), southeast ( taela^entris ), and west ( teyahalee ), the dark-bellied populations on the periphery of the range of shermanl may represent intergradation between shermani and these dark-bellicJ races of P. Jordan! . Bishop (19U1« 18-19) recorded a series of ten specimens from

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121 near Aquone, west of the Nantahala iijuntain range, that he regarded as intergrades between Plethodon shernani and Flethodon glutinosus . At that time, none of the dark-bellied races (except clems onae ) had been described. I have examined Bishop's specimens (CHHM 932U1-2; 93250-7) and all but one (CNHK 93?Ul) have a great reduction in the amount of red pigment on the legs, as in teyahalee . Their bellies vary from quite dark to fairly light and seem to bridge the gap between the lightbellied shermani and the dark-bellied teyahalee . It would seem best to regard them as intergrades between teyah.?.lee and shermani , but closer to teyahalee. Bailey (1937' 5) reports a series of five specimens taken about half way between Nantahala and Aquone, in which two resembled glutinosus , (presumably these had dorsal white spots), two had reduced red on the legs, and the other lacked both white and red pigment. These probably also represent intermediates between shermani and teyahalee . Specimens of P. ^. teyahalee from the west side of the Nantahala River in this area (near Junaluska Gap, UF 80?9; 8091; B17U; 8176) all possess small red spots on the legs, as well as the small dorsal white, characteristic of teyahalee (see below). Hairston and Pope (19U8) believed that specimens from looney Gap, in the southern part of the range of shermani , represent intergrades between shermani and rabunensis . Six of series of thirteen fresh specimens ( T IF P179; filP-lj) from near Mooney Gap possess a marked reduction of red on the legs, but they do not tend toward rabunensis in other characters. Their bellies are light and they appear to be well within the r-nge of eleven Wayah Bald sherrvmi (!JF PlOlj 8l06j B107) in amount of lateral yellow or white guanophore pigmentation. Much more field work is needed to definitely establish intergradetion between these two very

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122 distinct subspecies, wartof and Humphries (19£i> : 2h6) report 3hermani from extreme northwestern Rabun County, Georgia. They state that their specimens possess black bellies. These nay be intergrades ietween rabunensis and shermani . A series of eleven specimens (UF 8393) of typical shermani from Black Gap on the iacon-Clay County line and the record of Bailey (1937) for the Tusquitee ^fountains, establishes the presence of sher-vani on the west side of the Nantahala i&ver. Three specimens from Glade Gap (UF 8032), about three miles west of Black Gap, have very reduced red pigmentation on the legs, and the single adult has a dark belly. A specimen (UF 8U10) from near Tuni Gap, to the northwest of Glade Gap, on che Macon-Clay County line, seems to be typical teyahalee , yet Bailey (1937) reports shermani from .Weatherman and Tusquitee Balds, two miles further west. These records suggest slight overlap in the ranges of these two forms and this area should be more carefully studied in the future. More than half of a series of 22 specimens (UF 8301) from Jarrett Knob, on the east side of the Nantahala River, south of Hquone, have red on the legs either very reduced or absent. The belly color is variable, but would appear to be intermediate between the lightand dark-bellied forms. P. J, teyahalee was previously known only from the type locality, Teyahalee Bald, in the Snowbird Mountains. A comparison of living topotypee (UF 8166; 8168) from over hOOO feet elevation on Teyahalee Bald with specimens from Tuni Gap and Junaluska Gap, mentioned above, indicates that they are the same form. The original diagnosis

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123 of teyahalee does not distinguish it from rabunensis , and it would have to be synonymized with rabunensis were it not for the fact that most specimens may be distinguished from rabunensis by the possession of tiny (less than 0,3 mm. in diameter) white dorsal spots. These spots are similar to those of Plethcd on glutlnosus from western North Carolina, but are much smaller. Specimens of teyahalee usually possess small red spots on their legs, a ch?r»ctsr lacking in most rabunensis and glutinoeus . There are no records of teyahalee from the Snowbird fountains west of Teyahalee Bald, This mountain range extends to the west and joins the Unicoi range, with altitudes high enough to support a member of this group for most of the distance, Intergrades between unicoi and teyahalee may eventually be found somewhere in this mountain range. P. ^ e rabunensis (type locality, Rabun Bald, Georgia) is diagnosed as a dark-bellied form possessing lateral white spots. Specimens fron areas to the south and east of Rabun Bald (southern Rabun, Union, and Towns Counties, Georgia) have a much greater development of the lateral white spotting. It is apparent that Pstoan Bald specimens are intermediate in this character between southern Rabun, Union, and Towns County specimens and melaventris . Thus the type locality of rabunensis is located in a aone of intergradation. However, since the type specimen possesses lateral white spots, characteristic of the Georgia race, the name rabunensis can still be applied to this subspecies. Specimens of rabunensis with abundant white or yellow spotting are available from: Persimmon (UF 8it08), and 12 miles east of

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12U Clayton (UF 8U07), Habun County; jack Gap, Union-Towns County line (ChM, uncatalogued); and h miles east of Margaret, Union County (UF, uncatalcrued). These specimens are very similar in appearance to P. glutinosus , and the relationship between the two may actually be very close. The lack of a good systematic character to distinguish between rabunensis (and teyahalee ) and glutinosus makes the identification of living specimens difficult and preserved specimens almost impossible. The lack of dorsal spotting on most rabunensis usually serves to distinguish between the two in life, although occasional specimens of glutinosus from areas outside the southern Appalachians appear identical to it. (In certain localities in the Coastal Plain, P. glutinosus may often lack white pigment completely and cannot be distinguished from melaventris . ) Specimens of P. glutinosus from near the range of rabunensis that I have examined in life, from Tullulah Gorge, Habun County, and Potato Patch Mountain, Murray-Gilmer County line, possess small brassy dorsal spots similar to Florida glutinosus . however, all the P. glutinosus from southwestern North Carolina that I have examined possess white dorsal spots. I have collected this white-spotted form of glutinosus living in the same habitat with members of the jordani group at Soco Gap (Swain-Haywood County line), Cove Creek Gap (haywood County), U&x Patch Mountain (Haywood-iSadison County line), Cowee bald (Macon-Jackson County line), Highlands (iiacon County) in North Carolina, and at Jocassee (Oconee County), South Carolina. At the first four mentioned localities it was associated with nietcalfi , at highlands it was associated with melaventris , and at Jocassee with clemsonae. One specimen collected at rtabun Bald along with a series of topotypic rabunensis, also appears to represent this form of glutinosus. It has white

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12*dorsal and gular spots, unlike any other specimens in series of rabunensis, but is otherwise indistinguishable from them. Very likely the series of 3k specimens reported by Bailey (1937 J 3) from Blood Mountain, Union County, Georgia (IMS. 7633^-66), axe rabun ensis , since only two possessed an appreciable amount of dorsal white pigment, their throats are lighter than most glutinosus , and the lateral white pigment is concentrated into a band. This description almost certainly assigns this series to rabunensis , although both Bailey and hariston and Pope (19U8: ?75) identified the specimens as P. glutinosus . P. ^. raelaventris , an unspotted dark-bellied form, occurs commonly in the area around Highlands, tacon County, North Carolina. As pointed out by Howell and Hawkins (195U)> a small percentage (1$%) of topotypical melaventris possess lateral white spotting as in rabunensis . Both the extent and frequency of occurrence of this character in the Highlands population are not high enough to negate the usefulness of this character in diagnosing the two forms. Unspotted blackbellied specimens, probably referable to this race have been reported fro^n several localities; to the east of Highlands, but the range of this form, both geographic and altitudinal, an well as its apparent sympatric relationship with glutinosus are much in need of investigation. Two living specimens of melaventri s from the vicinity of Highlands had dorsal brassy flecks as in cleia3onae in life, indicating a low frequency of occurrence of this character in the t&ghlands population. P. ^. cleatsona e was described by Brimley in 1927 on the basis of its distinct ciorsal coloration (which usually disappears in pre-

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126 servatives), but was not again generally regarded as valid until bishop (19U1) obtained additional living specimens frora the type locality. This is a low altitude form, occurring in the vicinity of Jocassee, Oconee County, South Carolina. It apparently is very closely related to rabunensis and Tielaventrls , and differs frora Rabun Bald specimens only in the possession of small brassy guanophores that are scattered over the entire dorsum. No larger white spots are present on back as in glntinosus . Freserved specimens are usually not distinguishable from the other dark-bellied races with lateral white spots. An important problem which regains to be studied is the extent of intergradation between the lightand dark-bellied races of P. jordanl . Hairston (19$0) believed he had intergrades between melaventris and metcalfi from the top of the Balsam Mountains near Beech Gap, but it has been shown above that this population is very close to metcalfi . There is much evidence for intergradation between shermani and teyahal e e , and shermani may also intergrade with rabunensis . The greatest difficulty in studying this problem is that after years of preservation, changes take place in the intensity of ventral melanophore pigmentation. Moreover, individual variation is considerable and faiiv siaed samples should be analyzed. Hence one or two specimens may not be of much use in determining the nature of the population occurring at a particular locality. Except for jordanl and shermani , which have melanophore gaps in places where the red pigment is located in life, most faded preserved specimens must be identified almost entirely on the basis of locality. This adds to the difficulties for it means that fresh specimens from new localities are of limited value unless they can be compared with other appropriate living specimens. The

PAGE 135

127 fact that P. glutinosus is often almost, indistinguishable from some of the dark-bellied forms (especially rabunensis and teyahalee ) further compounds the difficulties in conducting systematic studies in the group. P. glutinosus is easily distinguished from the northern light-bellied races of P. jordani , but appears to be more similar morphologically to the southern dark-bellied races than the two groups of races of P. jordani are to each other. There is evidence of intergradation between the darkand light-bellied races of P. jordani and they replace each other geographically, indicating a subspecific relationship. P. glutinosus , on the other hand, is known to occur syrapatrically with several of the races of jordani without any evidence of interbreeding, according to King (1939* 55l), P. glutinosus and P. ^. jordani occur together on Gregory Bald, and metcalfl , clemsonae , and rabunensis have been collected in the same locality as glutinosus by the writer (see above) . liairston (19£l) also reports slight altitudinal overlap between glutinosus and shermani . On the basis of his field experience with unicoi, Dr. Grobman (personal coramunication) believes that this form is altitudinally separated from glutinosus on the north slope of the Unicoi Mountains in the vicinity of Stratton Gap. As yet, there are no data on this subject for teyahalee. In general, the northern light-bellied races are altitudinally separated from I\ glutinosus (Kairston, 1951) and the localities in which they occur together may happen to be in the narrow zone of overlap between the two species. However, I feel that extreme caution should be exercised in interpreting the type of negative data presented by Hairston. In yy experience, a locality that has been intensively collected has often yielded a species of Plethodon that was previously

PAGE 136

126 believed to be absent. P. glutinosus is often much less common than P. Jordan i where the two occur together, and can easily be overlooked, even though a thorough search is made. It seems to me very unlikely that there is competition between the two for space, food, or some other environmental factor. It seems much more likely that there are other limiting environmental conditions that restrict the alti
PAGE 137

129 and southern P. jordani is the result of parallel evolution rather than close relationship, as suggested by hairston and Pope (19ii8s 276). Grobraan (19uU) even places tne two species in separate groups within the section of the genus containing the Eastern Large Plethodons, but this interpretation is not followed here. Plethodon kentucld, described by Lfittleman (19$1: 10$) as a member of the jorda ni group from southwestern Kentucky, has been shown by Clay, Case, and Cunningham (1955) to be based on specimens of P. glutinosus . I have examined the type series and agree with their interpretation, ^any of these specimens still possess traces of the large dorsal, lateral, and gular spots characteristic of glutinosus . Most of the type series no longer is tagged so it is impossible to determine which specimens were designated as types by {.tittleman. CSNH 1$?1K was designated as the holotype of P. kentucki , CSNH 1521B the allotype, and If^lC-J as para types. There are now only three specimens in a series of 16 ("ittletnan designated only 10 as types) that are tagged ("H, n "I," and "J") and there is no specimen of the sex and measurements given for the holotype. It is therefore impossible to determine which of these specimens Mittleman designated as the holotype. This may become an important problem in allocating the name in future systematic work on raciation in P. glutinosus , but, for the present, it does not have to be considered further. Plethodon jordani jordani Blatchley Plethodon jordani Blatchley (1901: 762). Dunn (1920: 131). Dunn (1926: 1U5-6). Brimley (1927: 10). Bishop (1928: 159). Pope (1928: 2-3)

PAGE 138

130eller (1931: 29). Bailey (1937? 6). King (1939: $5l-j). Bishop (19U3* 261-1*). Grobman (19UU* 29U). riood (I9l*7a: l8£-8). hairston and Pope (191*6: 266-78). Deevey (191*9: 1367-9) Flethodon jordani jordani Blatchley. ii&irston (19$0: 271). hairston (1951: 266-7U ). Type:I>unn (1926: lhf>) states that the type specimen was originally in the collection of W. S. Blatchley, but that it has been destroyed. It was collected at Mb, Collins or Indian rass, Sevier County, Tennessee, by L. £. Daniels. Diagnosis tA light-bellied subspecies of Ple thorion jordan i possessing red cheeks in the adult and red dorsal spots in the young. Dorsal and lateral white spots are usually absent, except in juveniles. Range :The Great Smoky Mountains of North Carolina and Tennessee (figure 22). The eastern-most records of this race are -hite Hock Mountain (now »t. Cammerer) near the town of Mb. Sterling, haywood County, Worth Carolina (Dunn, 1926: 11*6) and on the east slope of Mb. Sterling (UF 8H-1). There are specimens in the GSmTJP collection representing many localities along the crest of the Smoky Mountains (North Carolina-Tennessee state line) from Mt. Canraerer to Gregory Bald, at the west end of the Great Smoky ^fountains National Park. This form apparently occurs at higher elevations throughout the Park, except in the southeast corner where it is replaced by metcalfi. Intergradee between the two forms are known from Hyatt Ridge (Hairston, 1950« 263) and Chiltoskie ilidge. The lowest altitude from which this race has been reported is 2$00 feet on the slopes of Mt. LeConte, Sevier County, Ten-

PAGE 139

131 nessee (Bishop, 19U3» 261 ). DescrigtionJThis form is characterized by its bright red cheeks. The dorsal and lateral surfaces of the animal are black and the underside is grayish, varying froa fairly light in the young to somewhat darker in the adults. The young possess dorsal red spots which are usually arranged in two rows, one on either side of the midline of the anterior half of the back. These are present on most individuals under JO mm. in snout-vent length, but traces of the red dorsal pigmentation may occasionally occur on specimens up to 50 ram. in snout-vent length, liost young individuals also have red cheeks. The red pigment of the dorsal spots of the young appears to be the same as the red cheek pigment in adults. Both occur in gaps in the melanophore background. There is considerable variation in the amount of red pigment on specimens from the central portion of the Smokies, where large series are available. Some have the red restricted to the upper half of the cheeks, most have the red confined to the sides of the head, but in a few, the red pigment extends ventrally onto the chin, posteriorly on the shoulders, and 10/o of the specimens from Indian Gap also have small red spots on the front legs. Most young specimens have small brassy flecks on their eyelids, but these disappear in older individuals. The young often possess small lateral spots on the anterior sides, but these also seen to disappear with age, so that most adults lack guanophore pigmentation, except in the iris of the eye. More than half the adults examined possessed brassy iridic guanophores. The costal grooves usually number 16, the trunk vertebrae 17. Vomerine teeth range from h to 12 in each series, increasing in number

PAGE 140

132 with larger size (see figure 16). This form is moderate in size, adults rarely attaining a length over 70 mm. in snout-vent length. Sexual maturity is reached between h5 and 55 mm. in snoutvent length. Flethodon jordan i metcalfi Brimley Ambystoma jeffersonlanum (Oreen). Phoads (1895: U02). Brimley (1912: 136). Flethodon metcalfi Brimley (1912: 138-9). Dunn (1917: 60U-6). Fowler and xlunn (1917: 23). Dunn (1920: 131-2). Breder and Breder (1923: 1U-5). Bishop (192U« 96). Tope (192U: 2). Brimley (1926: 79-80). Dunn (1926: lli8-5l). Noble (1927: U). Bishop (1928: 160). Walker (193U: 190). Bailey (1937: 6-8). Rankin (1937: 176, 180). Green (1939* k9)» King (1939: 553-U). Brimley (19U0: 6-7). Bishop (19U3: 26U-6). Green and idchmond (19U*: 256). Orobman (19UU: 289-92). Snyder (19U6: 17k). Hairston and Pope (19l8i 266-78). Hoffman and Kleinpeter (19l#a: 107). Hoffman and Kleinpeter (l9U8b: 605). Deevey (19U9: 1367-9). Hairston (19^9* 53-5). Bogert (1952: 16-30). Plethodon metcalfi metcalfi Brimley. Mittleman (19U8: U16-8). Flethodo n jordani metcalfi Brimley. iiairston (1950> 271). Hairston (1951: 266-7U), T ype *USNJ4 U9682, collected near Sunburst, haywood County, North Carolina, at an altitude of about 3500-l»000 feet, in late May, 1912, by Franklin Sherman and C. S. Brimley. Diagnosis 1A light-bellied subspecies of Flethodon jordani lacking guanophores and lipophores.

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133 Range ;From Cowee Bald, Jackson-Macon County line, Hyatt and Heintooga Ridges, Swain County, and Cataloochee Divide, Hsywood County, North Carolina, southeast in the Balsam Mountains to !.ft. Pisgah, thence northeast in the Blue Ridge Mountains of western North Carolina and adjacent Tennessee to rhite Top Mountain and Mt. Itogers in southwestern Virginia (figure 22). Also recorded from Burke's Garden, Tazewell County, Virginia, by Hoffman and Klednpeter (19b8bt 6o5). Description :P. ^. metcalfi is the only light-bellied subspecies that lacks both guanophores and lipophores in the adult. This race has the most reduced ventral melanophore pigmentation of any subspecies of P. jordani . The sides and back often have a reduction of black pigment resulting in a lighter over-all appearance than any of the other subspecies. One small juvenile from near Mt. Pisgah (UF 6108) possessed small dorsal guanophore flecks in life, ^evy large specimens occasionally possess a small amount of white pigment scattered through melanophore gaps on the sides and/or belly, but these guanophores are never concentrated into groups to form white spots as occur regularly in teyahalee , rabunensis , and unicoi . Guanophores are usually absent from the iris. The costal grooves usually number 16, the trunk vertebrae 17. Vomerine teeth range from U to lU in each series, similar to 1. ^. jordani . This is the smallest race, the largest specimen examined is 66 mm. in snout-vent length, but few adults attain a length over 60 mm. Sexual maturity is reached between hO and \i$ mm. snoutvent length.

PAGE 142

13UPlethodon Jordan! shermani Stejneger Plethodon shermani Stejneger (1906 j 559-62). Brimley (1912: 138). Dunn (1920: 131). Dunn (19?6: 1U6-8). Bishop (1928: l6o~l). Pope (1928: U-5). Bailey (1937: U-5). Grobman (19Uit 29U-6). P lethodon glutinosus sheraani Stejneger. Bishop (19U1* 18-9). Bishop (19U3« 253-6). wood (19U7b: 188). Wood (19l*7c: 27>U). Plethodon ahermani shermani Stejneger. Pope and Hairston (19U8: 106-7). Hairston and Pope (I9b8: 27U-5). Deevey (19U9: 1367-9). Plethodon jordani shermani Stejneger. hairston (1950: 271). Hairston (1951: 266-7U). Martof and Humphries (1955: 2^6). Type *USNM 3621U, collected near '.Vayah Gap (on the Franklin side of the Gap), Macon County, North Carolina, on August 2U, 190U* by Franklin Sherman. See Brimley (1912: 138) for correction of originally designated type locality. Diagnosis :A light-bellied subspecies of P. jordani with large red spots on the legs. Range :From northwestern Rabun County, Georgia, north in the Mantahala fountains of liiacon County, North Carolina to Tellico Gap; west to Tusquitee and Weatherman Balds, Cherokee County, and Black Gap, Clay County, North Carolina (figure 22). Description :The brilliant red legs of typical specimens of this race make it one of the most strikingly beautiful North American salamanders. The dorsum is black and the underside light in the young, becoming grayish in the adult. There are usually a few small lateral yellow or white guanophore spots, most numerous just behind the front limbs. The

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-135young possess dorsal red spots similar to those of F. £. jordani young. The red on the legs of the adults is usually most abundant on the proximal half of the limbs. In specimens vdth the red on the legs reduced, it is generally reduced on the hind limbs to a greater extent than on the front legs. Populations on the periphery of the range often have dark bellies and reduced red on the legs. The costal grooves usually number 16, the trunk vertebrae 17. Vomerine teeth vary from 3 to 13 in a series and the ontogenetic increase in number is veiy similar to jordani . A moderate sized form, with dimensions similar to P. ^. jordani . Plethodon jordani unicoi , new subspecies Type ;UF 8162, an adult female, collected along the trail to Haw Knob from ?ihigg8 Meadow, North Carolina (Graham County )-Tennessee (Monroe County) State line, between $000 and 5300 feet, on August 17, 1955, by Richard M. Johnson and Richard Highton. Diagnosis :A light-bellied subspecies of Plethodon jordani with lateral white guanophore spots. R ange ;Known only from Whiggs Meadow, haw Knob, Little haw Knob, Johns Knob, Stratton Gap, and Beech Gap, at elevations over UOOO feet in the Unicoi fountains along the Tennessee-Worth Carolina State line (figure 22). g atyp tatIrabu SamtQrj wit '-'•\-~ arc Goaa% , taaei wa ^tste line: UF 8162 (18) same data as holotype; UF 8030 (l5), Haw Knob frail} UF 8303 (26), .?higg8 Meadow; UF 8306 (8), Little Haw Knob; UF 8275 (3),

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-136between Mud Flat and Y/higgs iueadow; UF &312 (U), UF 6316 (3), Johns Knob; UF 8263, 0.7 miles south of Stratton Gap; UF 6j05 (5), 0.9 miles north of Stratton Gap; iJF 8l6l (7), 1.3 miles north of Stratton Gap; and UF 8213, 621k (2), Beech Gap. Tennessee, Monroe County: UF 8310 (2), 1 nile west of Stratton Gap. Description of the type in life :Snout to anterior angle of the vent, 5U mm.; snout to posterior angle of the vent, 58 mra.j total length, lUj mm. 16 costal grooves. 7 vomerine teeth on each side, ilelanophore gaps on chin and between front legs. Numerous si?all white spots on sides (now faded after preservation). No guanophores on dorsum, in eye or on eyelids. Variation ( based on 26 specimens examined in life);Nine (3$%) possess small dorsal guanophores (as in teyahalee ); one has two small red spots on one front leg; 13 (U5%) possess small brassy or white flecks on the eyelids; and all but one possess lateral guanophores. The average belly sntation of this series may be compared with the other races of P. ^ordani by referring to table VTI. The costal grooves usually number 16, the trunk vertebrae 17. Vomerine teeth vary from 5 to 12 in a series. * small race, the largest specimen examined is 59 mm. in snout-vait length. Sexual maturity is reached between hO and US mm. Plebhodon jordar.i melaTentri s Pope and Hairston Plethodon metcalfi Srimley (1912: 139). Fope (1928: h) . Chamberlain (1928: 51). Coker (1939: 10-1). Chadwick (19U0: 50). Bishop 19u3« 26U-6).

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137Plethodon clemsonae Brimley. Grobman (19U1;* 293-U). Plethodon shermani melaventria Pope and Hairs ton (19U8* 107). Hairston and Pope (19U9* 266-78). Hairston (19U9» 53). Deevey (19U9s 1367-9). Gordon and Smith (191*9* 175). Plethodon metcalfj clemsonae Brimley. Mittleman (19U8: Ul7). Plethodon jordani melaventris Pope and Hairston. Hairston (1950: 272). Gordon (1952: 679). Schwarta (1953* 156). Howell and Hawkins (I9&s 32-6). Howell (1951 J U2-3). Hoffman and Kubricht (195U*. 193). Schwarta (195U* 296-8). Type iCNHM U761U, an adult male collected at Highlands, fciacon County, North Carolina, at an altitude of 3800 feet, on July 27, 19U6, by Alexander Pope. Di agnosis *A dark-bellied subspecies of Plethodon jordani that typically lac' s both guanophores and lipophores. Range iSpecimens assigned to this form have been reported from Rwannanoa, Buncombe County (Grobman, 19UU, fig. 5), and Burkemont fountain, Burke County (Hoffman and Hubricht, 195U) south through Polk, Henderson, Transylvania, southern Jackson, and southeastern Macon Counties, North Carolina, and northern Greenville, Pickens, and Oconee Counties, South Carolina (figure 22). Description ;The variation in a series of lj8 topotypes has been studied by howell and Hawkins (195U* 32-6). 61 living specimens I have examined from the vicinity of Highlands usually lack iridic guanophores (92%), dorsal guanophores (97^), and lateral guanophores (82#), but all specithat possessed these guanophores had them present in very reduced number. In most cases they could be seen only with the aid of a microscope. The

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138 belly is dark (see table VII), but there is often a reduction of melanophores on the chin, and the chin is almost always much lighter than the belly. The costal grooves usually number 16, the trunk vertebrae 17. Vomerine teeth range from h to 17 in a series, tending to increase slightly in number with the size of the animal, as in P. ^. Jordan! . This is a large racej the maximum size recorded is 78 nim. in snoutvent length (Howell and Hawkins, 195&: 33). Sexual maturity is reached between 5>0 and £$ mm. in snoutvent length. Plethodon jordani rabunensis Pope and Hairs ton Plethodon shermani Stejneger. Howell (1909: 131). Plethodon glutinosus (Green). Bailey (1937: 3). Plethodon cl emsonae Brimley. Grobman (l9hU: 293-U). Plethodon shermani rabunenais Pope and Hairston (19U8: 106-7). Hariston and Pope (19U8: 27U-5). Hairston (19^9: 53). Deevey (I9h9: 13679). Plethodon metcalfi clemsonae Brimley. Vdttleman (19u8: U17). Plethodon Jordan 1 rabunensis Pope and Hairston. Hairston (1950' 272). Type :CNHM h7697, an adult female, collected at Rabun Bald Mountain, Rabun County, Georgia, on August 3> 19U6, by members of the C. H. Pope family, at an altitude between Ii200-b600 feet. Ringe :Known from Rabun, Union, and Towns Counties in northeastern Georgia (figure 22). Also reported from Habersham County, Georgia, by Wittleman (19U8: hl8). Specimens from the type locality appear to be intermediate between rabunensis and melaventris.

PAGE 147

139 Diagnosis :A dark-bellied subspecies of Plethodcm jordanl with lateral yellow or white spots. These may be so large that adjacent spots unite to form a continuous lateral band. Description :Pope and Hairston (19h8: 107) state that in their original series, the chief variation was in the amount of leteral white spotting. ihey state that 1$% of their specimens lacked the white lateral and cheek spotting characteristic of this race. However, most of their specimens were collected in 1939 and had been preserved for a number of years. They apparent^,' examined only ten specimens in life. I have examined living specimens from several localities on the north slope of Habun Bald Mountain and a large proportion of the specimens possess a small aaourt of lateral and cheek white guanophore spotting. It ia, however, poorly developed compared to specimens of this form from the south and west of the type locality. In the latter, the lateral guanophore pigment is very abundant, and often adjacent spots coalesce into a lateral "band" of yellow or white pigment. A small proportion, 2 of Hi (lh£) of a series of specimens from Brasstown Bald also have small dorsal guanophore spots, as in teyahalee . The costal grooves usually number 16, the trunk vertebrae 17. Vomerine teeth range from $ to 13 in a series. This is the largest race, adult specimens approaching the size of glutinosus a nd yonahlosgee . The largest specimen I hr.ve examined, from Brasstown Bald, is 82 ram. in snoutvent length. Sexual maturity is probably reached between 55 and 60 ram. in snoutvent length.

PAGE 148

IliO Plethodon jordani teyahalee Hairston Ple thodon metcalfi Brimley (191,?: 139). Plethodon glut irto sus (Green). Bailey (1937: 3). Plethodon Jordan! teyahaleo Hairston (19&): 269-72). Type :UJfiS 100807, an adult male, collected on Teyahalee Bald, GrahamCherokee County line, North Carolina, at an altitude of U525 feet, on August 23, 19U9, by Melson G. Hairston. Diagnosis ;A dark-bellied subspecies of Plethodon Jord an! with lateral white or yellow spots, usually with small red spots on the legs, and with very small (0.1-0.3 ma.) white spots on the dorsum. Range :Known from Tuni Gap, Clay-ilacon County linej near Junaluska Gap, Macon-Cherokee County line; and from Teyahalee Bald in the Snowbird Mountains, along the Cherokee-Graham County line, all in North Carolina (figure 22). Intergrades between teyahalee and shermani occur in the vicinity of Aquone, on the east side of the Nantahala River, in aeon County, North Carolina. Description :This race is distinguished on the basis of the presence of small dorsal white spots. These are of a different color than those of clerasonae, and resemble the dorsal spots of glutinosu s, except that they are much smaller. The mean size of ten spots measured at random in 11 living specimens range from 0.12 to 0.21 ram., with an average of 0.16 mm. The site of these spots is sraaller than in any sample of KLethoUon glutinosu s I have examined. The red spots on the legs are usually not more than 0.5 mm. in diameter. The lateral spots may be yellow or white, and are not as large as those of rabunonsis from the

PAGE 149

Ha southern portion of its range, although they are comparable in size to those froi (AbfBi i^ald. The same type of spotting that appears on the sides nay also be present an the chin. The costal grooves usually number 16, the trunk vertebrae 17. Vomerine teeth vary from U to 12 in a series. The largest specimen ex?aiined, from the type locality, measures 76 mm. in snout-vent length. Sexual maturity is apparently reached between li5 and 55 mm. in snoutvent length. Plethodon jordani clemsonae brimley Plethodo i clemsonae ^rimley (192?: 73-5). Brimley (1°U0: 7). Bishop (l9Ult 20). Bishop (19ii3* 239-U2). Grobman (19UM 293-U). Plethodon shermani clemsonae Brimley. hairston and Fope (19U8: 27U). liairston (19U9* 53). Deevey (19U9« 1367-9). Plethodon Jordan! clemsonae Brimley. liairston (1950: 272). Bairston (1951 J 266-7U). Type :USNM 838U9, a male, collected at Jocassee, Oconee County, South Carolina, at an altitude of 1200-1500 feet, by J. A. Berly, on April 8, 1927. Diagnosis iA dark-bellied subspecies of Plethodon jordani possessing numerous small dorsal brassy guanophores. Range 1 Knorn only from the vicinity of the type locality, Jocassee, Oconee County, South Carolina (figure 22). Description sI have examined onjy six specimens of this subspecies in life. There is apparently considerable variation in the frequency of

PAGE 150

1U2 the brassy dorsal spotting. In some specimens it virtually covers the dorsum, while in others it is somewhat reduced. This pigmentation contrasts markedly v/ith that on the lower sides, which consists of small guanophore spots. In the specimens I have examined, these were white in color. Presumably this form is similar to melayyntris in size, number of vomerine teeth, and costal grooves. Dr. Albert Schwartz of the Charleston Museum has completed a study of variation in this form and his information should be available in the near future. Flethodon glutinosus This species is the most widely distributed of the (^astern Large Plethodons. Workers have long realized that P. glutinosus is quite variable throughout its range. For example, bishop (19Uj« 19) states that he believed each of several salamanders, including Plethodon glutinosus , to be " a complex of species or subspecies requiring additional study before its components can be properly delimited." During the l?st twelve years, three populations of Plethodon glutinosus have been described as separate subspecies. The first, P. g. albagula , was described by Grotman (19hli) based on a series of light-chinned examples frora the region of the Dalcones Escarpment of Texas. In I9U9, Allen and Keill described the populations occurring in southeastern Georgia and peninsula Florida as another subspecies, P. j*. frobmani . In 1951, Mittleman described P. £. chlorobryonis , based on material from the Coastal Plain of North Carolina. The range of each race, as delimited by the above authors, is quite small, and the remainder of the species, including animals from every physiographic province in which this

PAGE 151

-11*3specios is known, is currently assigned to the typical subspecies, P. £. glutlnosus . The difficulties in studying geographic variation in P. glutlnosus are mainly due to the fact that much of the apparent variation in this species, as well as its close relatives, involves pigmentation characters, ttost of these pigments are altered by preservatives so that a study of living specimens is necessary in order to obtain accurate information on geographic variation. The recent studies of ilairston and Pope on P. jordani have shown the value of the examination of livin, specimens from the entire range of a species. The same method was attempted by the writer with t_, glutinosus, but much greater difficulties are involves in dealing with a species th?.t occurs over most of the eastern United states, from new York and Illinois south to rlorida and Texas, iierpetologists from every state in the range of the slimy salamander were contacted and informed of the study and a request was made for living specimens from each region so that animals from the entire range of the species could be compared. A great many interested persons responded to the request and sent me living specimens. As a result, during the last four years, I have examined slimy salamanders from all but two of the states in which it is known to occur, however, there remain many regions from which I have not had a sufficient number of specimens to obtain a clear picture of the variation within its glutin osus population. Several collecting trips have been made by the writer in order to fill in some of these gaps, but there are so many areas, including whole physiographic provinces, from which data on series of living specimens are not yet available, that the '-nowledge of the systematics of this

PAGE 152

na, species is still fragmentary. It is felt, however, that some progress has been made and the results, admittedly tentative and incomplete, are presented below. In comparing living specimens from various parts of the range of this species, it is apparent that there is much individual variation within populations, but very few consistent differences are present between aninulE from different regions. I have been unable to find any additional characters that vary geographically, other than those previously discussed by Grobman (19Ui), -sill (1°U8), Allen ana Weill (19k9)t or fctittleman (195>1). All of these characters have been evaluated using both living and preserved specimens. £• £• albagula is distinguished on the basis of the reduction of the melanophore pigmentation on the chin. As noted by Brown (19$0), the characteristic light chin of albagula may occur in few populations of giutinosus in other parts of the range. Specimens with lirht chins seem to be especially abundant in the Blue Ridge Province of Virginia. The data that Grobman (19l*h: ?P 3) presents, indicate, however, that the great majority of P. glutino3us , other than those from Texas, mry be distinguished from albagula on the basis of chin pigmentation. A study of additional material from Texas, not 6een by Grobman, raise additional problems in connection with the Texas populations that make it difficult to determine the proper systematic position of albagula . These data will be discussed below. P. g. grobman i was diagnosed on the basis of the color and size of its dorsal and lateral spots, lower costal groove count, and small size. The subspecies, P. £. clilorohryonis was characterised by a

PAGE 153

-Induction of dorsal white spots, the color of the lateral spots, bysmall "orange-gold dcts" on the venter, reduced number of vomerine teeth, and smaller average size than other populations of ?. glutinosus . Tburow (195l) has shown that the "orange-gold dots" mentioned by r.'ittleman are actually hedonic glands found in male glutincsus from its entire range, and the writer has observed the color to change in individuals that were kept alive for some ti^ie, presumably due to the accumulation of the secretion of the gland. These glands sometimes occur in females as well as males, and in both sexes the color ranges from clear or colorless to bright orange. There is seasonal variation in the colcr of these glands, as well as individual variation within a population collected at a single time, but I have seen no evidence of geographic variation in this character. The range of P. £, chlorobryonis was indie ted by c&ttleraan (19?1) to include most of the Worth Carolina Coastal Plain below the 100foot contour line. He also believed that the range of this subspecies almost certainly extends into South Carolina. :.^pecimens that he considered intermediate with P. £. glutinosus are from the lower Coastal Flr>in of adjacent Virginia, from Granville and Kake Counties in the Piedmont of North Carolina, and from Charleston and Berkeley Counties in the lover Coastal Plain of South Carolina. Schmidt (19^3: }h) has recently restricted the type localities of two synonyms of P. r lutinoems , Salamandra variola ta Gilliams (1818), and oalamandrfi cylindracea Harlan (18??) , to Charleston, South Carolina. Salamandra variolata was originally described as inhabiting the "southern states," but Gilliams (I8UB1 k6l) states that the specimens on

PAGE 154

Ili6 which the description of S. variolat a were based were received from the "Florida Party." Since they were apparently collected by a person residing in Florida, or an expedition to Florida, this state would appear more likely to be the type locality of S. variolate than South Carolina. harlan (182$) states that the specimens on which he based the description of Salamandra cylindracea were sent to him by Dr. Blanding of Camden, South Carolina, who collected other herpetological specimens referral to in the sare paper in the vicinity of Camden, bouth Carolina. Camden would therefore be a better choice for a restricted type locality than Charleston, South Carolina. Allen and Neill (19l9) give the range of P. £. grobmani as "from northern Emanuel and Screven counties, in the Georgia Coastal Plain, southward to Pinellas ?nd Hillsborough counties, Florida, eastward to the Savannah River in Georgia, and westward approximately to the eastern border of the Dougherty Plain in Georgia." This would include Savannah, Chatham County, Georgia, within the ran^e of P, £. grobmanl . Indeed, specimens from adjacent Effingham County, Georgia, are designated as paratypes of this form. If this is the actual range of grobmani , then this name is preoccupied by Salamandry albopunctata Vallenciennes (in Dumeril and Bibron, lB5k), type locality, Savannah, Georgia. Vallenciennes also described Galanan jra elungata in the same paper in which he described S. albopunctata , but failed to designate a type locality. This form was also probably based on specimens from the southeastern Coastal Plain, but it will probably never be necessary to designate a type locality for it, as there are several earlier names available for southeastern Coastal Plain P. glutinosus .

PAGE 155

-117There are, then, six names available for the Plethodon glutinosus of the southeastern Coastal Plains Salamandra variolata Gilliams, 1818 (type locality, probably Florida). Salamandra cylindracea Harlan, 182$ (type locality, probably Camden, South Carolina). Salamandra albopunctata Valenciennes, 185U (type locality, Savannah, Georgia). Salamandra elongat a Valenciennes, 18£U (type locality not designated) . Plethodon glutinosus g robman i Allen and Weill, 19U9 (type locality, Half-mile Creek Swamp, about one-half mile northeast of Silver .Springs, f&rion County, Florida.) Plethodon glutinosu s chlorobryonis Mittieman, 19£l (type locality, 13 miles north of New Bern, along U. S. Highway 17, Craven County, North Carolina). The subspecific identity of several Coastal Plain populations of F« glutinosus must be determined in order that the name Salaaandra variolata , which has priority over all the other more recent n.^ees, can be correctlv assigned, in order to attempt to correctly apply the above names and to determine the number of valid subspecies of glutinosus in the Atlantic Coastal Plain, living specimens have been obtained from the type localities, as well as from many other places in the region. The characters studied rdll be analyzed separately below. Nu mber of costal grooves t£>oth Allen and Weill (19U9* 112) and *«iittleraan (1951s 11) state that in southeastern Coastal Plain, P. glutinosus possesses one or two costal grooves less than more northern populations,

PAGE 156

Ill* which usually have 16 costal grooves. Data on the number of costal grooves and trunk vertebrae in several geographic sarnies of P. glut ino sus are given in table VIII. To avoid counting the same characteristic (number of body segments) twice, costal groove co-jnt3 are not included in the table for any of the specimens for which vertebral counts are listed. The modal number of trunk vertebrae in each sample is 17 and the corresponding number of costal grooves is 16. The only variation appears to be in the percentage of individuals with one segment more or less than the mode. In the Texas sample, 11 (8£) possess l p vertebrae, and there are none with 16 vertebrae. On the other hand, in the southeastern Coastal Plain, there is a slight tendency to possess one less segment than the modal number. Radiographs indicate that 9 of 22 (Ul£) specimens from peninsula Florida have 16 trunk vertebrae, but this is apparently an unusual sample, for only 20 of 1$0 (13^) additional specimens have 1? costal grooves. These slight differences may indicate the beginning of differentiation in the populations concerned, but the differences are not of the magnitude claimed by previous workers. SiseiBoth Allen and Meill (19U°: 112) and 'Ittleraan (l95l« 111) noted the reduction in site in southeastern Coastal Flain glu tine bus . 3iee is a difficult character to analyze in cold-blooded vertebrates, unless large series of specimens are available, so that an accurate estimation of maximum si?e or size at maturity can be obtained. Another difficulty results from the fact that the average eiae of mature animals varies from season to season, for as the group of immature animals begins to mature, the average size of sexually mature adults will be lowered. As growth continues, the average siae of mature animals will increase

PAGE 157

Ih9 -

PAGE 158

1$0 throughout the remainder of the year. That this factor is important in Florida p. glutinosus was shown by the writer (in press). As yet we have very little knowledge of the life history cf P. glutinosus in other parts of the range and have no basis on which to make comparisons of the average siae at .'maturity In Florida, in the fall, mcst specimens ovor U5 mm. in length are nature, whereas in the summer, most under 55> nm. are iimature. I have examined mature spoci .ens of ?. glutinosus from the southern Appalachian fountains that were between hS and 5>0 ma. in snout— vent length. At present, there is no evidence to support the contention that there is a difference in size at maturity between Florida and Appalachian mountain P. glutinosus . This does not rule out the possibility that differences in size at rnaturity between different populations do exist, but more information on the life history of this species outside Florida is needed to confirm this premise. The maximum size of over 700 peninsula Florida specimens that 1 have measured is 69 mm. in snout-vent, l&ngth (before preservation) . The largest Virginia Coastal Plain specimen I have measured is 72 mm. in snout— vent length (after several years of preservation). In small samples from many other localities in the range of tills species (Texas, Indiana, New York, New Jersey, Pennsylvania, J.Iaryland, and the Piedmont and Blue r idge Provinces of Virginia, North Carolina, South Carolina, and Georgia), specimens between 79 and 80 mm. in snout-vent length are common, and the larrest sp«ci^«^n exa-nined (UF Pll?, collected 5.? miles sout 1 if "ir>burst, Haywood County, North Carolina), a male, was 86 mm. in snoutvent lenrth before preservation. Cl«arly, the maximum nize of penir^snla Florida glutinosus is less than most other populations, but raaxi"m rr t size obviously cannot be use-* as a systematic character idth

PAGE 159

1*1 which to assign or identify individual specimens. Vomerine teeth iiiittieman (l9t>ls 109) published the vomerine tooth counts on the type series of P, g. c hloro b ryonis . He pointed out that they were much lower than most counts on P. g . glnt inosus . He also sm'geste^ that the vomerine corint in grobmnni might be expected to be even lower than those in chlorobryoni s . Vomerine tooth counts were made on 3? specimens from the Coastal "Plain of Virginia, 13 specimens from the Coastal Plain of North Carolina, and 269 specimens from peninsula Florida. The data obtained are shown in figures lh and 15. In addition, the series of 12^ specimens that Fope and Pope (19l8) used in their study of this character in montane Virginia glutinosus were also examined by the writer, ffy counts on this series were essentially the saiae as the Popes' counts. It is obvious that adult Coastal Plain Virginia and Worth Carolina specimens liave fewer vomerine teeth than either peninsula Florida or Virginia mountain specimens. An examination of the figures shows another difference between the three populations. In each sample there is an ontogenetic increase in the number of vomerine teeth. A t-test on the difference between the rate of increase (slope) of the equations for the Virginia mountain and Florida samples, shows that the probability of obtaining a difference as great or greater than that observed by chance alone is less than .TX)1. In spite of this significant difference in the rate of increase in the number of teeth with growth, there is wide overlap in the number of teeth in similar size groups of the two populations. In the Coastal Plain Virginia and North Carolina samples, there is a further reduction, both in number of teeth and the rate of increase with growth. Some slight overlap between the Virginia mountain and Coastal ! lain Virginia and North Carolina sam-

PAGE 160

-152ples remain, although a majority of adult specimens of each form could be segregated by a striaght line separating the dots representing the t/:o populations. For example, a line passing through the point (snoutvent length * U5» vomerine teeth « 16) and the point (snoutvent length * 70, vomerine teeth « 18) would separate 69 of 75 (92$) Virginia mountain specimens U5 mm. and over in length from 33 of 35 (9U^) Coastal Plain Virginia and North Carolina specimens U5 mm. and over in length. The condition in samples from the Virginia Piedmont is unknown, but the intermediate Florida sample shows wide overlap with both of the above sanples. A significant biological difference between the P. glutinosus of the three areas is obvious, yet this character could hardly be used as a key character to separate either of the Coastal Plain samples, or the Florida and the Virginia mountain samples into separate subspecies. The observed differences in the number of teeth may be the result of a gradual clinal change that is not usually recognized by the systematist as a basis for erecting subspecies. Size of dorsal spots:The size of the dorsal spots of living or freshly preserved specimens was measured with the aid of an ocular micrometer (1 micrometer unit = 0.0685 mm.). Since the size of the dorsal spots increase with growth, the ratio of the average size of ten dorsal spots chosen at random to the snoutvent length was used as a basis of comparison. Specimens of different lengths in the same sample usually show similar ratios when compared in this manner. The data obtained are shown in table IX. It is apparent that in the Atlantic Coastal Plain there is a decided decrease in the size of the dorsal spots from north to south. There is wide overlap between every adjacent population and nowhere is

PAGE 161

153 TABLE IX. GEOGRAPHIC VARIATION HI 0CR5AL SPOT SIZE OF F. GLBTIKOSUS Locality Actual mean (not 1.05.09.013.017.021.0over grouped) U.9 8.9 12.9 16.9 20.9 2it.9 25.0 „ Mean Spot 31 z e ,„„ Ratio -* x 100 Snoutvent length Coastal Plain:

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15Uthere apparent an abrupt change from small to large spots. This character, at least in the Atlantic Coastal Plain, would appear to represent an internal cline. There is not sufficient material available from the Gulf Coastal Plain, except for west Florida, to determine whether or not there is another decrease in the size of dorsal spots to the west. In other parts of the range of glutinosus , from which material is available, the average size of the dorsal spots appears to be fairly large, except for Texas and North Carolina specimens . In North Carolina, the average size of the dorsal spots for 12 specimens from the Piedmont is 13 & , while the average size of 37 specimens from the Blue Ridge Province is 10 .U. The available specimens would not support the contention that there is a west to east reduction in size of spots toward the Coastal Plain, since the Fiedraont series has spots averaging slightly larger than those in the mountains. One of the diagnostic characters given for chlorobryonis is the small size of its dorsal spots. Unquestionably, the size of the dorsal spots of specimens from the Coastal Plain of Virginia and North Carolina is much reduced. This reduction is approached by specimens from Texas, the Blue Ridge Province of North Carolina, and the Coastal Plain of South Carolina. Because of the clinal nature of the variation of this characteristic in the Coastal Plain, it cannot be used as a systematic character to separate the North Carolina and Virginia populations of Coastal Plain glutinosus from the remainder of the species, although it may be considered evidence that these populations show further biological differences from many other populations of glutinosus . All but one of the specimens in the Coastal Plain Virginia

PAGE 163

1# sample are well-preserved specimens in the Carnegie Museum Collection. The one living specimen has a mean spot size to snout-vent length ratio of 6.U, lower than most North Carolina Coastal Plain specimens. In the preserved Carnegie Museum series, only the dorsal spots of specimens in which they are well-defined and clearly visible are included, so that they are almost certainly comparable to the living material on which the remainder of the table is based. One of the characters that Allen and Neill (19U9: 112) used to diagnose P. g. grobmani was "back with tiny scattered dots of a metallic golden color." Data on the geographic variation of the color of dorsal spots will be given below, but it is obvious that the size of these spots in populations from the Coastal Plain of Georgia and Florida are well within the range of many other populations of P. glutinosus . Color of dorsal spots:The color of the dorsal spots is one of the diagnostic characters used in the original description of P. £. grobmani by Allen and ^oill (19h9'112). The dorsal spots of specimens from southeastern Georgia and northern peninsula Florida were described as a "metallic golden color." This character was described as being the most conspicuous diagnostic feature of P. g. grobmani . It is true that a very large proportion of specimens from the range of grobmani possess dorsal spots containing a large proportion of brassy guanophores. This type of dorsal spotting is not restricted to specimens from this area, however. Sinclair (19^0: 200) pointed out that specimens from Shelby County, in western Tennessee, possess dorsal spots that are similar in color to those described for grobmani . I have examined living speci-

PAGE 164

156 nans with as heavy brassy flecking as occurs in topotyic grobmani , from many localities outside its range as defined by Allen and Neill. Specimens in this category were collected in the Piedmont of Qeorgia (Pulton County); the Blue Ridge Province of Georgia (Murray and Gilmer Counties); the Blue 3Ldge Province of Tennessee (Monroe County); Greene County, New York; Pike, f fenroe and Alleghany Counties, Pennsylvania; as well as many localities in the Coastal Plain of Virginia, North Carolina, South Carolina, western Florida, southwestern Georgia, Alabama, and Mississippi. In some of these localities, only an occasional specimen shows a heavy amount of brassy flecking (e.g. the New York, South Carolina and the Georgia Piedmont samples), but in most of the other localities mentioned, it is the usual condition. This is especially true of specimens from southeastern Tennessee and Potato Patch iiountain, .iurray-Oilmer County line, Georgia, whore many individuals exceed the usual peninsula Florida glutinosus in amount of brassy flecking. In spite of the fact that the presence of brassy flecking is not limited to certain southeastern populations, there is considerable evidence that it does not occur throughout the entire range of this species. I have examined more than a dozen living specimens from ingle localities in the Blue Ridge Province of Virginia, North Carolina, and northeastern Tennessee, as well as the Piedmont Province of North Carolina and South Carolina. In most of these specimens the brassy flecking was extremely reduced or absent and in only about 10$ was there a moderate amount of brassy flecking present. In other localities from which I have studied more than a dozen specimens, the usual condition appears to be Intermediate, with a moderate amount of dorsal brassy

PAGE 165

157 flecking. P. glutlnosus of this typo have been exa^oined from central New Jersey (including the vicinity of Princeton, the type locality of P. glutinosus ); Cataraugus County, New York; Randolph County, Vest Virginiaj Giles County, Virginia] the Piedmont of Georgia; and eastern Oklahoma. An objective method of comparing the color of dorsal spots of specimens from different localities was necesr.-
PAGE 166

lse Specimens that were obtained from various parts of the range were maintained alive in a refrigerator at a temperature of about 11° C. At this temperature, adult Flethodon can survive for a year or two with a moderate amount of care and feeding. The maintenance of a collection of living speci-nens of this species was necessary in order to compare living specimens from different localities that could not be obtained alive simultaneously. Some loss of material before it could be thoroughly studied occurred, and as a result, a number of specimens th*t were casually studied on their arrival could not be included in some of the data in the tables because accurate notes on their color or pigmentation were not taken. The data on the color of dorsal spots indicate a large amount of variation in this character in specimens from a single locality. Three characteristics are measured using Villalobos and Villalobos* color guide. These are hue, lightness value, and degree of chroma ticity. Essentially these measure the color, lightness or darkness, and intensity of the color. AH three readings were taken from the spots of a given specimen. One way to analyse individual or geographic variation is to compare each measured quality separately. The color of dorsal spots in P. glutinosus varies from SSO to E (see Villalobos and Villalobos, 19U7t for the meaning of the symbols used here), and in addition there are some individuals in which color is lacking (see table X). In specimens in which the dorsal spots appear brassy to the naked 37s, the color characteristically ranges from to YYO. There appears to be little difference in color in samples with reduced brassy and those in which the brassy

PAGE 167

159m H g o m 3 8 to 1 > CM CM CM t-t ^ CM £> O W^CM fNU"\ C*\ CM cn^t r-» CM CM CM 4" C\ i-( * % d NO CM a a 1 1 o ell 4 3 1 >U) «rl
PAGE 168

160 O E 8 o O CO 03 V t I O s SO ft CM <•> r-. NO -* .-I «-» -* r\ r\ c\ CM SO H H ^t ^ CM i-< CM •* CO «H rv 45 1 CM .* NO CM 1 I a a r-t o o 5 B 2 o 64 « M 1 0) •9 M X

PAGE 169

161 pigment is abundant. In samples in which the brassy pigment is absent or very reduced (Virginia, North Carolina, and South Carolina Piedmont aid KLm ;, i.dge specimens) thern is a tendency for the color to be slightly more green (usual range to LLT) as well as a high percentage of specimens which do not show any color to the naked eye. There is wide overlap in the color of spots of specimens in which the brassy pigment is present and those in which it is absent, but the essential difference in the absence of color in those individuals that have white spots is often detected by this method. The lightness value ranges from 10 to 20 (see table XI), with very little difference between samples from different regions. The sample means vary from Hi. 5 to 19.0 with no apparent geographic trends. A significant difference may exist between samples with or without brassy flecking, but such a wide overlap exists between the two sanrples that this character is of little value in distinguishing between the two types. There is considerable variation in the degree of chromaticity, or intensity of the color, in the various samples (sen table XII). The entire range is from Cray to 9°. In this character, there is a decided geographic variation, the brassy flecked populations showing a much greater amount of color than those that usually 1-ick the brassy-type guanophores. It would be difficult to identify specimens on this basis, although there is a significant difference betv/een the two (cf . the Piedmont and . luo ddge Virginia, tlorth Carolina, and South Carolina samples with the Georgia, Florida, and Alabama Ocastal Plain samples). The Tennessee sample is mado up entirely of sped 1 ens of the brassy

PAGE 170

• 162 8 Os 00 SO rH W. PS. i-l r4 2 3 r\ fi n •* o a vr>^o • •••••••• O-SsOspOvOvOO-O. rHrHHr-lrHrHHr-irH SO CM a a 5 5 O SO O • • • VO SO VO v> H o\ r\ »-* r» rH C\ CM H OS "} ON CM c-> «h j* vg ©*«*%, r> rH rH V> C} .* J* H rH fl CM rH rH CM CM * CM OS V\ CM C»> <-j CM rH rH CM CM CM * CM V\ CM a, u o i o ° a I 3 3 m n CD «a 3 o o a m I 5 c c a a 3 c o O u 2 o I I hi I • p.

PAGE 171

-1638 o\ cQ p.. CM On CO »* *i &• •••••• jw Cs. O*^> «> «>t>. «-» H i"4 «-l 1-4 r-» H * a * CM CM r^ ov e cm •* cm -* • CM i-4 »-l VO i-4 »-» 3 I I CD o c I o r-i O <5 a u

PAGE 172

164 Ov NO f« <*> CM <-l «*» *rv en 8 § «-« H O O J I I VO VO cm cm -3vr\ O r-l H vrv \r\ 3r H cm CO ^ ON r-4 «-l c\ 1 " 1 CM o

PAGE 173

-165o o a I I o OS o o o o ^ °* c o CM O ! O tvr4 £ «>» ^ CC • ••••«• n n 4 h h n ia O O V% • • • O CM H c c. vn • » • CM .3 .=* C>CM H d 4 eg ef fa «K rH r-» V\ •-« .-I CM 5 fl) « 0) 6« | if « n

PAGE 174

166 flecked type, and the Georgia Vledmont and Blue ilidge sample includes both types. The results of the examination of the actual spots under the magnification of a dissecting microscope show a variation that corresponds to the data obtained fr parison with the color charts. The weakness of these data lies in hat the assignment of specimens to each category was tow what ctive. The specimens I used for standards are *oserve: o it will be impossible for other workers to repeat the work without obtaining living specimens from the localities from which individuals were used as a basis of comparison. I chose at from Torreya State Park, Liberty County, Florida, a i ax an animal with a large amount of brassy flecks in each dorsal white spot. A specimen from 5 miles east of Davidson, Cabarrus County.. Carolina, was chosen as the other extreme, completely lac! 1 leeks in the dorsal spots. Specimens that showed a very slight wt of brassy pigment, usually at the borders of the dorsal ;diite spots were regarded as "slightly brassy." Others, that had i ount of brassy flecking in the white spots were placed in an Intel ;y. In a few specimens, usually those with very small spots, no white guanophores could be seen, only the brassy type were present. These were included unchr the heading, "brassy oni . These data are summarised in table XIII (most of the spacim<*is Trm Tennessee included in this table belong to the white-spotted northeaster i essee sample rather than the southeastern Tennessee brass; •llation). The results of these data on the color of the dorsal spots

PAGE 175

167TABLE XLTJ GEOGRAPHIC VARIATION OF DORSAL PIGMENTATION IN PLFTHODOR GLUFINOSUS

PAGE 176

168 are difficult to interpret because of the paucity of material from a great many areas. The only large series are from the southeastern states. The condition in the northeastern portion of the range of glutlnosus appears to be more similar to that of Florida specimens than it is to the color of Piedmont and Blue Ridge specimens from Virginia, North Carolina and South Carolina. The data are sufficient to refute Allen and Weill's suggestion that the brassy condition is limitea to Georgia and Florida Coastal Plain specimens, for most individuals from New York, New Jersey, Pennsylvania, northern Georgia, and southeastern Tennessee have a moderate to large amount of this type of flecking. Whether the Piedmont and Blue Ridge specimens from Virginia, North Carolina, and South Carolina are different enough from animals in the remainder of the range to warrant nomenclatorial recognition on the basis of this character remains to be determined. This may be the case, but a study of much more material froi the northern and mid-western stax.es is needed before a decision can be reached with any degree of certainty . Abundance of lateral pigment :Grobraan (19hU« 262) has pointed out that there is a north-south cline in the amount of white lateral pigment in this species. Specimens from the southeastern Coastal xlain have large lateral spots and in many adults the spots coalesce to form a lateral band of white pigment. Specimens from the southern Appalachians show a reduction in the size of the lateral spots, and farther north, the lateral spots are no larger than those on the dorsum. Grobman Also pointed out that some rr> exas specimens possess a large amount of lateral pigment that forms a band on the sides of adults.

PAGE 177

169 One other difference has been noted In the lateral pigmentation of glutinoeus in the northern portion of itr range, compare' to southern specimens. In a large proportion of animals from Mew York, New Jersey, Pennsylvania, Maryland, and V,'ept Virginia, the pigment in the center of lateral spots seems much more concentrated than that on the edges of the spots, whereas in the south, where the spots are usually larger, the pigment is more evenly distributed. This character is probably also clinal in nature, but has not been analyzed in detail. It is difficult to measure and evaluate the size of the lateral spot*, since two adjacent spots lose their identity as they fuse during growth. I have not made measurements of the size of lateral spots, but superficial examination of specimens from different regions confirms the observations of earlier workers. The lateral b"nd seems to be most prevalent in specimens from the Virginia, North Carolina, and South Carolina Coastal Plain, as well as some Texas populations. Allen and Neill (19U9: 113) report that specinens from Burke, Jefferson, Washington, Qlascock, and Richmond Counties, Georgia, also have an abundance of white pigment on the sides. I have not examined living specimens from any of these counties. Color o f lateral pigment :The color of the lateral spots was used as a diagnostic character in the original descriptions of both grobmani and chlorobryonis . The lateral spots of grobmani were described as being grayish, while those of chlorobryonis are described as being greenish-white or yelloiv-green. A study of the color of the lateral spots similar to that made on the dorsal spots is sunraarized in tables XIV, XV, and XVI. The data on comparisons of lateral spots with

PAGE 178

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PAGE 179

171 | o CM O \Q P» CM CM « ft CM r-4 r-4 f4 s n o o Hi •5 o 8 CM

PAGE 180

4 & o o 8 X 172 o c f >\Ococo^r*'\'Avoo • ••••••••• VO 00 r-t »•* 1 •5 o O a « r\ N rt N W\ r> C"\ r-4 a 3U I 5 9 3 1 & \o l-l Q r\ a \© O 00 CM 4 -'3 :0 OT 1 «6 1-1 I 8 S 1 9

PAGE 181

3 C\ 2 o 173 r-* 00 V© p> » • O OS O * • • vO ""> 0\ l-l fi f-* o VO Ov r4 un CM 03 vr\ cm CM CM cv» f-\ r-l C"\ O CM N tf» Ifi r-» CM vO CM .* 0v v» H CM CM vr» iH vo r% VD CM vO -3" Ox CM <-» w r\ H CN .* r-i cm Ov CM c% r-» c<-\ vo CM CM 1 — • o o a o CJ Sr

PAGE 182

174 crt o Ox O co o o o o o o cm ! c CN ON

PAGE 183

17500 CM vO V* VC> M o a o o\ o 00 o o VO °^ o o o CVi g o o to © % u o v> m h r\ cvj Wi-ioNr-ir\«*..**n rtf-i^r4Qt>»QN ^^ #*^ t"1 r\ r\ r\ vo 3 1 ill A M m 1 1 1 1 * * 1 1 0) J £ 5 CO M 3 £ w © vr\ 3 o VO VO • * (M CM CI vo r-4 »-l ON «T» W 1 c C a 4f g CO

PAGE 184

-176Villalobos and Villalobos' Color Atlas indicate that a large proportion of Florida specimens lack color and appear gray, while a majority of North Carolina Coastal Plain specimens, as well as those from Texas, are found in the yellow to orange range. In most other samples there is a large amount of variation in the color of the lateral spots. There is slight variation in the lightness of the spots, but wide overlap between most samples. The amount of chroraaticity appears to be quite variable, with the Florida and Virginia samples having an especially large proportion of gray individuals. Although there may be significant biological differencebet een these samples in the various aspects of the lateral guanophore pigmentation, there certainly would seem to be no justification in naming either the North Carolina Coastal Flain or the Florida and Georgia Coastal Plain populations as recognizable subspecies on the basis of this character. Studies of the lateral pigment with the aid of a dissecting microscope indicate that the yellow pigmentation is not to the presence of brassy guanophores, but rather due to an actual difference in the color of the non-iridescent guanophores that appear to be the same type as those found on the dorsum, where they are always white. In classifying the color of the lateral spots, three categories were used, white, intermediate (yellowish-white), and yellow. The data on this character are tabulated in table XVII. In the Atlantic Coastal Plpin there appears to be an internal cline in the amount of yellow pigment, with a reduction in intensity from North Carolina to Florida. The Texas sample has lateral pigment that is extremely yellow, but the average condition of specimens in the remainder of the range is intermediate between the two extremes. A large amount of individual vnria-

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177 -

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178 tioai is present in this character within adults of each population. There is also an ontogenetic change in the appearance of lateral guanophore pigment, judging from the appearance of young speci ens in populations in which the normal adult spotting appears yellow. Young individuals of all samples I have examined possess white lateral guanophores and the change to yellow occurs at an earlier age in populations ir, which the normal adult condition is yellow. Absence of guanophores tSpecimens that completely lack dorsal and lateral guanophore pigmentation are occasionally found in samples of glutinosus from various parts of its range. In at least two areas, this characteristic is present in a large proportion of specimens and thus deserves special mention* Heill (19U8) reports that in Jasper County, South Carolina, there exists a population of P. glutinosus in which 100$ of the specimens are characterized by the complete absence of guanophore spotting. In adjacent areas, individuals in which guanophore pigmentation was absent were also recorded, but they occur with the normally spotted individuals. I have examined two series of P. glutinosus from Jasper County and in both series, numbering 16 and 7 respectively, every specimen was entirely black. Two of 12 speci' ens th^t I have examined from a locality in Dorchester County, South Carolina, also lacked guanophores, confirming the observations of earlier workers (Cope, 18R9: lli?; Schmidt, 192U» 67 J Neill, 19U8) that the unspotted type also occurs in the Charleston area. Several deviations described by Neill (19U8) for South Carolina Coastal Plain materinl are not confirmed by this study. In specimens examined by me, no consistant differences in the number of costal grooves, proportional length of tail, parasphenoid teeth, melanophore pigmentation, size, head

PAGE 187

17? shape, or structure of the vent in males could be detected. In collecting specimens at the locality mentioned by t!eill (I9U8), 7 tuiles north of Tillman, Jasper County, South Carolina, I was impressed by the unusual agility of the animals in escaping the collector, as well as the apparent delicacy of the skin, compared with other Coastal Plain giutinogus . 'Whether these differences are real or not, they would be difficult to measure objectively, although histological examination of the integument of the dark form might reveal structural differences other than the absence of white spots. Another Coastal Plain locality in which a large proportion of specimens lack guanophore spotting was discovered in Florida. On the east side of the St. Johns River in "folusia County, about 1 mile south of Astor, there is a population of F. glutlnosus in which about half (111 of 30) of the specimens are entirely black. On the west side of the St. Johns Iliver, in three nearby localities in Lake County (within 10 miles of Astor) only one of 107 specimens showed a complete absence of dorsal and lateral white spotting, so common on the east side of the river. Apparently the gene or genes responsible for this condition occur with small frequency in some glu t lnosus populations, but have become fixed in the Jasper County, South Carolina population and are present in much greater than usual frequency in the Volusia. County, Florida population . Grobaan (19UU* 261) states that "there is, apparently, a large glutinosus devoid of white marking which ranges from the Cumberland Plateau. Localities for this form are in Bibb, Chilton, Lawrence, and St. Clair Counties, Alabama; Chatooga, Drde, DeKalb, Fulton, Gilmer, and Jiurray Counties, Georgia; Edmonson, Harlan, Laurel, Morgan, and Whitley

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180 Counties, Kentucky; Pontotoc and Webster Counties, fctLssissippi; Avery, Buncombe, Cherokee, Graham, Haywood, Macon, Polk, Swain, and Transylvania Counties, North Carolina; and Claiborne, Cumberland, Davidson, DeKalb, Fayette, Fentress, Rhea, and Sevier Counties, Tennessee. " I have examined living specimens from Fulton County and the JAirray-Gilmer County line, Georgia; Harlan County, Kentucky; Haywood, Maoon, and Swain Counties, North Carolina; and Davidson and Rhea Counties, Tonnessee. Every specimen that I have exaadned from these counties, as well as many others from other counties in the region of the supposed unspotted form, possess both dorsal and lateral guanophore spots. Appajw ently the difference noted in preserved specimens is not present in life, and may merely result from the disappearance of the spots as a result of preservation. Abundance of melanophore pigmentation on chin :In 19UU, Grobman described a Texas population of Plethodon glutinosus as a separate subspecies, P. £. albagula . It was diagnosed as a "race of glutinoaus in which the arrangement of the black pigment to form small compact circles has been broken down in the gular region, thereby presenting a gross appearance of a lighter throat than that found in the typical race of glutinosus ." An examination of Grobman's map of the distribution of glutinosus shows a gap of approximately 120 miles between the McLennan County and the Upsur County records of glutinosus in Texas. If this hiatus actually represents an area in which f-lutinosus no longer occurs, then the population found along the Balcones Escarpment in south-central Texas would seam to be isolated from its nearest relatives and differentiation might be expected. However, specimens fro™ two localities in

PAGE 189

181 the region of the Balcones Escarpment were tentatively referred by urobman to typical P. g. glutinosus . ihese are from Bell and Blanco Counties, and the latter record, based on only one specimen (CM 6129), is only about 20 miles from localities at which paratypic albagula were collected. Brown (1950: 32) assigned specimens from Bell, Bexar, Blanco, Comal, Kays, Kendall, Real, and Travis Counties to P. g. albagula , although Grobman had indicated that specimens from two of these counties had dark chins. Brown also remarked that "the status of Plethodon glutinosus albagula as a valid subspecies needs further investigation. White throated forms have been reported from other areas supposedly in the range of P. g. glutinosus . " Several series of recently collected specimens from Texas, have confirmed the fact that both lightand dark-chinned forms occur in the region described as being inhabited by albj^ula. A series of 2k specimens (JSM 667-676, UF 8085 (U), 83U2 (U), 83U3 (6)) from a sink hole about 5 miles southwest of Austin, Travis County, are typical of Grobman' s description of albagula . They possess very light chins and in life, had an abundance of yellow lateral pigment, which, except in very young specimens, formed a continuous lateral band of guanophore pigment. This condition was mentioned by Grobman (19U*s 28U) as an additional characteristic of albagula , although he pointed out that this type of pigmentation was not entirely limited to albagula . The color of this lateral band in life is unlike that of glutinosus from other parts of its range. Grobman also mentioned that the dorsum of albagula is almost devoid of white pigment spots. The living specimens

PAGE 190

182 did not lack dorsal white spots, but the spots were very small in relation to the sise of the animal compared to most glutinosus. In addition to these characters, these specimens of albagula were found to differ from glutinosus in that a rather high proportion possess 18 trunk vertebrae. The vertebrae of one s«-fill specimen could not be counted accurately, but of the remaining 23, seven have 18 trunk vertebrae, four have 17A8 (the 18th trunk vertebra beers the sacral rib on one side, the 19th on the other side), and 12 possess 17 trunk vertebrae. Two freshly preserved specimens (UF 8009) from the Guadalupe iiiver bluff, 2 miles south of Sattler, Comal County, are identical in appearance with the above series. Both possess 17 trunk vertebrae. Available freshly preserved specimens from two other localities, UF 71U3 (3), 71U5 (1), from the head springs of Mill Creek, *>' miles northeast of Vanderpool, Bandera County, and UF 3^13 (6) from Bull Creek, west of Austin, Travis County, are strikingly different from albagula . The sides of these specimens, although preserved for a shorter period than many of the albagula , possess only a few small scattered white spots. In none of these specimens was there any tendency toward a coalescence of the lateral spots into a band. The chins of most of these sped' ens are as dark as their bellies. In only three young individuals are they slightly lighter than the bellies, but in none are the chins as light as in albagula of comparable size. Dr. Grobman has kindly examined the specimens mentioned and has informed me that the series on which he based the description of albagula were similar to the specimens that were collected at the sink hole, about $ miles southwest of Austin. The dark-chinned samples obviously do not fit the description of albagula and are thus similar

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183 to the specimens he examined from Bell and Blanco Counties. Additional material from Texas, not examined by Grobman (1°U0> was borrower) from Bryce C. Brown, Ottys Sanders, and the Strecker ttfr> seum. Most of the Strecker iuseum's collection is so faded and stained due to many years of poor preservation, that it is impossible to allocate some speci tens to either form. A series of 15 specimens (SM 16, 21*7-57, 1002, 1055) from Helotes, Bexar County, cannot be determined, although at least 11 have 17 trunk vertebrae. Two others probably have 17, while the other two are too small to make accurate counts from the radiographs . One specimen (S4J 5087) from Austin, Travis County, appears to have reduced melanophore pigmentation on the chin (although its chin now appears dark due to staining). SU 5213-16 and 5223, from Frio Canyon, Real County, may definitely be assigned to the dark-chinned form. They are presumably the specimens on which Strecker (1935* 32) based his Real County record, that Grobman (I9hkt fig. 3) included as a possible albagula locality on the distribution map. They are not albagula , but the dark-chinned form. Three have 17 trunk vertebrae, the other two probably also have 17 trunk vertebrae. A large series of UO specimens from San Marcos, Hays County (SH 1837, 32U1-2, 32Uh-76, U963-7) are in poor condition. 27 have 17 trunk vertebrae, 8 probably also possess 17 trunk vertebrae, and the vertebrae in the remaining specimens could not be counted. The melanophores on the chins of some of the specimens could not be discerned, but in most they appear to be much reduced as in albagula . One specimen

PAGE 192

18U(SM 3$ik), however, possesses as dense a melanophore network on the chin as on the belly. In most of the larger specimens of this series, it appears that white pigment was abundant on the sides, as in alba gula . Four other specimens from San ftercos (OS 613, BCB 113>, 2087, and 2ii23) also have light chins and 17 trunk vertebrae. Although all of the San ?iarcos specimens for which accurate counts are available possess 17 tnsik vertebrae, on the basis of their light chins they appear to be referable to albagula . Grobraan (19Uk: 283) designated 31 specimens from San Marccs as paratyoes of albagula . Three specimens (BOB 116-8) from Deep Eddy, west of Austin, Travis County, have dark chins and two (117-6) have 17 trunk vertebrae. BOB 5269-73 were taken from a cave near McNeil, Travis County. All five have dark chins and two have 18 trunk vertebrae, the other three 17. Ten specimens (SH 3162-3* BCB 7015-22) from Fern Bank, near Wimberly, Hays County, have light chins of the albagula type. Two have 18 vertebrae, the other eight 17. The presence of 18 vertebrae in only two other series from Texas, one of which is not albagula , would seem to indicate that the high proportion of 18 trunk vertebrae in the one sample collected 5 miles southwest of Austin, is either of local occurrence or the result of random sampling error ana not of taxonondc significance. Three specimens (CNHlf 37666-8) from y&mberly were designated as paratypes of albagula . A series cf 16 specimens frcra the vicinity of another paratypic locality of albagula (BCB 3lj31-U0, woods near Spring Creek, 11 miles northeast of Boerne, and BCB U 98 3-8, from Schneider Cave, Hi

PAGE 193

185 miles northeast of Boerne, Kendall County) are unusual in having a large amount of ventral guanophore spotting. All but three have light chins of the albagula type, and all (except U987, that is too small to obtain an accurate vertebral count) possess 17 trunk vertebrae, DGB 223U, near Cibolo Greek, 18 miles northeast of oan Antonio, Bexar County, in the vicinity of the type locality of albagula , has 17 trunk vertebrae and a light chin. It also has a large number of lateral spots, but it is a young individual and the spots have not yet coalesced to form a band. Several other specimens from the vicinity of Austin, Travis County (BCB 301, 1161-2, Zilker Park, Austin; BOB 799, Lake Austin; BCB 1163-70, 1956-7, Austin Caverns, west of Austin; BCB 1277-78, Barton Creek, south of Austin; and UF 3U1U (10), 3U6Q (1), junction of Barton and Little Barton Creeks), include specimens with both light and dark chins. All except BCB 1163, 1166-8, and 1277-8 which could not be accurately counted, possess 17 trunk vertebrae. It is difficult to analyse the distribution of the two forms, if indeed, there actually are two forms represented, because of the incoiqplete data on the characteristics of many of the specimens examined. Based on the freshly preserved University of Florida material, it would seem that we are dealing with two different forms, as different from each other as most species of Eastern Large Plethodons. The lateral band of albagula is very similar to the type found in the yonahlossee group. The light chin, a characteristic of the yonahlossee group, is also found in albagula . On the other hand, some Texas populations seem to have both lightand dark-chinned specimens represented. I

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166 have not examined living specimens of the Texas dark-chinned form, although the preserved specimens appear to be very similar to P. g. glutinosus . laving albagula are very distinct from P. £. glutinosus, and it may be that albagula is a valid form, possibly a distinct species more closely related to the yonahlossee group than to the glutinosus group, or else intermediate between the two groups. This would not rule out entirely a subspecific relationship with glutinous , if intergradation could be demonstrated. Another possible interpretation is that there may be some environmental influence that is acting on certain Texss populations of glutinosus (i.e. albagiLa), causing them to differ phenotypically from other populations of this species. If this were the case, then most systematists would recognize albagula as an ecotype, not to be given nominal recognition. Grobman (19lJbs 28U) points out that most of the available habitat labels accompanying specimens of albagula mention caves. *>re recent collections seem to indicate, however, that both types may be found either in or out of c?ves. Constant temperature conditions in caves might play an important part in producing a different type of pigmentation than that found in animals that undergo their embryonic development outside of caves. Plethodon glutinosus is known to lay its eggs both in and out of caves. Grobman (19bl* ?fi2) dlscuss«e the possible importance of temperature during embryonic development in the production of lateral pigmentation. Temper a tui*e might also have an effect on the number of somites (and vertebrae) in this species. The I-alcones Escarpment separates two major physiographic provinces, the Coastal Plain and the Edward's Plateau section of the

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167 Great Ilains. The change in the underlying rocks, vegetation, and rainfall at the Balcones Bscarpment is very abrupt (Smith and Buechner, 19U7). No correlation is indicated on the basis of the present distributional data, however, since both lightand dark-chinned specimens have been collected on both sides of the Balconee Escarpment (figure 23). A third possible explanation of the situation, the one reflected by the current nomenclature, is that the two forms are subspecifically related. The puzzling samples from Travis County, in which both lightand dark-chinned specimens, as well as intermediate types appear in one population, could be considered evidence for intergradation between typical P. g. glutinosus and P. g. albagula . according to this interpretation, the specimens from Bell, Blanco, Bandera, and Real Counties might be assigned to P. £. glutinosus , occurring, as they do, to the north and west of the albagula records, tost of the specimens from hays, Comal, Kendall, and Bexar Counties show the characteristics of albagula , while both typical albagula, typical glutino sus , and interrodiates occur in Travis County. Geographic replacement between the two forms is a definite possibility. If the true relationship is found to be subspeeific, it may be postulated that the isolation in which albagula differentiated was of considerable duration and that typical glutinosus has only recently reinvaded the region, vrith only a slight amount of secondary intergradation . A field study of the species in the area involved is much needed. A comparison of living specimens fron the entire region, a study of the life history, ecological requirements, and more ixiformation on the distribution of the animals will be needed before the proper

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188 systematic status of the Texas Large Plethodons can be determined. With our present knowledge, it would seera best to propose no change in the existing nomenclature with regard to albagula , pending t study that would clarify its true status. Wright and Wright (1938: 3U) report P. glutinosus from Medina County, Texas, but no specimens seem to be Available now from this county. Streckor (1902: lOOj 1908: 80) reports the species froir Me Lennan County on the basis of two specimens, one a sight record. Its actual occurrence in this county certainly needs confirmation. If P. glutinosus does not occur in ^.icLennan County, then the gap in the range of Texas glutinosus is greater than indicated above. Strecker (1915* 55) states that he collected two specimens near Cleveland, Liberty County, in the southeastern Coastal Plain of Texas. Since no other records of P. glutinosus are known for over 150 miles from this locality in any direction, this record should be deleted from the authenticated range of P. glutin osus until it can be verified. Summary of geographic variation in xlethouon glutinosus :In most of the meters studied, geographic variation was found to ue present. These characters were not chosen at random, nowever. They were selected for detailed analysis because previous workers had found evidence of geographic variation in them and this had been supported by preliminary observations made by the writer. In a criti cai study of other characters of living specimens from various parts of the range, I could find no others that could be successfully correlated with geography. In the number of trunk segments, the differences between

PAGE 197

189 P. glutinosus Q glutmosus A olbagula •-... J,., KERR ,«END* L l\y , / ~* ,i ^ ' \ -; % a/ -' '"Figure 23. The distribution of the darkchinned (gl utinosus ) and light-chinnod ( albagula ) populations of P. glu tinosu s in central Texas, Solid symbols represent localities from which specimens have been examined by the writer. Hollow symbols represent literature records of Grobman (19UU).

PAGE 198

190 geographic saoples are very slight, but in the other characters studied, consistent differences between some samples are demonstrable. Southeastern Coastal Plain specimens are significantly saaller than most other P. glutinosus . The number of vomerine teeth is reduced in Florida glutinosus , compared with the Virginia mountain sample, end the North Carolina and Virginia Coastal Plain specimens exhibit a further reduction. The size of the dorsal spots shows clinal variation in the Atlantic Coastal Plain, increasing from north to south. The spot size of other populations, except those in the Blue Ridge Province of North Carolina, and in Texaa, appears to be about the same as southern Coastal Plain animals. The color of the dorsal spots is eat* tremely variable, but uniformly white in the southern Piedmont and Blue Ridge Provinces north of Georgia, and uniformly brassy in the Gulf Coastal Plain from klssissippi to Florida. Lateral spots are larger and mors yellow in color in the Atlantic Coastal Plain and in Texas. Texas animals usually possess a great reduction in the number of melanophores on the chin. On the basis of the above data, several tentative conclusions on raciation in P. glutinosus may be suggested. The Texas populations are apparently isolated from the remainder of the species. Living Texas specimens that have been examined differ in several ways fro.T. glutinosus in other parts of its range and are the most distinctive of all of the samples of this species studied by the writer. For the present, it would seem advisable to continue to recognize al bagula as a separate form, although there remain many questions that must be answered before its correct systematic position can be determined.

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191 nether population that differs from raGst other P. g lutinosus in a number of ways is the one that occurs in the South Carolina, North Carolina, and Virginia Coastal Plain. Its reduced vomerine teeth, small dorsal spots, lateral yellow band of guanepbore spots, and srall size serve to distinguish a large proportion of these animals from the remainder of the species. No single one of these characters is United to this population, however, and all but the last appear to be the remit of a gradual clinal-type change from south to north in the Coastal Plain, with wide overlap between adjacent populations. Most systeroatists would not recognize the end products of such a cline as j*or»ar«te subspecies. For those that care to recognize the northern end of this cline as a nominal form, the name cylindi-acea is available for the South Carolina population. P. g_. chlorobryonis tfittleraan, based on specimens from the North Carolina Coastal Plain would not appear to be valid, since tnere ia wide morphological overlap between the North Carolina and South Carolina populations. Similarly, for those who would recognize the southern end of this sline as a separate subspecies, the name grobmani would not be applicable, nince it is preoccupied by at least two earlier navies, yarielata being the oldest. This name would be available for all Coastal Plain P. glutinosus if, in the future, it is shown that these populations should be given racial recognition. At present, there are no 't.otti characters, other than size, that would support this thesis. Although, as yet, there is not sufficient reason to recognize a southeEstern Coastal Plain race (or races), the information on geographic variation of several characters is extremely suggestive in explaining

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192 some of the distributional problems and relationships of these populations of Piethodon g lutinosus . The Florida and Georgia samples are closer to the nonCoastal Plain samples than are the Virginia and Carolina Coastal Plain populations in every character except maximum size. If the wideranging P. glutinosus was first able to adapt itself to the Coastal Plain environment in Georgia (or, perhaps to the west of Georgia), it probably migrated to the northeast into the coastal areas of the Carolinas and southeastern Virginia. There are no records of glutinosus from the Coastal Plain of northern Virginia, the Del-Mar-Va Peninsula, southern New Jersey (south of Ocean County), or Long Island, although Piethodon cinereus has been able to adapt to Coastal Flain conditions in all these areas. It would therefore appear that northeastern Piedmont glutinosus have been unable to move into the Coastal Plain. The fact that Virginia Coastal Plain glutinosus are so different from upland Virginia glutinosus lends support to the theory of a more southerly origin of the former, rather than a close relationship to glutinosus populations in the adjacent Piedmont. One other interesting situation has been discovered as a result of field work being done in southeastern Tennessee by )r. Richard M. Johnson. Nicholls (1950: 312) has suggested that there is more than one type of Piethodon glutinosus in eastern Tennessee, but does not describe them. Field work in western North Carolina has indicated that glutinosus of that region are unusual in invariably possessing white dorsal spots. Mr. Johnson has also collected this white-spotted glutinosus at several localities in eastern Tennessee.

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193 In other localities, however, eastern Tennessee glutinosus populations exist in which the amount of brassy flecking is greater than in most Florida specimens. (This type of brassy flecked glutinoaue also occurs in northern Georgia at Potato Patch Mountain, ftirrayGilmer County line). It is quite possible that more than one form is represented in this area and it is hoped that Mr* Johnson's field studies will clarify the altitudinal, ecological, and geographic distribution of the two forms, as well as to shed some light on their relationships . Plethodon glutinosus glutinosus (Green) Salamandra glutlnosa Green (1818: 357). (Type locality probably Princeton, New Jersey) Salamandra yariolata Gilllaras (I8l8s U6o). (Type locality, Southern States, probably Florida) Salamandra cylindracea Harlan (1825: 156). (Type locality, South Carolina, probably the vicinity of Camden) Plethodon glutinosus (Green). Tschudi (I838: 58). Bishop (19Ul: 219-32). Plethodon glutinosua (Green). Gray (1850: 39). Salamandra albopunctata Valenciennes in Dumeril and Eibron (185U: 81). (Type locality, Savannah, Georgia) Plethodon variolosum (Gilliams). Dumeril and Bibron (185U» 83). Salamndra elongata Valenciennes in Dumeril and Bibron (185U: 8U) (Type locality not designated) Salamandr a melanoleuca led (1865: 130). (Type locality, Nazareth, Pennsylvania) Plethodon glutinosus glutinosus (Green). Dunn (1920* 131). Bishop (19U3: 250-3). Grobman (19W*: 278-83).

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19U Plethodon glutinosus grobmani Allen and Weill (19U9* 112). (Type locality, near Silver Springs, tiarion County, Florida) Plethodon kentucki '&ttleraan (1951: 105-8). (Type locality, Big Black 'fountain, Harlan County, Kentucky) Plethodon glutinosus chlorobryonis Mittleman (1951* 108). (Type locality, 13 miles north of New Bern, Craven County, North Carolina) Plethodon jordani kentucki 'fittleman. Schmidt (1953* 37). Type :Dunn (1926* 138) states that the type is not known to exist, but that the type locality is obviously Princeton, New Jersey. Diagnosis :A dark-bellied, dark-chinned Eastern Large Plethodon with dorsal and lateral guanophore spots, and without red pigment. Kange iCentral New York west to central and southern Illinois, south throughout all the states east of the i&ssissippi Hiver, except for southern Florida, and the Atlantic Coastal Plain north of southern Virginia. Aest of the :<*ississippi River, it occurs in east-central and southern Missouri, the northwestern half of Arkansas, eastern Oklahoma, and extreme northeastern Texas (figure 2h). Dark-chinned specimens which may be referable to this subspecies are also known from Real, bandera, bianco, Travis, and Bell Counties in the region of the Balcones Kscarpment of Texas (figure 23). Descriptio n:This form is extremely variable and the reader is referred to the above discussion of geographic variation in this species for descriptions. The costal grooves usually number 16, the trunk vertebrae 17. Vomerine teeth range from h to 17 in a series.

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* 195 Figure 2U. The distribution of Plethodon glutinosus in eastern United States. Solid symbols represent localities from which living specimens have been examined. Hollow symbols represent literature records and localities from which preserved specimens have been examined. See figure 23 for a map of the distribution of the subspecies of P. g lutinosu s in Texas.

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196 There is geographic variation in size, the largest peninsula Florida specimen (from Eureka, Marion County), is 69 mm. in snout-vent length, while the largest specimen on record (Orton, 19U6) is 88 mm. in snoutvent length. Sexual maturity is reached between UO and 56 mm. in snout-vent length in Florida, but there is little or no information for other areas. Plethodon glutinosus albagula Grobman Plethodon glutinosus albagula Qrobman (19UU: 283). Brown (1950: 32-3). Type:CM 9652, an adult male, collected 20 miles north of San Antonio, Bexar County, Texas, on February 2U, 1935, by Wesley Clanton. Diagnosis :A subspecies of Plethodon glutinosus that differs from the typical race in possessing a reduction of melanophore pigmentation on the chin. Range :Known from Bexar, Kendall, Comal, Hays, and Travis Counties, Texas. Description :Living specimens from a sink hole, about $ miles southwest of Austin, Travis County, Texas, besides possessing the diagnostic light chin, also have exceptionally yellow lateral spots, which are often so large that adjacent spots fuse to form a lateral band in adults. The dorsal spots are snail and white. As noted above, this population is unusual in the number of trunk vertebrae, so it may not be typical in other respects. The costal grooves usually number 16, the trunk vertebrae 17. Vomerine teeth range from h to 11 in a series. This form is large in

PAGE 205

-197siae, the largest specimen examined is 75 nn. in snout-vent length. MflNUPE A systematic study of the North American salamander genus Plethodon indicates that there are three major groups in the genus. These are called the Western Plethodons, the Eastern Small Plethodons, and the Eastern Large Plethodons. Special emphasis has been placed on a study of pigmentation characters in living specimens and in the variation in trunk vertebrae measured by means of X-ray photographs. A method of correlating the number of costal grooves with the number of trunk vertebrae has been suggested. ihe Western Plethodons are divided into four species groups. The Plethodon vandykei Group, with one species, vandykei , includes three subspecies. The Plethodon vehiculum Group includes two laonotypic species, dunni and vehiculum . The third species group includes a single species, formerly known as P. elongatus , for which a new name is proposed, P. productu s. The fourth species group of Western Plethodons also contains a single species, P. neomexicanus . The Eastern Small Plethodons are divided into two species groups, the Plethodon welleri Group and the Plethodon cinereus Group. The first includes two species, welleri and richmondi (with three races). The Plethodon cinereus Group consists of two species, dorssiLis with two subspecies, and cinereus with three races. The Eastern Large Plethodons are divided into three species groups. The P. wehrlei Group includes one species, wehrlei , with three

PAGE 206

198 races. The Plethodon y onahlossee Group Includes three species, yonahlossee, ouacnitae , and caddoensis. The Plethodon glut,inosus Group has two species, jordani with eight subspecies (one, unicoi , is new), and glutinosus with two races. The Eastern Large Plethodons and the Eastern Small Plethodons are more closely related to each other than either is to the Western Plethodons. The Eastern Small Plethodons appear to be closer to the Western Plethodons than are the Eastern Large Plethodons. LITERATURE CITED Allen, E. Ross, and Wilfred T. Neill 19U9. A new subspecies of salamander (genus plethodon ) from Florida and Georgia. Herpetologica, 5 (?): ll?-llU. Bailey, Joseph R. 1937. Notes on plethodont sala-nanders of the southeastern United States. Occ. Papers *5us. Zool., Univ. &ch., (36U): 1-10. Baird, S. F. 1850. !tevision of the North American tailed-batrachia, with descriptions of the new genera and species. Jour. Acad. Nat. Sci. Phila., 2nd ser., 1 (l)« 281-292. Barbour, Roger W. 1953. The amphibians of Big Black iaountain, Harlan County, Kentucky. Copeia (2): -U-89. Bishop, Sherman C. 192U. Notes on salamanders. Bull. N. Y. State ^ius., (?53)» 87-102, pis. 1-3. 19?7. The amphibians and reptiles of Allegany State Park. N. 1, State Mus. handbook (3): 3-lUl, figs. 1-58, map. 1928. Notes on some amphibians and reptiles from the southeastern states with a description of a new salamander from North Carolina. Jour. Misha Mitchell Sci. Soc, U3 (3-U)s 153-170, pis. ?3-?6. 193li. Description of a new salamander from Oregon, with notes on related species. Froc. Biol. Soc. ^ash., U7* 169171, 1 pi."-

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199 Bishop, Sherman C. -_«*«•*«.«* 19Ula. The salamanders of New York. Bull. N. Y. State Mm*! (32U)» 1-365, figs. 1-66. 19Ulb. Hox.es on salamanders with descriptions of several new forms. Occ. Papers Mus. Zool., Univ. Mich., (U51)* 1-25, pis. 1-2. 19U3. Handbook of salamanders. Ithaca. Comstock Pub. Co.: i-xiv, 1-555, pi. 1, figs. 1-lUi, maps 1-56. Blanchard, Frank N. _ 1926. A collection of amphibians and reptiles from southern Indiana and adjacent Kentucky. Papers Mich. Acad. Sci., Arts and Letters, 5* 367-3P8, pis. 22-23. Blatchley, W. S. 1901. On a small collection of batrachians from Tennessee, with descriptions of two new species. 25th Ann. Rept. Dept. Geol. xncLUna, 1900: 759-763. Bogart, Charles M. 1952. Relative abundance, habitats, and normal thermal levels of some Virginian salamanders. Ecology, 33 (l) s 16-30, figs. 1-6. Bonaparte, C. L. 1839. xcon. Fauna Italica, 2 (26): no pagination. Rome. * 1882! Catalogue of the Batrachia Uradientia S. Caudata and Batrachia Apoda in the collection of the British i^useum, 2nd Ed.: i-viii, 1-127, pis. 1-9. Bragg, Arthur N. ,. ,. 195?. Amphibians of r/cCurtain County, Oklahoma, .asmann Jour. Biol., 10 (2): 2U1-250. 1955. The Amphibia of Cherokee County, Oklahoma. Berpetologica, 11 (1): 25-30. Bragg, Arthur N., and W. F. Hudson 1951. New county records of Salientia and a summary of the known distribution of salamanders in Oklahoma. Great Basin Nat., 11: 87-90. Breder, C. M., Jr., and R. B. Breder. 1923. k list of fishes, amphibians, and reptiles collected in Ashe County, North Carolina. Zoologica, U (l): 1-23, figs. 1-8.

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200 Brimley, C. 3. 1912. Notes on the salamanders of the North Carolina mountains with descriptions of two now forms. Proc. Biol. Soc. ash., 2$: 135W), pis. 6-7. 1926. Revised key and list of the amphibians and reptiles of North Carolina. Jour. Elisha ^tchell Sci. Soc., U2i 7^-93. 1927a. Some records of amphibians and reptiles from North Carolina. Copeia (162 s 10-12). 1927b. An apparently new salamander ( Flethodon clemsonae ) from S. C. Copeia (16U): 73-75. 1928. Yellow-cheeked Desmognathus from Macon County, N. C. Copeia (166): 21-23. 19U0. The amphibians and reptiles of North Carolina. Carolina Tips, 2 (2): 6-7. Brooks, Uaurice 19U5. Notes on amphibians from Sickle's Knob, West Virginia. Copeia (It): 231. 19U8. Notes on the Cheat Mountain Salamander. Copeia (U): 239-2UU, fig. 1. Brown, Bryce C. 1950. An annotated check list of the reptiles and amphibians of Texas. Baylor Univ. Studies, i-xii, 1-258. Brown, . C, and J. R. Slater 1939. The amphibians and reptiles of the islands of the state of •' ashington. Occ. Papers Dept. Biol., College of Pu*et ound, (kj* 6-13. Burns, Douglas M. 195U. A new subspecies of tJie salauander Flethodon vandykei . Herpetologica, 10 (2): 83-87. Chadwick, Claude S. 19U0. Some notes on the burrows of Plethouon metcalfi. Copeia (l): 50. Chamberlain, *). B. 1928. Some salamanders from Caesar's heac, iiouth Carolina. Copeia (167)' ^1-52. Chermock, Ralph L. 1952. A key to the amphibians and reptiles of Alabama. Geol. Surv. Alabama, !\*us. Paper (33)' 1-88, figs. 1-10.

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201 Clay, (Tilliaii M., iioberta Burckhardt Case, and Robert Cunningham 1955. On the taxonomic status of the slimy salamander, " lethodon glutinosus (Green), In southeastern Kentuc'^. Trans. Kentucky Acad. Sci., 16 (3) J $7-65, figs. 1-2. Coker, V. C. 1939. Opportunities for biological work at Highlands and report of progress. Highlands Mus. and Biol. Lab. Pub. (U): 1-16, fig. 1. Cooper, J. G. i860. No. h report upon the reptiles collected on the survey, itept. ISxpl. and Surv. from Miss. Hiver to Pacific Ocean, 12 (2): 292-306, pi. 31, fig. k. Cope, S. D. 1867. A review of the species of the Ambystotaidae. Proc. Acad. Nat. Sci. Phila., 19: l6o-211. 1869. A review of the species of the Plethodontifee and Desmognathidae. Proc. Acad. Nat. Sci., Phila., 21: 93-118. 1883. Notes on the geographic distribution of Batrachia and Reptilia in western North America. Proc. Acad. Nat. Sci., Phila., 35: 10-35. 1889. The Batrachia of North America. Bull. U. S, Nat. Mus., (3U): 3-525, pis. 1-79, 83, 86, figs. 1-119. Cowan, I. M. 1937. A review of the reptiles and amphibians of British Columbia. Rept. Irov. ifois. Nat. hist, for 1936: 16-25. Cox, Philip 1907. Liaards and salamanders of Canada. Proc. Miramichi Nat. Hist. Assn., (5)s U6-55. Davis, Mathan Smith, Jr., and Frank L. Rice. 1683. Descriptive catalogue of iiorth AaorUaw Batrachia and Reptilia found east of the Mississippi liiver. 111. State Lab. Nat. hist. Bull., 5* 1-67. Deevey, E. S., Jr. 19h9. Biop.eop;raphy of the Pleistocene. Part I. jiurope and North America. Bull. Geol. Soc. Amer., 60: 1315-LU16, Tigs. 1-27. Duellrnan, William E. 195U. The salamander Plethodon ri chnondi in southwestern Ohio. Copeia (1): L.0-U5, fig. 1. Dumas, Philip C. 1955. ^ggs of the salamander Plethodon dunni in nature. Copeia (1): 65.

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202 Dumeril, A. i. C., and G. Bibron 185L. Fdstoire Neturelle des Reptiles, 9'i-xx, 1-liUO. Dundee, Harold A. l°i±7. Notes on salamanders collected In Oklahoma. Copeia (?)i 117-120. Dunn, B« P. 1917. .eptile and amphibian collections from the North Caroline mountains, with especial reference to sala/aanders . Bull. Aaer. Mus. Nat. Hist., 37* 593-63U, pis. 57-61, figs. 1-7. 1918. The collection of Amphibia Cauda ta of the ituseum of Comparative Zoology. Bull. IAis. Comp. Zool., 62: UiS-U71. 1920. Some reptiles and amphibians from Virginia, North Carolina, Tennessee and Alabama. I-roc. idol. Soc. V : ash., 33: 129-137. 1926. The salamanders of the family Flethodontidae. Smith College Ann. Pub,: 1-UUl, pis. l-3» maps 1-86. Dunn, E. R., and A. A. I;ein?e 1933. A new salamander from the Ouachitae fountains. Copeia (3): 121-12*, 1 fig. Dury, Ralph, and Tdlliam Gessing, Jr. 19U0. Additions to the herpetofauna of Kentucky. Herpetologica, 2 (2): 31-32. Fitch, H. S. 1936. Amphibians and reptiles of the Rogue River basin, Oregon. Amer. Hid. Nat., 17 (3)s 63U-6l?2. Fitzinger, 7,. J. I8k3. Systema isptilium. Amblyglossae. iii-vi, 1-106. Fowler, H. ,;., and S. n. Dunn 1917. Notes on salamanders. Proc. Acad. Nat. Sci. rhila., 69 » 7-28, pis. 3-U. Fowler, James A. 1951. Preliminary observations on an aggregation of Plethodon dixl . Herpetolopica, 7 (3)' 1U7-1L8. Garman, Harrison 189U. A preliminary list of the vertebrate animals of Kentuclsjr. liull. "3esex Inst., 26 (1): 1-63. Garman, Samuel 1 6U. The North American reptiles and batrachians. Bull. Essex Inst., 16: 3-U6, figs. 1-6.

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203 GiUiams, Jacob 1818. Descriptions of two new species of Linnaean Lacerta. Jour. Acad. Nat. Sci. Phila., 1: U60-b6l, pi. 18, fig. 1. Gordon, Kenneth 1939. The Amphibia and Reptilia of Oregon. Oregon State Mon., Studies in Zool., (l): 1-82, figs. 1-5U. Gordon, Robert . 19$2. A contribution to the life history and ecology of the plethodontid salamander Aneides a^neus (Cope and Packard). Amer. Midland Hat., U7 t3)« 666-701, figs. 1-6. Gordon, Sobert E., and Richard L. Smith 19U9. Notes on the life history of the salamander Aneides aeneus . Copeia (3): 173-17$. Graf, William, Stanley G. Jewett, Jr., and Kenneth L. Gordon 1939. Records of amphibians and reptiles from Oregon. Copeia (2); 101-10U. Gray, I. B. 1939. An extension in the range of Plethodon yonahlossee . Copeia (2): 106. Gray, J. E. 18 50. Catalogue of the specimens of ajnohibians in the collection of the British ftaseum. ?t. 2, J3atrachia Gradientia. London: 1-72, pis. 3-1*. Green, Donald M., and barren F, talker, Jr. 19!?U. A northern extension of the range of Plethodon richmor.di in Ohio. Copeia (1): 60. Green, Jacob 1818. Descriptions of several species of North American Amphibia, accompanied with observations. Jour. Acad. Nat. Sci. Phila., 1 (2): 3h8-358. Green, N. Bayard 1938. A. new salamander, Plethodon net tingi , from »»est Virginia. Ann. Carnegie f^us., 27s 295-299. 1939. The pygmy salar.ander Desraognathus yrr ighti King, on hite Top Mountain, Virginia"; Uopeia (i): U9. Green, N. Bayard, and N. D. Richmond 19UU. Courtship of Plethodon metcalf i. Copeia (I4): 256. Grinnell, Joseph, and C. L. Camp 1917. A distributional list of the amphibians and reptiles of California. Univ. Cal. Fub. Zool., 17 (10): 127-208, figs. 1-llu

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20U Grobman, Arnold B # 19\ih . The distribution of the salar and^rs of the genus Plethodon in east err) United Stai.es and Canada. Ann. H. I. Scad. Sci., h$ (7)i 261-316, figs. 1-11. 19b9. Some recent collections of Plethodon frorr Virginia with a description of a new form. Proc. Biol. Soc.
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205 Hilton, lUllM A. 19U6. Salamanders from northern California. Copeia (1): 16. 19U8. Salamander notes from the northwest. Herpetcloglca, h (3)* 120. Hoffman, Richard L. 1953. Flethodon welleri talker in Tennessee. Jour. fenn. Acad. Sci., >P (l)'i 56-87. Hoffman, Richard L., and Leslie Hubricht 1951u Distributional records for two species cf tq^hodon in the southern Appalachians. Herpetologica, 10 (2): 191193. Hoffman, Richard L., and H. I. Kleinpeter 19U8a. A collection of salamanders from Mount Refers, Virginia. Jour. Wash, Acad. Sci., 36 (3)* 106-108. 19U8b. Amphibians from Burke's Garden, Virginia. Aner. Midland Nat., 39 (3): 602-607. Holman, J. Alan 195?. Fall and winter food of Plethodon dorsaiis in Johnson County, Indiana. Copeia (?): 11*3. Howell, A. H. 1909. Notes on the summer birds of north Georgia. ;-uk, 26 (2): 129-137. Howell, Thelma 195U. Methods of counting costal grooves in salamanders. Jour. Tenn. Acad. Sci., 29 (1): U2-UU, fig. 1. Howell, Thelma, and Ann Hawkins 195U. Variation in topotypes of the salamander flethodon jordani melaventris . Copeia (l): 32-36, fig. 1-2,' ' JeAett, Stanley G., Jr. 1936. Notes on the amphibians of the Portland. Oregon, area. Copeia (1): 71-72. Kermode, Francis 1926. Accessions in Rept. Prov. Mus. Nat. Hist, for 1925: 35. King, iillis 1939. * survey of the herpetology of Great Smoky Mountains National Park. Amer. i&dland Hat., 21 (3)' 531-532, figs. 1-9, 1 map. Lachner, Ernest A. 19U2. An aggregation of snakes and salamanders during hibernation. Copeia (h): 262-263.

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206 Logier, E. B. S. 1932. Some account 3 of the amphibians and reptiles of British Columbia. Trans. Can. Inst., 18 (T): 311-336. Logier, S. b. i>., and J. C. Toner 1955. Check-list of the amphibians and reptiles of Canada and Alaska. Contr. Hoyal Ontario Ms. Zool. ': Pal eon. (Ul): k-v, 1-88, 77 maps. Lowe, Charles H., Jr. 1950. The systematic status of the salamander Plethodon hardii , with a discussion of biogeographical • robleits in Aneides . Copeia (2)a 92-99, fig. 1. 1955. The salamanders of Arieona. Trans. Kansas Head. Sci., 58 (2): ?37-25l, figs. 1-5. Martof, Bernard, and Robert, L. Humphries 1955. ObservAtions on some aiaphibians from Georgia. Copeia (3): 2U5-2it8. M&yr, Srnst 1931. Notes on Halcyon chloris and some of its subspecies. Aaer. &1us. Nov., Ui69): 1-10. Mittleraan, Nyron B. 19U8. American Caudata. V. Notes on certain Appalachian salamanders of the s»enus P lethodon . Jour. .»ash. ~cad. Sci., 38 (12): U16-419. 1951. American Caudata. VII. Two ner/ salamanders of the genus Plethodon . Herpetolofica, 7 (3): 105-112. Molir, C. E. 1937. Notes on cave vertebrates, troc. Penn. Acad. Sci., 11: 38-U2. 1952. The eggs of the zig-zag sala lander, Plethodon cinereus dorsalis . The American Caver: 59-60. Muchmore, William B. 1955. Brassy flecking in the salamander P.lqthodon c. cinereua , and the validity of lethodon huldae . Copeia (j): 170-172. Neill, Wilfred T. 19'-;8. An unusual variant of Pl c thodon t 'lutin orus in South Carolina. Copeia (b)t 2h 7-251, fig. 1.

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207 Netting, M. Graham 1936a. Vehrle' s salamander, Plethodon we hrlei Fowler and Dunn, in "est, Virginia. Proc.' *v. Va. Acad. Sci., 1C : 8993. 1936b. 'ehrle's salamander, Plethodon wehrlei Fowler and Dunn, in Pennsylvania, i'roc. I'enn. Acad. 3cii, 10* ^8-30. 1939. The ravine salamander, Plethodon richmondi Netting and Mittlaraan, in Pennsylvania. rj roc. Penn. Acad. Sci., 13s So-Sl. 19ii6. The amphibians and reptiles of Pennsylvania. Commonwealth of Pennsylvania, Board of Fish Commissioners: 1-29. Nesting, H. Graham, N. Bayard Green, and Neil D. Richmor"* 19U6. The occurrence of Wehrle's salamander, Plethodon wehrlei Fowler and Dunn, in Virginia. Proc. Biol. ~oc'. wash., 59: 1S7-160. Netting, Is. Graham, and M. B. Mittleraan 1938. Description of Pl ethodon richmondi , ? now salamander from west Virginia and Ohio. Ann. Carnegie 'ius., 27 J 287293, pl. 30. Neinan, .alter B. 195U. * new plethodontld salamander from southwestern Virginia. Herpetolcgica, 10 (1): 9-1U. iciolls, J. C, Jr. 1950. .Votes on the salamanders associated with Dosnognathus ocoee Nicholls. Jour. Tenn. Acad. Sci., 2£ (U): 31031h. Noble, G. K. 1927. The plethodontid salamanders; some aspects of their evolution. Amer. fctas. Nov., (2U9)« 1-26, figs. 1-10. Orton, Grace L. 19U6. The size of the slimy salanander. Copeia (2): 107. Paivcer, alcolm V. 1937. Some amphibians and reptiles from Reelfoot Lake. Jour. Term. A c ad. Sci., 12 (1): 60-66, figs. 1-18. 1939. The &:q?hibians and reptiles of Reelfoot Lake and vicinity, with a key for the separation of species and subspecies. Rept. lieelfoot Lake Biol. Sta., 3* 72-101, figs, l-lli. 19U8. A cortrlbition to the herpetolojy of western Tennessee. Jour. Term. Acad. Sci., 22: 20-30.

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208 Pope, Clifford H. 192U. Notes on North Carolina salamanders with especial reference to the egg-laying habits of Leurognathus and Pesmognathus . Amer. Jftxs. Nov., (153)* 1-15, figs.' 1-2. 1928. Some plethodontid salamanders from North Carolina and Kentucky with the description of a new race of Leurognathus. Amer. Mus. Nov., (306): 1-19, fig. 1. 195>0. A statistical and ecological study of the sala.-aander Plethodon ycn ahlossee . Bull. Chicago Acad. Sci., 9 (5): 79-106, figs. "1^7 Pope, Clifford H., and James A. Fowler 19U9. A new species of salamander ( Plethodon ) from southwestern Virginia. Nat. Hist. Misc., Chicago Acad. Sci., (U7): 1-U. 1 ope, Clifford H., and Nelson G. Hairston 19U8 . Two new subspecies of the salamander Plethodon shermani. Copeia (2)i 106-107. Pope, Clifford H., and Sarah P. Pope 19U9. Notes on growth and reproduction of the slimy salamander Plethodon £lutinosus. Fieldiana (Zoology), 31 (29): 251-261, figs.-"60^>5. 1951. A study of the salamander Plethodon ouachitae and the description of an allied form. bull. Chiacgo Acad. Sci., 9 (8): 129-152, figs. 1-6. Rabb, George B. 195l>. Observations on the identity of the salamander Pleth odon huldae . Copeia (3): 261-262. Rankin, J. S. 1937. An ecological study of parasites of some North Carolina salamanders. Ecol. Monogr., 7 (2): 169-269, figs. 1-15. Rhoads, Samuel N. 1895. Contributions to the zoology of Tennessee. No. 1. Reotilcs and amphibians. Proc. Acad. Nat. Sci. Phila., U7: 376-U07. Richmond, Neil D. 195 9 . First record of the green salamander in Pennsylvania, and other range extensions in Pennsylvania, Virginia and West Virginia. Ann. Carnegie Mus., 32 (7)» 313-318, fig. 1. Sager, Abrara 1858. Description of a new genus of perenni-branchiate amphibians. Peninsular Jour. Medicine, 5 (8): U28-lj29.

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209 Sanders, Ottys, and Hobart M. Smith 19U9. Some noteworthy records of amphibians in Texas. Trans. Kansas Acad. Sci., 52 (1): 28-29. Savage, Jay M. 1952. The distribution of the Pacific giant salamander, Dicamptodon en sat us, east of the Cascade Mountains. Copeia TT7 amp tod nto; Schmidt, Karl P. 192U. A list of amphibians and reptiles collected near Charleston, S. C. Copeia (132): 67-69. 1953. A checklist of North American amphibians and reptiles. Sixth Ed. Amer. Soc. Ichthy. & Herp.: viii, 1-280. Schwartz, Albert 1953. A new subspecies of the crowned snake ( Tantilla coronata )from the southern Appalachian fountains'. Herpetologica, 9 (3)« 153-157. 195k. The salamander Aneides aeneus in South Carolina. Copeia (U)« 296-298. Sinclair, Ralph M. 1950a. Notes on some salamanders from Tennessee. Iferpetologica, 6 (2): U9-51. 1950b. Some noteworthy records of amphibians and reptiles in Tennessee, herpetologica, 6 (7)* 200-202. Slater, James R. 1933. Notes on Washington salamanders. Copeia (1): ljiu 193U. Notes on northwestern armphibians. Copeia (3) : 1U01U1. 1939. Plethodon dunni in Oregon and Washington. Herpetologica, 1 (6;: 151i. 19U0. Salamander records from British Columbia. Occ. Papers Dept. Zool., College of Fuget Sound, (9)» U3~liU. 19U1. The distribution of amphibians and reptiles in Idaho. Occ. Papers Dept. Biol., College of Puget Sound, (lli): 78-109. 1955. Distribution of Washington amphibians. Occ. Papers Dept. Biol., College of Puget Sound, (16): 133-15U, figs. 1-2. Slater, James R., and V. C. Brown 19U1. Island records of amphibians and reptiles for Washington. Occ. Papers Dept. Biol., College of Puget Sound, (13)* 7U-77.

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210 Slater, James K., and J. Ifl. Slipp 19h0. A new species of Plcthodon from northern Idaho. Occ. Papers Dept. Biol., College of Pup;et Sound, (6): 38-b3, figs. 1-3. Slevin, J. R. 1928. The amphibians of western North America. Dec. Papers Cal. Acad. 3ci., 16: 1-152, pis. 1-23. 193U. A handbook of reptiles and ampld-bians of the Pacific states. Spec. Pub. Cal. A c ad. Sci.i 1-73, pis. L-73, figs. 1-9. Smith, Hobart M., and H. K. Buechner 19U7. The influence of the Balcones Escarpment on the distribution of amphibians and reptiles in Texas. Bull. Chicago Acad, Sci., 8 (l)s 1-16, fig. 1. Smith, Philip iff. 19U8. Noteworthy herpetological records from Illinois. Nat. Hist. Misc., Chicago Acad. Sci., (33)* 1-iu Snyder, Richard C. 19U6. Plethodon welleri from Flat Top ^fountain, North Carolina;; Copeia U): 17U , Stebbins, Robert C. 19U9. Speciation in salamanders of the plethodontid genus Knsatlna . Univ. Cal. Pub. Zool., U8 (6): 377-525, figs. 115, pis. 1-6. 195l. Amphibians of western North America. Berkeley: University of California Press, ix, 1-539. 195U. Amphibians and reptiles of western i>Jorth America. New York: ^raw-Hill Book Co., Inc., xxii, 1-528. Stebbins, Robert C, and H. C. Reynolds 19U7. Southern extension of the range of the Del Norte salamander in California. Herpetologica, li (2): U1-U2. Stebbins, Ro'oert C, and H211M J. Riomer 1950. A new species of plethodontid salamander from the J^raez Mountains of New Mexico. Copeia (?): 73-80, pis. 1-2. Stejneger, Leonhard 1906. A new salanander from North Carolina. Proc. U. S. Nat. aus., 30: 559-562, figs. 1-6. Stejneger, Leonhard, and Thomas Barbour 1917. A check list of North American amphibians and reptiles. Cambridge: Harvard University Press: 1-126.

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211 Storer, Tracy I. 1925. A synopsis of the Amphibia of California. Univ. Calif. Pub. Zool., 27: 1-3U?, figs. 1-U2, pis. 1-18. Storm, rtobert M. 1955. Northern and southern range limits of Dunn's salamander, Plethodon dunni . Copeia (1): 6U-65. Strauch, Alexander 1870. Revision der Salarandriden-Gattungen . Wfemoires de l'Acad. Imp. 3ci. St. Petersbourg, ser. 7, 16 (U): 1-110, pis. 1-2. Strecker, J. K. 1902. A preliminary report on the reptiles and batrachians of McLennan County, Texas. Tran. Texas Acad. Sci. for 1901, U (2): 95-101. 1908. The reptiles and batrachians of McLennan Countv, Texas. Proc. Biol. Soc. I'»ash., 21 j 69-4 3. 1915. Reptiles and anphibians of Texas. Baylor Ball., 18 (lt)s 1-82. 1935, The reptiles of tiest Frio Canyon, Heal County, Texas, baylor Bull., 3? (3): 32. Svihla, Arthur 1933. Extension of the ranges of some Washington Amphibia. Copeia (1): ^. Svranson, Paul L. 1939. Horpetological notes from Indiana. Amer. Midland Nat., 2? (3)» 68)^695, pis. 1-3, map. Thurow, cordon R. 1951. An analysis of variations in the pigmentation of Plethodon glutinosus . Unpublished iaaster's Thesis, Unive»sity of Chicago. 1955a. An albinistie individual of the sala mender Plethodon dorsalis . Copeia (1): 62-63, pi. 1. 1955b. Plethodon netting! in Virginia. Herpetologica, 11 (2): 102rioj: Tschudi, J. J. 18 38. Classification der Batrachier mit Berttcksichtigung der fossilen Thiere dieser Abtheilung. Mem. Soc. Sci. Nat. Neuchatel., 2s 1-100.

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212 Van Denburgh, John 1906. Description of a new species of the genus Plethodon ( Plethodon vandykei ) from kount Ranier, Washington^ Proc. Calif. Acad. Sci., h (!*)» 61-63. 1916. Pour species of salamanders nev? to the state of California, with a description of Plethodon elongatus , a new species, and notes on other salamanders . Froc. Calif. Acad. Sci., 6 (7)» 215-221. Villalobos-Dominguez, C, and Julio Villalobos 19U7. Atlas de los col ores. Buenos Aires » 7U pp., 38 pis. Halker, Charles F. 1931. Description of a new salamander from North Carolina. Proc. Junior Soc. Nat. Sci., Cincinnati, 2: U8-51. 1933. °>ome new amphibian records for Ohio. Copeia (U)» 22lu 193U. Plethodon welleri at -^hlte Top Mountain, Virginia, Copeia {h)'190. rtatney, uertrude S. 1938. A new record of Plethodon vehlculus (Cooper) from Vancouver, British Columbia. Copeia (2): 89. .Veller, ?/. H. 1931. A preliminary list of the sala-oanders of the Oreat Smoky Mts. of North Carolina and Tennessee. Proc. Junior Soc. Nat. Hist., Cincinnati, 2 (1): 21-32. Vied, !.5ax P. zu 1865. Verseichniss der Reptilien welche auf einer i:eise im nordlichen America beobachtet wurden. Nov. Act. Acad. Leopold Carol. Nat. Curios, 32: 1-lUU, pis. 1-7. #ood, John Thornton 19U5a. >varian eggs in Plethodon richmondi . herpetologica, 2: 206-210. 19U5b. Plethodon richmondi in Greene County, Ohio. Copeia (1): U6. 19U6. Additional records of Plethodor: richmondi . Copeia (3)* 169. 19U7a. Juveniles of Plethodon jordani Elat.chley. Herpetologlca, 3 (6): 185-181 , fig. 1 19U7b. Description of Juvenile Plethod on glutinosus sherraanl Stejneger. Herpetologica, 3 (6): lB8.~ 19U7c« Notes on North Carolina salamanders . Copeia (Lt): 27327U.

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213 Wood, John Thornton, and William E. Duellman 19U7. Preliminary herpetological survey of Montgomery County, Ohio. Herpetologica, I4: 3-6. >.ood, Wallace F. 193k . Notes on the salamander, Pleth odon elongatus . Copeia (h): 191. 1939. Amphibian records from northwestern California. Copeia (?): HO. right, Albert H., and A. A. right. 1938. Amphibians of Texas. Tran. Texas Acad. Sci., 21 J 1-38, figs. 1-6, pis. 1-3.

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211*BICGRAPKICAL ITEMS Richard Highton was bom on December ?U, 19?7» in Chicago, Illinois. He graduated from the Bronx High School of Science in New York City in June, 19h5. He received the Bachelor of Arts degree with a major in biology from the University College of Arts and Sciences, New York University, in October, 1950. He entered the University of Florida in September, 195l, and received the .'faster of Science degree in biology in June, 1953. He served as a graduate assistant in the Department of Biology at the University of Florida from July, 1952, to January, 195U. From February to June, 195U, he held a graduate fellowship. From September, 195U, to the present, he has been a teaching assistant in the Department of Biological Science. During the sunnier of 1955, he was employed as a Hanger-Naturalist by the Great Smoky Mountains National Park. He is a member of the American Society of Ichthyologists and Lerpetologists, the Herpetologists League, and Sigma Xi.

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This dissertation was prepared under the direction of the chairman of the candidate's supervisory committee and has been approved by all members of the committee. It was submitted to the Dean of the College of Arts and Sciences and to the Graduate Council and was approved as partial fulfillment of the requirements for the degree of Doctor of Philosophy. June U, 1956 SUPERVISORY COMMITTEE: Chairrn W//A^ ifJJ1. #