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The Distributional History

of the

Biota of the Southern Appalachians


A Symposium Sponsored by
Virginia Polytechnic Institute and State University and
the Association of Southeastern Biologists,
Held at Blacksburg, Virginia
June 25 -27, 1970

Edited by Perry a. IIolt

qJJith the Assistance of
Robert A. Paterson and John P. Hubbard

Research Division Monograph 4

Virginia Polytechnic Institute and State University
Blacksburg, Virginia

December` 1971

Distributional Interactions Among Eastern North American

Salamanders of the Genus P/cffroc7oce
Richard Highton

The largest salamander I amily, the Plethodontidae, has its probable center
of origin in the ancient Appalachian Mountain system of eastern North Amer-
ica. All of the more primitive genera (those with aquatic larvae) are still
restricted to North America east of the Great Plains where their lungless-
ness probably evolved as an adaptation to the mountain stream environment
(Wilder and Dunn, 1920; Dunn, 1926). The more advanced groups (in
two of three tribes in one of the two subfamilies recognized in the classifica-
tion o Wake, 1966) have dispensed with the aquatic larval stage. Complete
independence from the aquatic environment has enabled these two groups to
spread to western North America, Central America, northern South America
and Europe, across barriers that have apparently restricted the very successful
and abundant, but more primitive, aquatic breeding plethodontids to eastern
North America.

The type genus of the family, P/cJfroJoce Tschudi, is the genus with the
largest number of species in the LTnited States. Seven (tJczredyAeg., J7fcJJz.,

#orJo%G., d2J72%z., tJcfec.c"/"772, j`}or7%. and eJo7zgcr/ztJ) occur in the Pacific North-
west of the United States and adjacent Canada, and one (cecomcA%.cc7z%J) is
known only I rom the Jcmez Mountains o New Mexico. The remaining
thirteen species of the genus occur east of the Great Plains. Two of these
(ctZdoereJg.J and o%cfeG.e) are confined to the mountains o Arkansas and
oklahoma, and appear to be closely related to the eastern J)07zoJJcc. The
distributional interactions of the eleven species occurring east of the Mississippi
River is the subject of this paper; all occur in the southern Appalachian Moun-
tains. Two new species are described herein (P. feo#7"7z. and P. P%recZaZ%J).
Only one of the eastern species (cz.%e7.e#J) ranges extensively north of the south-
ern limit of the Pleistocene glacial boundary; the rest are largely restricted
to non-glaciated territory.

The eastern species were divided into two groups by Grobman (1944). The
salamanders in one group of smaller species (cc.73cre%J, rz.cfemorad2., fro//772dr%g.,

recJf.#gz., Jo7Ja/z.J and ttJe//erg.) are usually more elongate than the larger species,
although eve//cr. has a body form more similar to the eastern large plethodons.
Both f.#,c7.c2J and dorJa/z.J have isolated races in the ouachita and ozark high-
lands (P. c. j:er7Jz/J and P. d. 7?gz"Zz.c/t%.z") which are outside the area o
concern here. Three of the five larger species (#/%/c.%oJz", /'ord72c. and J;ocefe-
/a"cc) have a rather similar body form, but ouc'Ar/cz. and z/7zcZZz are more

elongated and tend to be somewhat intermediate between the larger and smaller
groups in other characters as well (Highton,1962b). P. /07!yz.crz" Adler and
Dennis (1962) is here regarded as a geographic color variant o P. Jo72crfeJoJJcc,
and probably represents an unusual population adapted to a somewhat differ-
ent (rock-crevice) habitat.

Groblnan (1944) showed that the distributional patterns of the two eastern
groups of P/cZfeodo%, are related and that there are several species pairs, each
including a large and a small species with similar ranges. Thus in both the
large and the small groups there is one wide-ranging species (g/a/Jz.%os#J and
cz.7zcrc2#) and other species whose ranges are much more restricted. The same
historical factors which produced isolation and speciation could have thus had
parallel results in both groups. More recent work has shown that there are
many dif ferences in the distributions of the syngeographs as outlined by Grob-
man. Much field work remains to be done before the distributions of all the
species discussed here will be thoroughly known. Our field work in the Mid-
dle Atlantic states (Pennsylvania to northern Georgia) during the last 15 years
has helped to clarify in detail the ranges of several Appalachian species and
has shown several new elements in their distributional patterns, especially in
the distributional interactions of closely related species.


One conspicuous dif ference between the two most widely distributed east-
ern large and small woodland salamanders is the marked seasonal variation
in their surface activity patterns. In the northern part of its range, P. J/z/c.-
7zoJz" first appears at the surface in early spring and is active during wet
weather throughout the summer and early fall. At the onset of sub-I reezing
temperatures it disappears f rom the surf ace and is rarely seen even during
warm spells in the winter. In the southern part of its range, it may be found
during wet weather throughout the year. Its ability to expand its range north-
ward into glaciated areas has been quite limited in comparison with that of
cg.#crcJ (figs. 1 and 2) and this may well be due to the lessened activity

period of g/"/z.7zoJz# in colder climates. P. cz.7z,ercz" is much more active dur-

ing colder weather in the northern part of its area of sympatry with F/z/f3.#oJzJJ.
During extended periods of above-f reezing weather in the winter, c3.72crc"s
may be found abundantly in the Middle Atlantic states. Conversely, cz.73cre2"
is much less active than g/z/fz.7zoJ#J during the summer months (except in the
mountains), even during cool wet periods of seemingly favorable weather.
Table I shows the relative seasonal varf ation in surface activity of the two

FIGuRE 1--Distribution of the seven species of eastern large P/cAodo#.

rr^BL_E 1=Seasonal cv_arial.ion_ in the relative abundance of Plethodor\ glndinosus and

P._ c;\nereus at Cunriingham Falls, Frederick County, Maryland, from Tq956-1971.

fin equfl amornt_of _collecting qJ)as not done during each month so the monthly
co.mP.arsons of afiut.d.aT.ce cwitPin a species are biased, but the relaticve frequen;y
of. the two species _(express.ed as a ratioJ indicates the s`easonal diff;re;ces ;n
the comaraticve surf ace aciiq)ity of the tqwo secies.

Month z s:-ou=====s=o h La Number Ratio

cineretts glutinosus.. cinereus

January |- - v\
March 143 .01
April 192 .17
May -+\+ 444 .28
June 163 .56
July 162 .68
A"g"st -+t r,- .60
September 60 .37
october 9 .14
November 127 .05
December 133 -o

species at a single locality in the Blue Ridge physiographic province o Mary-

land, at an elevation of 1350 feet. The seasonal activity of gJ#Jz.73oJe# at
Cunningham Falls State Park, Maryland, is typical of that throughout the
Middle Atlantic states (except in the southeastern Coastal Plain where g/z/-
!z.7!of2" is more active during the winter). The considerable surface activity
of cc.#crec" at this locality during the summer months is unusually high for
c!.crcwJ at lower elevations in the region. This locality was selected for in-
tensive I ield work over a long period for life history studies of the two species
(Highton, 1962a; Sayler, 1966) and for other work in progress because of
the abundance of the two species there and the presence of cz.72crcz/J at the
surface during the summer months. In spite of the unusual (for low eleva-
tion cz.7zcrcJ) summer activity, the differential seasonal activity of the two
species is clearly evident. It would probably be even more different at most

FIGURE 2-Distribution of the six species of eastern small P/c/AoJo#. One other spe-
Cies, P. #cocAf2.fcz#J, is extralimital.

low altitude localities in the Middle Atlantic states. The adaptations which
permit cz.7Gcrcz" to forage at the surf ace during cool weather are probably very
important elements in its successful colonization of the vast glaciated territory
of northeastern United States and southeastern Canada since the relativelv
recent retreat of the Wisconsin continental ice sheet (fig. 2). Such a specie.s
would seem better adapted for survival during a shorter I rost-f ree season in
more northerly areas. Conversely, its limited surf ace activity during very warm
weather may explain why its southern distributional limit is more northern than
that of y/#fz.7toJzJJ: a limit which may not yet be accuratelv known. P. c.73crcz"
has recently been reported in Alabama (Mount and .Folkerts, 1968) and
Louisiana (Keiser and Conzelmann, 1969), and mav be much more wideli7
distr!.buted in the south than is now known. There are few records froin
North Carolina and Georgia and none I rom South Carolina, although its pres-
ence is to be expected there. The fragmentation of the ranges of the small
plethodons is probably an indication of rather widespread recent contraction
of their ranges, particularly in the south (fig. 2). Much of the collecting in
the south is done during the warmer months, and cz.7zcrcz/J, active there pri-
marily during the cooler months, has apparently often been overlooked. It is
able to forage extensively during the summer at high altitudes in the south-
ern Appalachian Mountains and in the ncJrthern part of its range where sum-
mer temperatures are cooler.
Two c)ther small plethodons have similar seasonal activity patterns. High-
ton (1962b: 312) showed that /2o//77zcz73z. (under the name rz.c%77zo7zJz.) is less
active than g/z(Jz.77oJ#J in the summer than in the spring and fall at a locality
in Pennsylvania. Similarly, there are very few museum records for dorj`aJz.s
during the summer months. Although no quantitative data f rom throughout
the year are yet available, our field work indicates that the other large species
of eastern P/cffeodoce have seasonal activity patterns rather similar to those of
The eastern small plethodons, cg.72crc%f, dorJ/z.J and 7.z.cfe777ocedz., are largely
allopatric species, but there is no doubt that all are good species because each
occurs with the other two in one or more areas of overlap with little or no
evidence of hybridization. The three overlap in the Appalachian Mountains
and considerable new information on the zones of contact between c.%c/c?/J
and 7.cfr772o%dg. and between cz.7c/.e%s and f/orJ/z.I will be presented below. The
taxonomic status of populations previously assigned to rz.cfe772o%dz. is reconsid-
ered below.
The eastern large plethodons J/oce//oJJcc and etJcfrr/cj. are allopatric spe-
cies that as yet have not been taken together at the same locality (with
the possible exception of one cocferJcz. from Whitetop Mountain, Virginia, dis-
cussed in Highton, 1962b:322). Hoffman (1967) has not taken the two
together in southwestern Virginia. The remaining two species of large wood-

land salamanders, /.orJ7z. and y/"Z.#oJz/s, are very closely related and the dis-
tribution of both species is influenced by the other, particularly in the south-
ern Appalachians (Highton, 1970). The two species hybridize extensivelv
in the southwestern part of the range of /.07.dcz7zz., but not elsewhere.


It is desirable to reconsider the relationships of the small eastern woodland

salamanders before analyzing their distributional patterns, since there is diS-
agreement in the literature on their taxonomic arrangement. Grobman (1944)
suggests that cz.%crc#f and c7orj`/!.J are closely related and that tug//crz., rg.cfe-
fflocedc. and 7effG.7zg; form a second related group. Thurow (1957) and High-
tcm and Worthington (1967) have named two more isolated montane popula-
lions most closely allied to 73c/zz.#gz.. These are fectbrz.cfeJz. and Jfec72fl%Joarfe, both
apparently high altitude relicts, as is 73c#7.#gz.. Highton ( 1962b) considered %cZ-
fz.7zfz. and fa"brz.cfeZG. conspecific with rg.cfe772o#dG. and all of these close to cz.#cre"J,
but placed dorJ&/3.J and wc//c'rz. in a separate group and suggested that the lat-
ter two are probably not as closely related to each other as are the members
of the c!.ceere#J group. Thurow ( 1968) suggested that cecffc.cegc. and fe%Gr3.cfej.
are most closely related to oweJ/e7.G., while c2.cec/e%J, JorJa/.J and r.cfrfflocec7!. were
considered a second group. Thurow also felt that Jfrc7icrcadocfe is not a valid
form and tentatively synonymized it with cg.#crc#L but Jaeger (1969,1970)
has made an exhaustive study of the ecological relationships of j:rfec#Jocz/! and
cz.#crcz/J that he believed conclusively demonstrated their specific distinctive-
Since Grobman (1944), Highton (1962b) and Thurow (1968) are in such
disagreement as to the taxonomic relationships of these forms, the taxonomic
characters are reconsidered here (see tables 2 and 3 and fig. 3). The first
eleven listed in table 2 are those used by Highton (1962b) to suggest rela-
tionships in the group. One additional character, a physiological one, is listed
in table 2. This refers to the almost complete lack of response of gravid
females of r2.cfemoced!. and fro//ma%g. to injections of mammalian gonadotropic
hormones, compared with all of the other forms (except do7Jdr/G.J, not yet tested)
that readily deposit eggs after such treatment (unpublished data of the wrI.t-
er). Using all of these characters, the close similarity o 7z.cfronocedG., fro//-
maniJ .ne.ttin,?i, hqrichti, shenandoah a,nd cinereus fs aiparent, a,s we+l` as .tie
more isolated positions o JorJa/7.J and eueJ/cr.. Thurow (1968) dismissed some
of these characters and placed considerable emphasis on resemblances in body
proportions, a type of character I have usually avoided in the analysis of taxo-
nomic relationships in the genus because it is of ten strongly correlated with
the number of trunk vertebrae . Eastern small plethodons have interspecific
dierences and some also show intraspecific geographic variation in the pro-
portional length of the limbs and width of the head, but much of this varl.a-

JO!J3,sod (1uaSqt?) (1uaSqe) q}!JIAHOtZ'q o,}qS![s 9}t'JaPOur TStI!'}OdS }uE1)unqe

[ t?uns }u3Sqt: s)ods 3,!uJIA ]tI3!IS 3uas9Jd
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rySt,z-S!z T'uS!EJ}S Jos}unourt} paJ`H.I,lq 'ut'PUnqt:

S?10SJ,OP IO!J9]SOd 1U9S9|d Put:aP!A| |JIAOJJEu a}!HA 29 'uanbaJ
I(I:uns s}ods It3nba JO3uou
6T uB!ls _

/ /(,I tI1!JIAH0Elq
stopyair)ueJlfSt7oy3!wXO?10J;A-ZFIT8:PL abt e1|at JO!Ia)ue zJIA' JIgU }u8!tzJ'S
6I 93Jel uaSqt? ad!J)S 'tIS![s 3)!LIA Su!"ods }qg![s 3uou u3nbaJ

!1tlJ?Jqnl JO!J9,ut' zJIAOJJt:U

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9SJEI u9Sqt' q8!t,I]S 9d!I]S }qS![s 3t!VJIA 8u!] ods ]tIS![s 9U0u I(J9A
0Z s9J{ oJ{qur!J{lO}U9Saldz

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?unul!foq IZZ-TZ IO!I3tuE JHS!eI,

tI)!A Ht:[q 9t'J6POun 'UgPunqt! luanba.IJ
9SJt'[ AIIt3nSn 'u3qC AIOJ t2u 3d!J`S 3}!VA1 SUE,,ods auou

29}tIS!t,I S 1_Pa}EJ aS }tl'Punqt!

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3P!JIA ad!I)S It,nb9 S)tlnourt: H,e[qJ0 9}!tIJA29 3uou
o],qS![s T}U3nb9J:I

Jou!Jt u9rS!d3I0udot![eI -H9[J{StZIql?sJop}unorv

-aJd9IeunJ:ods }S3It}T ast?ldp9!IJso93us3Jd a6EtldJo pad!J}sun!Su J{quo!}sd!AoJu!}npul
HunJ]o3qun lt3poliI uo!}t?]uarS!duVOa]!VA !doJ]pt!uo8E!itunrt:u
u3IJA)9d!J}SOtI}P!A oJ{qur9JouI91}t!dPaH
PUE[SIt2tu3M q]a9}J{ t,Il!Xt,un
('uasaJd (,uas@Jd
S3IJt'A0 euounJoT
lion is strongly correlated with the number of trunk vertebrae. Thus geo-
graI)hic populations of cc.cecre2/J with the highest number of trunk vertebrae
tend to have smaller heads and shorter limbs. The same trends can be seen
in the various species of eastern small plethodons with different numbers of
trunk vertebrae. Figure 4, for example, shows what appears to be a very
significant dierence in the relative width of the head o Hawsbill Moun-
tain Jfecre%docfe (mean number of trunk vertebrae, 19.0) and a population o
cc.%cre2/J from nearby Culpepcr County, Virginia (mean number of trunk
vertebrae` 20.5). Yet another nearby population of c.#crc"J from Skyline
Drive.` mile 62.3, Greene County, Virginia, (mean number of trunk vertebrae,
19.9) has a head-to-body ratio closer to that of j`fecceorado/2 than to the
Culpeper County cz.racrc%J (fig. 5). Thurow's data show that samples o
cz.%"zo from Georiga where the species has a very high number of trunk

vert.ebrae have a relative head width even lower than all his r2.c/!772073df sam.
ples. It would therefore appear very likely that Thurow has based his taxo-
nomic arrangement of the eastern small plethodons on similarities in sev-
eral highly correlated characters. Pending data that clearly show there are
significant differences among the species in body proportions that are mostly
independent of the number of trunk vertebrae, I prefer to continue to use only
the latter character in order to avoid the bias resulting I rom treating correlated
characters as independent. Figures 4 and 5 were originally prepared for use
in the original description o Jfee7#cec704fe, but were not used when it became
obvious that the relative head width was not a good taxonomic character in
this group.I do not deny that the differences between the species that Thu-
row has found exist, but I feel that they are all so correlated with each
other, and with the number of trunk vertebrae, that they should not be given
the status of independent characters and then used to set up a classification
that ignores most other characters. It is possible that some of the taxonomic
characters I used are also correlated with each other. The amount of white
spotting on the belly and chin may well be correlated. The same could be
true of some of the features of the stripe; yet none are so often associated


I,.:;,.:, :, ;, !;./,; .,:i



FIGuRE 3-Phenetic relationships of the eastern smaH P/cfAocZo# based on the percentage
of shared characteristics (from table 3).

with each other as are the limb length, head width and body segmentation
characters used by Thurow.
Brodie (1970) showed that relative head width and limb length may vary
considerably between closely related species in western members of the genus

lO 20 30 40 50 60


FIGURE 4-Ontogenetic variation in relative head width of P/efAodo# 7zcJ!!.#g!. JAc##.

Jo4A from the Hawksbill isolate (indicated by triangles) and P/ejAodo# c.#c7c#J
from Culpeper Count.v, Virginia (indicated by circles). The regression lines were
fitted by the method o least squares.

O IO 20 30 40 50 60

FIGURE 5-Ontogenetic vaI.iation in relative head width o P/eAodo c.#ere#J from

Skyline Drive, mile 62.3, Greene County, Virginia. The solid line represents the
regression line for these data and the two dashed lines represent the regression
lihes of the two populations in figure 4.
150 RicHARi> HIGHTor\T

P/cf4oc!o#. He suggested that head size may be related to feeding specializa-

lions. The significant variation in relative head width and limb length be-
twecn species within all three species groups of western P/cJ%ocZo7z does not
support the use of this character by Thurow as a good taxonomic measure
of the relationships of eastern small plethodons.
Table 3 shows the number of characters that each of the forms has in com-
mon with all of the others (from table 2). With data on onlv twelve char-
acters, conclusions are certainly very tentative and subject to future revision
as more characters are discovered. The number of available characters is too
small for a good numerical taxonomic analysis, but a crude approach can be
attempted. The phenetic relationships of the eight recognized forms, based
on the percentage of characters each has in common with its most similar rela-
live is shown in figure 3. The most similar forms are rg.cfa772oceJJ. and fro#-
77z7zz., then Jbc7zcz73Joczfe, 7zefZ2.7j7z. and %`67.2.cfeZ2'. The next most similar form

is c7.72crc'z4J, with Jo7-`frJ//.J and awe//crc. less similar. This arrangement agrees
well with the phylogeny suggested by Highton (1962b), except that cZor`fJz.J
is a little closer to the c7.72cre%f group than to owc7J/crz.. New characters are
needed to better clarify the relationships of these species.
Since hybridization is expected to be more frequent between closely related
species, it is of importance that the only two reported cases of hybridization
among sympatric species of eastern small plethodons are between c.z.reerc%J and
ffre77czcedocrfe (Highton and Worthington,1967) and between c.#cJ.c%`f and rz.cfr-
7#0%d2. (Thurow, 1968). This may be considered additional support for a
phylogenetic arrangement similar to that suggested bv the phenetic relation.
ships in figure 3.
The taxonomic arrangement of the isolated alTopatric populations of dark-
bellied forms (rz.c/c77"77J7., #.cZfJ.#y7., fez{4rz.ffezz. and Jfec72cz7zo%) has been dis-

cussed by Highton and Grobman (1956), Highton (1962b), Highton and

Worthington (1967) and Thurow (1968). Since none of these forms are
sympatric with each other (or with feofJ."J&ca2.) it is impossible to apply the
criterion of intergradation to determine their biological relationships. All were
considered conspecific by mghton and Worthington although we suggested that
the separation of the three isolated montane forms as a species separate from
rz.cfe77207zJz. would be an equally satisfactory arrangement. With the apparent
separate specific relationship between r7.cfe77zo7talc. and an even more similar form
(feo#on%z., described below), it now seems more likely that the relationship
between rc.c/z777o#Jc. and 7zcZfz.re7z. should also be regarded as at the species level,
as suggested by Thurow ( 1968). I am therefore considering the c?.recre%f group
to Pe `compo:.ed. ot four sp?cies, cinereus, richmondi, hoff`'mani zLnd nettingt:.,
and the zuc//err. group to have two species, e4JeJJcrz. and dorJc7/G.a (Since I
believe that the taxonomic characters that JorJ/z.J and ztJc/Jer3. have in com-
mon are more 1.mportant I.n determI.ning phylogeneti.c relationships, I still regard

dorJ4r/z.J as more closely related to oue//crg. than to the members of the cc.#ercz

group, even though the total number of characters listed in table 2 shows Jor-
j`cz/z.J to be closer to cg.7zcrcc" than to zue/Jcr.). It could easily be argued that
the forms %cZz.73gg., fact4rz.cfeZz. and j`%c7ec77zJofe have reached the specific level
of divergence, but, for the present, I feel that their association together as
subspecies of a single species (7z,e#z.7zyz.) better reflects their phylogenetic rela-
tionships. This association is quite open to subsequent modification as fur-
ther information is obtained on the degree of differentiation of these forms.
It has been known for some time that the population heretofore assigned
to rz.c477zo73d. in the Valley and Ridge physiographic province of Pennsylvania,
Maryland, Virginia and West Virginia, diers from rc.cfr77zo%c7G. in the remain-
der of its range by having more white mottling on the belly, and especially
on the chin (Highton,1962b; Highton and Worthington,1967). Our field
work has revealed that this form is probably geographically isolated from other
rc.cfe772ocac7z. (fig. 6), although the two are known to occur extremely close to
one another on opposite sides of the New River in Virginia and West Vir-
ginia. I suggest that the consistent differences between the two are sufficient
to consider them distinct species. Since the Valley and Ridge form is unnamed,
it is a pleasure to name it for my friend, Richard L. Hoffman.

Plethodon hoffmani, rlow ST}eCjles

DIAGNosls.-An eastern small P/effeocZoce of the cz.#crc%J group with a modal
number of 21 or 22 trunk vertebrae (usually 22) ,. a dark belly with a mod-
Crate amount of white mottling,. and a chin heavily mottled with white pig-
ment. It differs from 7zefZz.7zyz., fez/brc.c%Zc., and j`%e7cz7zJoerfa in possessing more
trunk vertebrae and a more heavily mottled belly and chin. It diers from
c7.#`4rci/J in usually having more trunk vertebrae (sympatric populations of the
two species always differ by a modal number of one to three vertebrae), in
usually lacking the stri.ped phase (when present in feo#ov72z. the stripe is dif-
erent from that of cc'cee7e%J in being narrower) , and in having less white mot-
tling on the bellv. It differs from rc.cfe772oradz. in possessing more white mottl;ng
on the chin. wihere the known ranges of feo#77zace. and rz.cfr7"0%dc. are closest
(in the New River Valley), the two also differ in number of trunk vertebrae,.
rz.ffe772o%cZ2. has a modal number of 2l, and /!o//mc77zz. 22. USNM l35203, an adult male taken at Clifton Forge, Alle-
ghany County, Virginia, April, 1954, by Richard L. Hoffman.
4//ojyc: USNM 135204, an adult female with the same data as the holo-
Pcrr#f.+'cJ: USNM 135205, same data as the holotype; USNM 190220-3,
topotypes, collected 20 October 1956, by Richard L. Homan and Richard
Highton; USNM 127578L9, North Clifton Forge, Alleghany County, Vir-
gI.nia, 28 December 1946; USNM 133044, 2 miles north o Clifton Forge,
AIleghany County, VirginI.a, 28 April 1950,. USNM 127589, Lowmoor Mines,

Alleghany County, Virginia,1 June 1947,. CM 34992, topotype, 16 April

1944. (Unless otherwise noted, all specimens were collected by Richard L.
Hoffman. )
OTHER MATERIAL.-Specimens of feo#7ca#z. have been examined from all
of the localities shown in figure 6 (museum specimens are indicated by holr
low symbols, while living material has been examined from the remaining
localities, indicated by solid symbols). Geographic and individual variation
in the number of trunk vertebrae of fro//79ce;. are reported in Hfghton ( 1962b :
310 and in table 7 under the category o Ridge and Valley Province r!.cfe-
77zo7?dz.) and Highton and Jones (1965: table 1). Variation in the species,
based on new material collected during the last 10 years, will be reported in
a subsequent paper.
DEscRIpTIoN oF HoLoTYpE.-An adult male with a large mental gland of
the rz.cfe77307zJz. and cz.ceerc%J type (mghton,1962b: fig. 2C) and premaxillarv
teeth with enlarged anterior cusps (mghton, 1962b: fig. 3D). The length


F[coR# a:i.RneACr`dh:a..I `=.l~e:..h_O..On_ _h`Of`fT?a7i. and+I eth.odon Tichmondi in Pansylvaalia,

Maryland, West Virginia and Virginia. The dotted lines indicate the limits of
the Valley and Ridge Physiographic Province. Solid symbols represent my locality
records, ho]]ow symbols literature records aI]d ]oca]ities for lnuseum specimens.

I rom the tip of the snout to the anterior angle of the vent is 54 mm, to the
posterior angle of the vent, 58 mm, and the total length is l16 mm. The
length of the head (measured to the gular fold) is 10 mm and the width of
the head at its widest point is 6 mm. There are 21 costal grooves on both
sides (equivalent to 22 trunk vertebrae) and the vomerine teeth number 6
on the right side and 3 on the left. After 16 years of preservation, there
are no traces of the abundant brassy dorsal flecking present on all topotypic
indI.viduals in life. The belly is heavily mottled (with white pigment in life,
now represented by gaps in the melanophore bed). The abundance of white
chin mottling on the specimen in life is indicated bv the presence of melano-
phore pigmentation on less than half of the area of the chin in the preserved

DISTRIBUTIoN.-The Valley and Ridge physiographic province from the Sus-

quehanna River in central Pennsylvania southwest in the western two-thirds
of the Valley and Ridge province o Pennsylvania, Maryland, West Virginia
and Virginia to the New River. Its range extends westward into the Appa-
lachian Plateau physiographic province in two areas: along the valley of the
West Branch of the Susquehanna River in central Pennsylvania,. and in south-
ern West Virginia. Its range extends eastward across the eastern Vallev and
Ridge physiographic province to the top of the Blue Ridge in the Blue Ridge
physiographic province of Botetourt County, Virginia (see fig. 6).
VARIATION IN P. fro#77273z..-This species is remarkably uniform in appear-
ance throughout most of its range. Geographic variation is apparent in one
small area of its range. On Shenandoah Mountain, along the West Virginia-
Virginia state line, some populations of feo#772fJ#c. have a modal number o 2l
instead of the usua122 trunk vertebrae (Highton,1962b), and some individ-
uals possess a narrow dorsal red stripe (Highton and Jones,1965).
AFFINITIEs.-A comparison of specimens of feo#77z73G. with examples of rg.c.fe-
fflo7zJ. from any adjacent population in Pennsylvania, Virginia and West Vir-
ginia, results in good separation of the two species on the basis of amount of
chin mottling in all but the smallest individuals. On the other hand, west-
ern specimens of r.cfr772072dz. from ohio and Kentucky are often intermediate
between the two species in belly and chin pigmentation, a possible example
of character displacement (Brown and Wilson, 1956) that needs further inves-

Besides the trunk segmentation dif ference mentioned in the diagnosis, there
are other minor dierences between rz.cfr772o#cZ?. and feo#7reczcez. in the New River
Valley. Southern rz.c.faffloceJz. tend to have a greater average amount of dorsal
brassy than do foo#77272c., although there is considerable overlap in the
range of vari.ation. Red pl.gment is often present in both species, but it is usu-

ally confined to the dorsum in fro//7##f. (in the region where the narrow
stripe appears in some individuals of that species), while in southern rc.c%77?07zdc.
the red pigment occurs more frequently on the cheeks, front legs and anterior
sides. The geographic distribution of feo#7#cz7ez. and rc.c%7#o#Jz. and their taxo-
nomic relationship is discussed further below.
The ranges of seven of the I ifteen combinations of the six species do not
over+ap or meet. rThese hoffmani and nettingiJ hoffmani aLnd dorsalis, ho ff-
mani 2Lnd welleri, n.etlirigi zmd dorsalis, nettingi and welleriJ nettingi zLnd rich-
mondi, and dorsalis a,nd welleri. rT`he raLng;es of hoffmani zLnd richmondi cone
within less than a mile of each other on opposite sides of the New River in
Virginia and West Virginia, but they have not yet been taken together. The
range of fro#772&#z. also closely approaches that of both P. 72. 73efJz.%gz. and P. 7z.
frzJG7.a.cfaf7., so that it is possible that further field work may show some con-
tact or overlap in their respective ranges. The wide-ranging cz.72L?rGz/J occurs
with all of the other five species, and we have found it together in the same
habitat with all five in the Appalachian Mountains (including all three sub-
species of nettingi).

INTERAorIONS BETWEEN P. 7zcJZz.7g7z. AND P. cz.72crezJJ.-The three isolated

montane populations, #cf}f.aeg7., fe"arc.cfefz. and Jfrcra73doczfe, are clearlv relicts of

a formerly more widespread species (fig. 7). In the Cheat Mountai-ns o West
Virginia, 73cZfz.73G7z. is almost always found asso.ciated with spruce forests ( Brooks,
1948) which are relicts of the formerly more widespread northern coniferous
forests. In some of these spruce forests 77eZZz.#gc. appears to exclude fc.7ce/C#J,
the only other small P/f'Z4oJo~ in the area, but in most localities the two
are found together, Figure 8 shows the localities where we have taken the
two s_Decies in the Cheat Mountains and vicinity. A detailed ecological study
of the two species in that area would be welcome. It would appear that %cZ-
fz.#gc. is unable to successfully compete with cc.72erc%J in the deciduous forests
of the Cheat Mountains and has become extinct over most of the region.
Highton and Worthfngton (1967) found that j`%c73c7zJofe is restricted to
talus slope habitats on the northwest slopes of three of the h;ghest mountains
in Shenandoah National Park in the Blue Ridge province of Virginia. On

NOTE: Thurow (1968:20) published a figure which he interpreted as showing that

the geographic variation in belly mottling in P. rz.cA77207zd. suggested in my figure
(Highton, 1962b: fig. 30) docs not hold for southern /c.cA%072Jc.. Yet he claimed
(1968:23) that some of these same individuals are intermediate between rz.cA77zo7zdz.
and cG.%%" It is not at all clear how supposed hybrids between the dark-bemed
/2.cAcoo#Jz. and the mottled-bellied cz.cee7.c%J show that pure Tennessee /3.cAfflo#dc. are
not dark-beI]ied (with reduced white mottling compared with Ao//ovo7zz').

two of these mountains (Hawksbill and Stony Man) it appears to exclude

c.%crc#J, while on the third (The Pinnacle), the two are found together
throughout the entire talus slope. Jaeger (1969,1970) reported on a detailed
study of the ecological relationships of Jfec73cz7zJofe and cz.7crcJ on Hawksbill
Mountain. He showed that both species have similar habitat preferences for
moist soils, but that j`fec7z72dofe is more tolerant of dry conditions than cz.7zcrczJJ,
which is excluded I rom the drier talus slope habitat by insuicient moisture
rather than by the presence of j`%c7zcz7zdo4r%. On the other hand, cz.eJ does
exclude Jfec7zcz7Jo% from the surrounding woods by its better ability to com-
pete for limiting environmental resoui.ces, primarily food. Thus Jfec#'72dofe
survives only in a suboptimal habitat where cz.7zcrez/f cannot penetrate. When
the talus slope becomes invaded by soil, c2.#ercz/`t enters, and the ultimate result
probably will be the extinction of j`fec777zJofe as the talus habitat disappears.
Figure 9 shows the distribution of the two species in the Shenandoah National
Park o Virginia.
The ecological relationship of fr%6rz.cfefz. and cz.73c#J needs study. We have
taken %zJbrc.cfeZg. in abundance at five localities along the top of the Blue Ridge
Mountains in a distance of 4y2 air miles. P. cz.7zc'rc#J has been taken at only
one of these localities, but it appears to be less abundant there than is fezJZ7rz.cfe/z.

(10 c.7zerc%J: 33 A2tG7z.cfr}z.). It would seem that fe#Grz.cfeJz. is much less re-
stricted ecologically than either 73cZ.#g3. or .ffec%cedoftfe but its very limited
range may be indicative that it, too, is able to survive in competition with
cz.#c'rc2/J only under unknown special conditions that occur at high elevations
in the Peaks o Otter area. Perhaps even now fe#Grz.cfefc' is being displaced
by cc.72erc2JJ. It is not restricted to the talus slope habitat and there are no
spruce forests in its range. Fl.gure ]O shows the localities where we have
taken the two species.
All three of the forms currently assigned to the species P. 7zcJfz.77,p/. are obvi-
ously relicts of a more widely distributed form now displaced bv c3.%cre%J, eX-
cept in marginal habitats where it still survives. We have searched for other
populations at high altitudes in several other mountains in the Blue Ridge,
Valley and Ridge and Appalachian Plateau physiographic provinces o Vir-
ginia. and West Virginia without success. However, at some of these places,
ofcaslonal cz.73crc%J have unusually dark bellies, perhaps an indication that
cz%"% has "swamped out" a formerly existing population of #c#z.7zgz. through
hybridization. In the vicinity o Cow Knob, in the Shenandoah Mountain`s
of Pendleton County, West Virginia, and Rockingham County, Virginia, occa-
sional feo#77zzzz. also have unusually dark chins and bemes. This could hardlv
be the result of introgression from r2.cfr77207zJz., since this area is on the eastern
edge of the range of Ao#77zcz7zz.,. so, it may also be the result of genetic influ-
ence of a preexisting population o %eJJz.%4z. in that mountai.n range, in this
case in fro/J-7724#j. rather than c7.72czc%^ Evidence of hybridizatI.on between


/ /-
/ 780

^^ N ETTINGl /
I `l
r-. `\--_`/,/



FIGuRE 7-Locality records of the three subspecies of P/c!feodo #c#z.p. in Virginia and
West Virginia.

fz.crcacf and Jfec7z%Jocrfe at the Hawksbill isolate of j`fec7zdrJocfe was presented

by Highton and Worthington (1967). This possible influence o formerly
existing %cZ3.grz. populations on the present high mountain populations of
cz.7zcrc#f and feo#772fl7zz. is much in need of further investigation.

INIBRA.c"Ot`s BBrrWBBr+ P. and P. hoffmani.-:The grea.ter Pa.rt

of the range of 4o#77t72z' ;s not inhabited by cz.7zc'rc."J (fig.ll). In most parts
of the range of fro#mcz#z., c2.#c7.c%J occurs with fro//zB4#. on1;' at the eastern

and western edges of the latter's range. There are, however, three areas where
there is considerable overlap in the ranges of the two species. The first of
these is in the northernmost portion of the range of fro#773drce. in central Penn-
sylvania. At several localities, well within the range of fro//z73G., cG.#ere%J has
been taken in abundance. There is a considerable area in southwestern Clin-
ton County and in northeastern Clearfield County in the Appalachian Plateau
physiographic province of Pennsylvania where we have not taken cz.7ze/e%j`. It
is surprising that feo//7724r7z. should exclude cz.crc#J from this particular part
of the plateau when cG.73cre%J occurs to the eastern edge of the plateau province
in all other areas at the western limit of the range of feo#7%ace.. There is no
striking ecological dif ference in this section of the plateau that would explain
this unusual feature of the geographic interaction of the two species. P. fro//-
7;#ceJ. has apparently penetrated the range of cG.%ere#J along the valley of the
West Branch of the Susquehanna River for a considerable distance.
A second area where cg.7.crcz/J is regularly found within the western part of
the Valley and Ridge physiographic province in sympatry with fro#m47z. is in
the high area in the vicinity of Monterey, Highlands County, V;rginia, and
adjacent counties. It would appear that c?.%erc%J has entered the range of Ao//-
7"a". both I rom the east and from the west. This can be determined by the

FIGuRE 8-Distributional interactions o PJeAoJoce c.cecrc%J and PJeAodo% #cf!c.#g!.

#eJJ!.#grG. in West Virginia and Virginia. Only my records are included. The 4000
foot contour line is indicated.

differences in the cz.72crcz" populations of the Appalachian Plateau and those

of the Valley and Ridge province. The cz.72crc%J of the plateau are all of the
striped phase and usually have a modal number of 19 trunk vertebrae, while
those of the Valley and Ridge contain both color phases and usually have a

FIGURE 9-Distributional interactions of P/c!AocZo7z cz.7jcrcc" and P/cfAocJo7z 72c#;7z4z. JAc'#-

47Jo4A in the Shenandoah National Park o Virginia. Only my records are in-
cluded. The 3500 foot contour line is indicated. The north, east, south and west
boundaries of the map are 3840' N ]at. 7815' W long. 38a25' N lat. and 7830'
W long.



FIGURE 10-Distributional interactions of P/cfAodo# c3.7?crcwJ and P/cZAodo7c 7zc#z.7zfz.

AwZJr2.CAJ; in the Peaks of otter region of Virginia. Only my records are included.
The 3500 foot contour line is indicated. The north, east, south and west boun-
daries of the map are 373333' N lat. 7925' W long. 3725' N lat. and 7940'
W long.

modal number of 20 trunk vertebrae (Highton, 1962b). The cz.#c'rcc" that

entered i rom the east can easily be distinguished I rom those that have pene-
trated the range of feo#77zcz7#. from the west, and it is apparent that both
types are present in this area. Usually, fro//7#czz. is found together with c.77crcz/a
in this region.

The third area where the ranges of the two species overlap is at the extreme
southwestern limit of the range of %o//7727z.. In this region feo#777czz. and
cz.7zcrcs are sometimes found together at the same locality, but we have found
only one or the other species at a number of places. This is apparently the
only other section where feo#7J2dr#z. has been able to penetrate into the Appa-
lachian Plateau physiographic province for more than lo miles.

In any case where two closely related species exclude one another over most
of their ranges, it is of interest to determine which one is expanding its range
at the expense of the other, or i, indeed, the contact zone is stationary. The
range of feo#7727. is almost completely surrounded by that of cz.#,crez/J, a fac-
tor which might lead to speculation that fro//mzzz. is retreating at the present
time The detaild nature of th contact zone should be studied I.n several
areas. both where there is considerable overlap and where there appears to

.8.--.8.=to . . o::
a oO -

FIGuRE ll-Distributional interactions o P/cZAodo% c2.#erc%J and P/efAodo% Ao#coa#Z

in Pennsylvania, Delaware, Maryland, West Virginia and Virginia. Only my
ords are included.

be very little overlap in the ranges of the two species. Long-term field studies
may provide valuable information on the stability of the contact zone. At
present, however, it is of interest to look for isolated pockets of one species
or the other as possible indications that one species is expanding its range at
the expense of the other. We have taken one species well within the range
of the other at only two places (4 miles south o Moorefield, Hardy County,
West Virginia, and at Wells Tannery, Fulton County, Pennsylvania) ; and
at both, it is cG.z}erez4J that appears to be surrounded by froJ:/onace.. Whether or
not this me.ans that feo#7cecec. is presently extending its range at the expense
of cz.7zercz" is not yet established, but this possibility should be investigated. The
Valley and Ridge province, where fao#on&72c. occurs, appears to be generally a
drier region with better drained soils than adjacent physiographic provinces.
Particularly in the summer, the region is much drier than the higher Appa-
lachian Plateau to the west and the Blue Ridge to the east. It is therefore
possible that cJ.ceerc%J is excluded from this area by the drier or by

the presence of feo//77zcz7gz., or by a ccmbination of both. It is of interest that

cz.#c'rcz" is apparently able to penetrate into the western part of the Valle}`
and Ridge province - otherwise occupied exclusively by flo//fflcz7%. - only in
the higher moister areas in the central part of the range of feo#77?cz7zz..

Although feo#772cz. and cc.7zcrcJ have been collected together at 40 locali-

tiesj only at one place (in Augusta County, Virginia) were a few salamanders
taken which appeared somewhat intermediate. Even if these are hybrids, the
two species appear to overlap in most areas without any indications of inter-

Extensive data on the geographic interactions of %o#77.7zz. and rz.cfe77zo72cZc.

with c!.cre"JJ and with each other, have been obtained during the course of
a 15-year field study on microgeographic variation of P. cz.7zcrc#s in Pennsyl-
vania, Maryland, Delaware, Virginia and West Virginia. Our procedure is
to attempt to collect a series of 25 or more individuals of" at each
locality where we find it. Thus far, we have obtained samples of 25 or more
cz.7cre"J from over 1200 localities in the above states, and a smaller number
from almost 400 additional places. Early, it became obvious that only feo#-
773cz7zj or rz.cfe772072cJG. were present in some regions so that cz.crc#f was not dis-
tributed throughout the entire area of the study. These facts have made the
survey of microgeographic variation more interesting because of the isolation
of certain cz.73ercz populations by the other species, and have also provided
valuable data on the distributional interactions between the three species pre-
sented here. The distributional work is not yet completed, but the major
patterns appear to have been determined.
P. fz.#ercacf is usually much more abundant than either rz.cA7#077Jz. or feo#-
onar7zz.. However, I feel that if either of the rare species is present, we should
have found it while spending the time to search for 25 c.c.72crez/5`. Undoubtedly
we have missed the rarer species at certain local;ties, but it is hoped that the
number of localities from which collections were taken in each physiographic
province is suf ficient to fairly accurately determine the ranges of the three

Throughout the area (Pennsylvania, Delaware, Maryland, Virginia and

West Virginia) we have been successful in finding c3.%erc%J at, or very near,
all but two of the places where it has been previously collected (based on
literature records and records of museum specimens). Thus, only my own
locality records are included on the distribution maps involving cz.7crc#f in
the above states. The only two places where cJ.72ere%J has been reported where
we were unable to find it are at Gloucester, Gloucester County, Virginia (Cope,
1889). and just north o Cumberland, Allegany County, Maryland (Keller,
1945). Thus far, we have been unable to find cc.z2crc2/J anywhere on the
Coastal Plain peninsula between the York and Rappahannock rivers in south-


eastern King and Queen County, and in all of Middlesex, Gloucester and
Mathews Counties, Virginia. Cope's record is therefore in need of confirma-
lion before being accepted. The specimens on which Keller's record is based
have been eaxmined by the writer,. they are all feo#772%c., so this record should
be deleted I rom the natural range of cz.7zcrczJJ.

P. cc.rocrc%J is common in the southeastern Coastal Plain o Virginia, but

its distribution in the Piedmont of southern Virginia appears to be spotty. It
has not been found at many apparently suitable places we have collected. This
is close to its southern range limit, and it may have become extinct over much
of southern Virginia.

As mentioned above, r.c/37#ocec73. and fro//7re4rec. are often rare and diicult
to collect, especially during the summer months. Literature records and rec-
ords of museum specimens from the above states do add considerable informa-
lion on the distributions of these two species, and so these records are included
(indicated by hollow symbols) in figure 6, but not in figures ll and 12. The
reason they have been omitted from figures ll and 12 is that these figures
concern the interactions of the two species with cz.72erez", and museum speci-
mens and literature records usually provide no data on whether or not cz.7zcrc%J
is also present at the locality along with rz.cfeono#dz. or feo#77z#c.. Therefore
only our own records, that provide this information, are included. The same
is true for figures l3, 15, 16, 19 and 20. Literature and museum records for
certain species are included in figures 7, 14, 17 and 18.

IN`TBRACTlorIS PBrrYBBN P. hoffrnani and P. richmondi.-Ffgnre 6 Shows

the known records of feo#ffl722. and rz.cfa772o%d. in VIrginia, West Virginia,
Mrayland and Pennsylvania. The two have never been taken together. Their
ranges come closest in the New River Valley in Giles County, Virginia, and
Monroe` Summers and Mercer counties, West Virginia. Here we have found
only %o#77?#z. east of the river and only rz.c%772o#dz. west of the river, I.nclud-
ing localities on both sides of the river within a half mile of its banks. The
fact that the ranges of the two forms come so close without any evidence of
interbreeding between them at this point has been a major factor in the deci-
sion to regard the two as distinct species. Rivers are not normally barriers to
the dispersal of any species of the genus P/effrodo#. Thus all the species that
occur on either bank of the Mississippi, Ohio, Hudson, Columbia, and most
other major rivers, also occur on the other side. P. rz.cAmo%d2. itself occurs on
both sides of the New-Kanawha River system to the west where this river is
wider. It would thus appear that major rivers are rarely barriers to the dis-
IJersal o P/effeodo#, except in this particular case. It would seem likely, then,
that the presence of the closely related form on the opposite side of the river
is the factor which prevents the species from crossing - rather than the rl.vcr
itself acting as a barrier to the dispersal of the two forms.

The ranges of the two species also approach each other rather closely in
two other areas. We have taken r!.cfr7ceoradc. at Cowen, Webster County, West
Virginia, only 27 air miles northwest of the nearest fro//773a!ce. locality at See-
bert, Pocahontas County, West Virginia. Records of feo#fflcz7z.I. near Wilpen,
Westmoreland County, and I rom near Kittanning, Armstrong County, Penn-
sylvania, are both only 33 miles I rom the nearest records of rz.cfe77zo%cJ3. in the
vicinity of Pittsburgh, Allegheny County, Pennsylvania. In both of these areas.
we have found only cJ.#crcz(J in the intervening regions (see figs.ll and 12).
Further search is continuing in the hope of finding localities where /2o//772c%z.
and rz.cfe77zo7zcZz. are in contact.


o oo oa:oo. o
OOO oO. oooo




^ o = _A.a . i i.i*e-,,',,'``.---->i:
I I Ag a _. :..i.,,-,:
. . . co.;:..:::~.:..._..#ro: a .a:S*..::.``*:S
: . _..:.:.:..`.`f ;,,_.;iJ.;:_.;a:_._`.:.=o h` LZt

A ;..-.:..:.ey.ct:i /.i
^ _ ._out ::
I .a _ .ooa a

)``, \+ . :.:o
oo oo o a.g`'o..Oooi
'`,`\-I I A&O`^o
-? I
S . o a
`';.::-;'''kfffftyr ao:,a

a a
oO ^

---o-:5-i_ ^`..

FIGURE 12-Distributional interactions of P/e"7odo# c2.%cre%J and P/c!Aoc!oce r2.cAaco#d;

in Pennsylvania, Delaware, Maryland, West Virginia and Virginia. Only my
records are included.

One literature record o 7c.cfa77207zd3. or fao#on4cez. deserves comment. This is

a specimen (CM l3939) supposedly collected at Lake Terra Alta, Preston
County, West Virgi.nia, and indicated as a rz.cfe77?oredc. record on its distribution
map in Highton (1962b). This unstriped specimen was originally catalogued


83 82 8l.



/ A A
` _L\ `,_


,i.-. '-,i LEI

.``.L./: . : .

FIGURE 13-Distributional interactions of P/cj4oJo# cz.7zcrc#J and PJffAocJo# rc.cAaco#df

in northwestern North Carolina and northeastern Tennessee. Only my records
are included.

as cg.73crczJJ, and it has a typical mottled cz.72erc"J-Jyc belly. I identified it as

rc.c.fe7#o#dG. because of its very high vertebrae count. In that area of the high
Appalachian Plateau o West Virginia, cz.erc"J occurs only in the striped phase
and usually has from l8 to 20 trunk vertebrae, while this individual has 22
trunk vertebrae. I concluded that it could not be cz.73erez/f even though it has
a belly typical of that species. I now think it is more likely that the speci-
men is one of cz.%ere%J I rom somewhere else and docs not have correct locality
data. Lake Terra Alta is a locality where several herpetologists have doric
extensive field work, and no other specimens of rc.cfeono7zdG. or fao#772cec. have
ever been taken there. I suggest that this record needs confirmation before
being accepted as a valid record,. therefore, it is not included on the distribu-
tion map (fig. 6).

INTERAcTIONS BETWEEN P. rz.c.fe77zo7zJz. 4r%d P. cz.%erc#J.-There is rather

broad overlap in the ranges of these two species in southwestern Pennsylvania,
West Virginia and southwestern Virginia (fig, 12), in northwestern North
Carolina and northeastern Tennessee (fig. 13) and I.n most of the state o

83 82

A +^

LroL ^ I
A _+L^ ^
I /:-ivL{ `





FIGuRE 14-Ditributional interactions of PJeAOJoce rz.cAcoo#dG. and P/cAoJo# coc//ere.

in southwestern Virginia, northwestern North Carolina and northeastern Tennessee.
Only my records of rG.c/zaco#d?. are included, but all known localities of qx/e//cr;.
are shown.




j.::. + ~.

` `--.-/ . ,

1 I,
I ,

830 82.
FIGURE 15-Distributional interactions o P/efAodo# c.ceerc%J and P/c/Aodo# dorJ.a/G.J in
eastern Tennessee and western North Carolina. Only my records aI'e included.







J .{

: `?RE.:.` _.-`:


FIGURE 16-Distribution of the three phenotypes o P/eJAoc7o# #/%f3.#oJ#J and their

intermediates in Pennsylvania, Maryland, West Virginia and Virginia. Only my
I.ecord are included.

Ohio (fig. 2). In ohio, Duellman (1954) and Seibert and Brandon (1960)
often found the two species together, but at a number o localities only one
was taken. The same type of ecological relationship appears t.o exist in the
Appalachian states, for we have taken the two together at 31 (44 percent) o
our rc.c.fe77zo7zdg.localities and r/.c.fe77207zJc. only at 39 (56 percent) places in the
area where the ranges of the two species overlap in Pennsylvania, West Vtr-
ginia, Virginia, Tennessee and North Carolina. Thus, a somewhat dierent
pattern of ecological interaction would appear to be involved in the areas of
geographical overlap between rG.cfe772072dg. and cc.ceere%J compared with fao#mre3.
and cz.cec~e#J. In the case of fro//7727z. and c.#c7e%J, the two usually exclude
one another from their respective ranges (except in three areas discussed above),
while there is generally considerable overlap in the ranges of rz.cfefflofflcZz. and
cG.7zerez/J in their zone of contact. We have never taken animals that appear
to show introgression between cfcecre2JJ and #.cfe772orodG., although Thurow
(1968) reported a population containing apparent hybrids between the two
species jn northeastern Tennessee.

Our field work in Pennsylvania and northern West Virginia has indicated
that cz.7zcrcz" is usually much more abundant than rg.c%77zo72dz.. It therefore
might appear that cz.73crcz" is able to utilize the available resources of the en.

: ;r.ounl:t: :: :eonrsei t; I `aitci:n::? lt.hcaaTi t:::.fe onfira.d;e : :r: scianncet h;I,ii!Ly. ce-,: i :st ai:u aal I; i gr:: :
whether or not c.z.7zerc'%f is also present, it would seem more likely that there
is some factor in the population biology of northern rz.cfa772073Jz. that apparently
keeps its population density much lower than that of most other plethodons.
In southern West Virginia, southwestern Virginia and adjacent North Caro-
lina cind Tennessee, rz.cfe772o#c7z. is far more abundant, and the density of its popu-

FIGuRE 17-The distribution of P/cfAodo# qLueA//cz. and P/cZAodo# ##cf%J in New

York, Pennsylvania, Ohio, West Virginia, Virginia and North Carolina. My
records are indicated by solid symbols and those of literature reports and museum
specimens by hollow symbols.

lations there often appear to be as high (sometimes even higher) than those
of sympatric c.recrcztJ. Quantitative comparisons of geographic gradients in rela-
tive abundance of sympatric species o P/eZfeodoz are much to be desired,


FIGuRE 18-Distribution of P/c/AoJo# J'07g4A/oJJcc and its interactions with P/eZAodo#

gr/z/fz.#oJ#J and P/cZAodo# jorcJ7g; in southwestern Virginia, northwestern North
Carolina and northeastern Tennessee. The 3500 foot contour line is indicated
Except for literature records for JJo#4A/oJJe4, only my records are included.

INTERAcTIoNs BETwEEN P. rz.ffa`mo#d; AND P. dorj.a/c.j`.-The geographic

contact between these two species occurs west of the Appalachian Mountains
in the Interior Low Plateaus physiographic province. I have done no field
work in this area of contact, but records of museum specimens indicate a rather
broad area of overlap in Kentucky. Dr. Roger Barbour has told me that he
has found them together at several localities in the vicinity of Lexington, Kenl
tucky. P. dorJa/c.J apparently does not enter the Appalachian Plateaus physio-
graphic province in eastern Kentucky,. but farther south, in Tennessee, it has
extended its range eastward across the Plateau and the Valley and Ridge prov.
inces to the western edge of the Blue Ridge province. Ccmversely, rc.cfe7raocad3.
extends westward into the Bluegrass section of thes Interior Low Plateaus
physiographic province o Kentucky, but has not been taken in the Cumber-
1and Plateau o Tennessee. The ecological interaction between these two
very different species in thef r area of overlap needs study.

INTERACTIONs BETWEEN P. r!.c/}mocadZ AND P. z~eJJcr..-These two species

are among the most different of all the eastern small plethodons. Their ranges

FIGURE 19-The distribution of P/efAodo# jo/d#G. in the southern Appalachian Moun-

tains. The 3500 foot contour line is indicated. Only my records are included.
Localities of hybrid populations between jorda7G{. and 7/#.#oJ%J in the Nantahala
Mountains of North Carolina and Georgia are omitted.

overlap in southwestern Virginia, northeastern Tennessee and northwestern

North Carolina. Because of the rarity of owe//erg. or perhaps because of its
spotty distribution, we have little information on the interactions, if any, be-
tween these two species. The two occur on Whitetop Mountain and near-by
Mount Rogers, Virginia, but were not taken together by Dr. James organ
(personal communication) during his extensive field work in that region. In
that area, eueJJcr2. appears to be restricted to high elevations and rz.c/tonocadj.
to lower altitudes.
In eastern Ten.nessee, cocJJerz. also occurs at lower elevations. The only local-
ity where eve/Jerc and rG.cfefflo#d2. have been taken together is the place where
Hoffman (1953) first reported owe//c~ at a low elevation in Tennessee, near
CardervI.ew, Johnson County. Figure 14 shows all known wc'//f'r. localities
and my records of richmondi,

INTERACTIoNS BETWEEN P. c{.raerc?/J AND P. Jo7j`/2.J.-The ranges of these

two species overlap in several widely separated areas. These include areas in

|I +
\ \
. -


+ .
~ .
`.,..... , . .

.... -.
.. ...I ..

\.`. . . I....


. ,. ,:. .

.-a. .
i.'i#. :. II



35 I

85 84 83 82 8i 8o
FIGURE 20-The distribution of P/c!4odo g/!/fz.%o_f"J in the southern Appalachian Moun-
tains. Only my records are included. Localities of hybrid populations between
jorJ#G. and g/"Zz.oJ#J in the Nantahala Mountains of North Carolina and Georgia
are omitted. The absence of records, southeast and northwest of the Appalachian
Mountains is due to lack of field work. The species is common throughout those

Oklahoma, Illinois, Indiana, possibly ohio, northern Kentucky, eastern Ten-

nessee, western North Carolina, northwestern Georgia and adjacent Alabalna.
The ranges of the two species come in contact on the eastern slopes of the
Great Smoky Mountains (King, 1939, Grobman, 1944, mghton, 1962b).
Unfortunately, dorJJ2.J is extremely diicult to find during the summer when
most of our ficld work ;n that area was accomplished,. however, some new
information on the interactions between these species is available, but in only
a small part of their probable zone of contact (fig. 15).
P. dorJJz.J was col-
lected in 193l from l3 miles south of Asheville, North Carolina, in the cen-
ter of a region where many herpetologists have done extensive field work for
many years, but the species has not been found there since. The recent dis'

fovery of dorj.J2.J in the Piedmont o South Carolina (Harrison, 1969) makes

it very likely that I.t does occur at other locaI! I.n the BIue RI.dge and pled-

mont provinces o North and South Carolina, but has probably been overlooked
by herpetologists who do most of their field work in the area during the sum-
A striking interaction between dorJ4/2.f and c2.cecre%J in their zone of con-
tact on the east slope of the Great Smoky Mountains is the absence in this
area of the striped phase of c7orJdr/!.J. This appears to be the only known area
of its range where the striped phase seems to be absent. In fact, the striped
phase is usually much more common than the unstriped phase in this species.
(The unstriped phase of cJorf&Jc.J does possess some red pigmentation on the
dorsum in what is obviously the region of the stripe,. so this phase, unlike
unstriped cz.72crczJf, usually possesses the faint outline of the zig-zag stripe.)
In eastern Tennessee and western North Carolina, unstriped c.%ere%J have
been taken at only one locality (in Cades Cove, Blount County, Tennessee).
The two species are, therefore, very easily distinguished by their color patterns
in the area of overlap in Tennessee - an obvious example of character dis-

Table 4 presents data on the altitudinal distribution of the two species at

the only two places where we have obtained sufficient material at several dif-
erent elevations along a transect. It is clear that at both places the two
species replace each other altitudinally at elevations between 1700 and 2000
feet. Our other data from scattered localities are in agreement with this
conclusion. It is possible that the range of JorJ/3.J extends east into the moun-
tains along some other river valleys besides the known penetration into the
Blue Ridge physiographic province along the French Broad River in the vicin-
ity of Asheville, North Carolina. Since all of the major rivers draining all
but the easternmost areas of the southern Blue Ridge Province flow westward
into the Tennessee River, Jo/f&/z.J might occur at low elevations in some o

TABLE_ 4-Altitudinal distribution of Plethodon and P. cinereus in tcwo traiisects

from the Great Smoky Mountains National Park, in eastern Ten"esSee.

the other river valleys of the area. The a.bsence of c.#crcz/J in all our collec-
tions from the major valleys (French Broad, Little Tennessee, Pigeon) may
be indicative of the presence of dorJarJz.J at these low elevations (see fig.15).

We have not taken cz.%crczJJ above 5400-feet elevation in the southern Appa-
lachians. It has not been found in the spruce-I ir Canadian Zone forests that
occur in the higher mountains of the area (in North Carolina and Tennessee).

INTERACTIONS BETWEEN P. cz.73Crc#J AND P. zoc//crz..-As in the case O

7c.ffrmoced!. and etJe/Jerz., these two species are very distantly related and do not
show any unusual interactions. Part of this may be due to the scarcity of
tc//crz. records, so possible interactions between the two species may not yet
be apparent because of lack of data. The range of zuc//c.rz. appears to be
entirely within the range of cz.72crc%J (see figures l3, 14 and l5).


Four species of eastern large plethodons are presently recognized in east-

ern North America east of the Mississippi River. One species (g/z/Jz.72oJ#J)
ranges throughout most of the area; and four others, with restricted ranges,
are largely confined to the Appalachian highlands (I;g. I). Highton (1962b)
placed these species in three separate species groups.
Adler and Dennis (1962) described a population from Bat Cave, Hendcr-
son County, North Carolina, as a full species, P. /om7.c.r%J. This form di-
ers from yo72fe/oJJee in having longer limbs and in the dorsal color pattern.
Dennis has done considerable field work in the area between Bat Cave and
the nearest J/oz2"oJJec localities in the Swannanoa Mountains and has found
intermediates between the two phenotypes to be common. He now believes
the two forms are conspecific (pers. comm.), and I agree that they should be
so considered. It would appear that this southernmost population of yocacfr-
/oJj`ce, living in a rock crevI.ce habitat, has become slightly dicrcntiated from
typical yorafe/oJJec, but there is still extensive gene flow between the two
populations. P. Adler and Dennis should therefore be included in
the synonymy of yo72feJoJJce.

Highton (1962b) pointed out that there is considerable geographic varia-

lion in #/ztjz.7zoJztJ of the easterli United States. Populations currently assigned
to this species I rom the Piedmont and Blue Ridge provinces of Virginia, West
Virginia, Tennessee, North Carolina and South Carolina, and some immedi-
ately adjacent parts of the Valley and Ridge Provinc.e o Virginia, West Vir-
ginia and Tennessee, have lighter chins and white dorsal spots, with very re-
duced brassy flecking in the white spots compared surrounding popula-
tions. These are referred to here as the white-spotted t~vpe, Populat;ons to

the east of this area in the Coastal Plain of southeastern Virgiiiia, North
Carolina and South Carolina, are characterized by being much smaller, by`
having a darker chin and very small dorsal spots with considerable brassy
pigmentation in them (here called the Coastal Plain type). Populations to
the north, in Maryland, Pennsylvania, New Jersey, New York, and to the
west I rom West Virginia, Virginia and Tennessee to the Mississippi River,
are characterized by large size, a dark chin and large dorsal white spots with
heavy brassy pigmentation (here referrcd to as the brassy-spotted type). Our
extensive field work in the Middle Atlantic states has provided new data on
the distribution and relationships of these three types of slimy salamanders, all
now referred to the species G7/z/2.7zoJztJ.

Figure l6 indicates the distribution of the three phenotypes in Pennsylvania,

Maryland, West Virginf a and Virginia. In this area the three forms appear
to have dierentiated to, or very close to, the species level of divergence. Mem-
bers of this complex appear to be absent from the Coastal Plain physiographic
province o Delaware and Maryland. The brassy-spotted type occurs through-
out Pennsylvania and in Maryland west of the Fall Line, but is replaced
south of the Potomac River by the white-spotted type. Only in the vicinity
o Hancock, Maryland, do populations north of the river show any reduc-
lion in the amount of brassy flecking in the white dorsal spots. Whether this
is due to introgression of genes I rom the white-spotted type on the south side
of the river is not known, but these populations are indicated as ;ntermediate
on the distribution map (fig. 16). The brassy-spotted type occurs south of
the Potomac River in Mineral County, West Virginia, and southwestward
in the western portion of the Valley and Ridge physiographfc province in West
Virginia and Virginia, apparently replacing the white-spotted type along a long
zone of contact, with almost no overlap in the ranges of the two forms. We
have taken the two forms together at one place (Duncan Knob, Bath Coun-
ty, Virginia) with no evidence of intergradation. At a locality just west of
Monterey, Highland County, Virginia, one unusual specimen may represent a
hybrid between the two forms (locality indicated as intermediate in fig.16).

^%t interbreeding
of `.==a.\t^e^rA.?_L=CeL
between_al_=?g thl:S forms.
the two long Zene -of c_optaci
Clearly, the i=v=--=i-`i*.a::
two have reached+Side+nUc'e.
species level of divergence in this area. I am at present engaged in a I urther
stud.`' of the genetic relationships of th.ese two forms, using additional inde-

pendent characters.

The white-spotted type occurs throughout the Coastal Plain o VirgI.nia as

far south as King and Queen, Kin_ Wlmam and northern New Kent coun-
ties. To the south, it is replaced by tI`_c Coastal Plain type, the latter rang-
ing south to South Carolina, Geor\:a and Florida. There is considerable cl!.nal
geographi.c varI.ation I.n the Coastal Pial.n type toward the south, the

spots increasing in size (Highton, 1962b) so that they are proportionatcty as

large as those of the other two types at the southern end of this cline. The
Coastal Pla].n type does not increase in size to the south (Highton, 1962a).

We have collected the Coastal Plain type in the Piedmont of southern Vir-
ginia in southern Dihwiddie County. The Fall Line apparently does not re-
strict the western distribution of the Coastal Plain type in this area any more
than it restricts the eastern range of the white-spotted type into the Coastal
Plain further north. To the west, we have taken a number of individuals
that appear intermediate between the Coastal Plain and white-spotted types
in Chesterfield, Powhatan, Cumberland, Amelia, Nottoway and Brunswick
Counties, Virginia. These localities are indicated by separate symbols in fig-
ure 16. Not all individuals from these localities are intermediate; some appear
to be typical of one type or the other, but there appears to be strong evidence
of introgression between the two very different forms in this part of the Pied-
mont of southern Virginia. The g/zJfc.7zoJz" of this area are also under study,
using additional independent characters for analysis of the genetic interactions
of the two types.

If nothing were known about geographic variation in g/z/j.77oJz" in other

areas, I would conclude that the white-spotted and the brassy-spotted types
were distinct (sibling) species. (If recognized taxonomically, the white-spotted
type would have to be known as P/c/%odo72 zcyczfe4/cc Hairston, 1950, the
only published name that refers to the white-spotted type, but Highton (1970)
showed that the population from the type locality of feJ/4Acz/ce in the Snow-
bird Mountains o North Carolina is a white-spotted type of g/zJzz.7zoJ#J which
shows some influence of introgression from /.orJcr7#.). However, in southwest-
ern North Carolina and southeastern Tennessee there is considerable evidence
of interbreeding of the brassy~ and white-spotted populations of g/2tZj.72of"J
(Highton,1970, and Highton and Henry,1970). Highton and Henry showed
that there is geographic discordance in the variation of plasma protein char-
acters in the two forms, evidence that they are not distinct species in this area.
This relationship is also under current investigation, so I prefer not to recog-
nize the two types as taxonomically different species until their relationships
throughout their entire zone of contact is thoroughly investigated. Such an
investigaticm must also include a study of the zone of contact between the
brassy-spotted and Coastal Plain types in Georgia and South Carolina, where
the two forms probably ccme in contact.
Although there is considerable evidence of interbreeding between the white-
spotted and the Coastal Plain types in southern Virginia at present, I would
regard this as hybridization between good species. Again, however, the rela-
tionship between the two forms must be investigated along their zone of con-
tact further south, (in North Carohna and South Carolina) before any firm

conclusion can be reached as to their true taxonomic relationships. If the

Coastal Plain type should eventually prove to be a good species, separable
from both the white- and brassy-spotted types, the oldest name available is
probably SczJcz77ireJr z/a!rz.o/% Gilliams (1818), with type locality listed as
"southern states," but collected by the "Florida Party." Schmidt (1953) sub-

sequently restricted the type locality of t,tr3.o/ to Charlestcm, South Caro-

lina, which if accepted, would clearly be referable to the Coastal Plain type.
The brassy-spotted type would continue to be recognized as P. 9/ztZz.7zoJ"J
(Green, 1818), since the population at its type locality, Princetcm, New Jer-
sey, is of this phenotype.
For the present, then, the three types will be referred to in this paper as
P. g/%J7.72oJz{J, with the knowledge that they may actually represent a com-
plex of two or three sibling species, pending more complete I uture analysis
of the taxonomic relationships of the three phenotypes that occui` in the Mid-
dle Atlantic states.
Two populations related to P. ouefer/c. from southwestern Virginia were
described as full species, P. dz.#3. Pope and Fowler (1949) and P. /.c4Jocez.
Newman (1954). Highton (1962b) reduced both to the synonymy o P.
evefer/c!.. Neither can be distinguished I rom zuefer/e2. from the rest of its range
by other than minor color dierences that appear to vary clinally. We have
found a population of salamanders closely related to oucfar_/e2. that diers strik-
ingly in color pattern I rom all other ovefar/c. and also has a greater average
number of trunk vertebrae than all other populations of the species. This
form has been taken only at higher elevations on Shenandoah Mountain and
nearb}. North Mountain, in Shenandoah, Rockingham and Augusta Counties,
Virginia, and Hardy and Pendleton Counties, West Virginia, in the Valley
and Ridge physiographic province. We have not found typical euefrr/ec. in the
Vallev and Ridge province west o Shenandoah Mountain in West Virginia;
the nearest records are from the Appalachfan Plateau phys].ographic province,
about 25 miles to the west. Apparently, a population has become isolated from
the main eve far/ez. stock in this area, is restricted to high elevations and thus
cannot exchange genes with the plateau populations to the west. This 25-mile
hiatus has resulted in far more dierentiation than occurs among all other
known populations in the entire range of zuefer/ez. (a north-south distance of
approximately 400 miles) (fig.17). The Shenandoah-North Mountain form
(named below as P/cJfroJo% zJ73cZZ2") resembles ttJe/r/cz. and differs from all
other eastern large plethodons ln possessing a modal number of more than H
trunk vertebrae and in having more webbing on the toes. It strongly resem-
bles g/%fz.7zoJ%J, superficially, because of its very abundant and large-sized dorsal
iridophore spots. The pigmentation of the dorsal spots o #%cJ4/#J appears
identical to the lateral spots of cocA7/cc. and z/cecfZz/j`. The dorsal

spots of /%}z."oJz/J are almost never yellow; they are either white (in the white-
spotted type) or white with numerous associated brassy flecks (in the brassy-
spotted and Coastal Plain types). The lateral spots of many individuals (usu-
ally large adults) of all three types of gr/#Z2.73oJ"f are often yellow, but this
type of spotting has never been observed in the dorsum. Brassy flecks, more
common in central and southern ouGfer/c; than in northern populations, are usu-
ally absent from the dorsum of fJ%cff"J, as are the red spots usually found
on juveniles and often on adults o West Virginia and Virginia gucfer/eG.. The
statistically significant dif ference in the mean number of trunk vertebrae be-
tween #7.cfczfz" and wcAr/ez. are indicated by the data in table 5. An addi-

TA\BLB S-Variation in number of trunk cuertebrae in Plethodon cu)ehrlei and P. uncta-


Number of Trunk Vertebrae

Species 17/17 17/18 18/18 18/19

qJ) ehrl ei 36 9 262

Bun ctatus 35

tional ecological difference between zucfer/cz. and z/72cJf2JJ exists in the nature
of their interaction with the sympatric populations of gJ%}G.%oJ%J. Throughout
the entire range of owc'ferJeG., it is almost always found in the same habitat to-
gether with gJ%Jz.#oJ%J,. but on Shenandoah Mountain we have usually found
only one or the other species. This is of interest in light of the similar appear-
ance o #cecfZ2/J and 7J%Z2.ceoJ%J and the quite distinct color patterns of etJcfrr/cz.
zmd glutinosus.
The question of the taxonomic relationship between etJeferJe. and the new
form is a diicult one to resolve. The new species obviously is very closely
related to t"efer/ez. but is geographically isolated from it so there is no possi-
bility of applying the criterion of intergradation (or lack of it) in determin-
ing their biological relationships. Because of the several differences mentioned
above, I tend to slightly favor the hypothesis that the two forms are as well
differentiated as several other closely related sympatric species in the genus
(`o= exELTpl=, .glut_i?osus zLnd jordani, or hoffina;i zLnd ;iner;UST. --irltf=:ii
their real relationship remains to be determined, I feel it best to regard them,
for the present, as separate species.

Plethodon Punctattts, neNI EsT!eCi+es

DIAGNosls.-An eastern large P/effeodoca of the ecArJcG. group (mghton,
1962b) with a modal number o 19 trunk vertebrae and many white or yel-
lowish white dorsal spots. It diers from all other eastern large plethodons
I.n possessing a higher average number of trunk vertebrae (although there is

overlap in the range of variation of the number of trunk vertebrae with

owcfrr/ef.J the modal number is 18 in evc'ferJc2. and 19 in %necf4!Zz/j:, see table 5).
It also differs from coefrrJec. in color pattern. Most owefrrJez. (except for north-
ern populations from New York and Pennsylvania) have young with red dorsal
spots. In owefrrJe;. there is usually dorsal brassy flecking, although this char-
acter is geographically variable. Neither the red nor the brassy pigments have
been observed on the dorsum o #%cf}%a The dorsal yellowish white spots
o %reffafc" are much larger and more abundant than those of etJefe7Jc2..

fro/ofJJc: USNM 190224, an adult male collected between 0.I and 0.2 mile
north-northwest of the top o Cow Knob, Pendleton County, West Virginia,
on 22 September 1970, by Rudolph T. Danstedt, Douglas F. Fraser and
Richard Highton.
t4J/ofye: USNM 190225, same data as the holotype.

USNM 190226-34, collected 0.2 mile east-southeast of the top
o Cow Knob, Rockingham County, Virginia-Pendleton County, West Vir-
ginia state line, on 29 April l966, by Richard Highton, Robert G. Jaeger
and Richard D. Worthington.
OTHER MATERIAL.-.Living specimens o P%73tc`/f#J have been examined from
all the localities shown in figure 17.

DESCRIpTION 0F HoLOTYpE.~An adult male. Before preservation, the length

I rom the tip of the snout to the anterior angle of the vent was 58 mm, to
the posterior angle of the vent, 61 mm, and the total length 128 mm. The
length of the head (measured to the gular fold) was 12 mm and the width
of the head at its widest point was 9 mm. There are 18 costal grooves on
both sides (equivalent to 19 trunk vertebrae) and the vomerine teeth num-
ber 3 on both sides. In life, the top of the head and dorsum had many scat-
tered, small yellowish-white spots. There are a few brassy flecks only on the
irises of the eye, eyelids, the dorsal surface of the tail and the top of the head.
Yellowish white iridiphore spotting also occurs on the cheeks, sides, legs and
lateral and dorsal parts of the tail. Melanophore gaps on the anterior belly
are occupied by yellowish iridophore pigment, but there is only a trace of this
pigment in the larger melanophore gaps on the chin.
The color pattern of the allotype is similar to that of the holotype. It lacks
the mental gland, present in all adult male eastern large plethodons, but
never in females. It has 17 costal grooves on both sides, and its vomerine
teeth number 6 on the right side and 7 on the left.

DIsTRIBuTIoN.-Higher elevations (over 3000 feet) on Shenandoah Mountain

in Pendleton County, West Virginia, and Augusta and Rocingham counties,
VirginI.a, and on North Mountain (over 2800 feet), Hardy County, West
Vfrginja, and Shenandoah County, Vl.rginI.a. We have not done sufficl.ent field

work in either area to determine its lowest altitudinal limits accurately, but
it is quite probable that the populations on Shenandoah Mountain are now
isolated I rom those on North Mountain.

VARIATION IN P. #7zc/crfz/J.-Variation in the number of trunk vertebrae is

shown in table 5. No evidence of geographic variation has vet been detected
in individuals I rom the two mountain populations.


Although there are fewer interactions between the five large species than
there are between the six small species of eastern plethodons, one of these (be-
tween p/z/fz.ofz" and /.orcZcz7%.) is of considerable evolutionary interest. A sum-
mary of an extensive field study of their ecological and genetic interactions
in the southwestern part of the range o ;orcZ7G?. has recently been published
(Highton,1970, and Highton and Henry,1970). During the past two years,
I have extended this work to include the remainder of the range of /.orJ&72z..
The I ield work is as yet incomplete, but some information on the ecological
interactions of the two spec;es is now available and can be reported here in
addition to a summary of the interactions between the two species already pub-

It has already been indicated that the ranges o %#cJJ%J and owefer/ez. do
not meet. Homan (1967) has not found J/ocefa/oJJcc and ovefrr/ci. together
in southwestern Virginia, but their ranges come so close that some overlap
may occur. This possible zone of contact needs to be identified and studied,
but no new information on it is now available. The ecological interaction of
gJ%f.7.73oJ%J and the two species of the coefer/c2. group (discussed above) needs
further study because of the apparent dif ferences. This leaves the interactions
of the three sympatric species, /.orJ7zz., g/"z.%oJz/J and J/0%Woj.Jcc to be dis-
cussed in detail. The latter species is placed in a different species group (High-
ton, 1962b) from the other two and, as might be expected, the ecological
interactions between the two members of the same species group are much
die=ent from those between either of these and yo#ofJcc.

INTERAcTIoNs BETWEEN JJo%/z/oJJcc AND /.orJ4%!..-Both of these species are

largely confined to higher elevations of the southern Blue Ridge physfographic
province. Although less is known of the detailed distribution o J;072afe/ofJcc
than /.ordz2z., the former appears to be fairly continuous in distribution through.

?ut its range (fig.18), while the range of the latter is subdivided into approx-
lmately 2l geographic isolates (fig. 19). The two are often found together
in the same habitat. In none of our transects have we found an altitudinal
replacement o 7.ord%G. by J7o%fa/a"" Thus far, it would appear as if the
ecological (altitudinal) distribution of the two species fn their area of over-

lap is largely independent of the presence of the other species. Hairston (1949)
reports that J/073WoJJcc is restricted to a zone within 100 feet of streams in
the Black Mountains of North Carolina, but Pope (1950) shows that this is
not the case in many parts of its range.

P. /.07.J#z. and yo73czfeJoJJce differ considerably in size, and thus there may
be decided differences in their ecological requirements; but the difference in
size between .vo7zcrWoJJce and /.07-J&#g. is about the same order of magnitude as
that between g/2/Zz.7zoJa!J and /.orJcr73z.. So the differences in the interaction be-
tween jorc7722. and the other two species are much more likelv due to stronger
competition between the two more closely related species (J.o.rJ4%z. and g/z//.7z-
oJ"J) than to the size differences.

The apparent absence of yo73/!/tJJJce from the contiguous higher sections of

the Blue Ridge province south of Bat Cave has not yet been explained. Since
this species can successfully compete with /.orcJz. and p/z/fg.7zoJz/J throughout
the rest of its range, their presence would hardly be expected to be a barrier
to the spread of J/07zczWoJJcc into this region. It may be significant that the
southernmost known population of Jio7crferoJJcc is the most differentiated one
in color pattern and in habitat preference (see above discussion of /,
p.172). In his study o JJo72frJoJfee, Pope (1950) showed that most of the
known records of yo72cz%JoJJcc are from intermediate elevations between 2700
and 5700 feet. Our field work is in general agreement with that of Pope,
although Hoffman (1967) reported .vo7z4feJoJJec at 2100 feet elevaticm at the
extreme northeastern part of its range fn Virginia, while at the extreme south-
western limit of its range, the Bat Cave population occurs at altitudes as
low as 1430 feet elevation (Adler and Dennis, 1962). It is not clear just
why the species should be able to occupy lower areas at the extreme periphery
of its range and not in the central area where it reaches its greatest abun-
dance. The presence or absence of /.07-J7z/. does not seem to influence the alti-
tudinal distribution of .vo73czfe/oJJce to any measurable extent.

INTERAcTIONs BETwEEN yo#czfe/oJJee AND gJzJZG.ce,oJg/J.-We have collected

J707zc7WoJJce at a total of 50 localities. At 45 (90 percent), g/z/Zz.72oJz/J has also
been taken. At the remaining 5 localities where gJz/J.ceoJz/J was not taken with
J;07zcWoJJcc, 7.orJcz7zz. is present, and these probably mostly represent places
where J.orJczcez. has excluded g/%Jc.72oJ%J. There is no evidence that yo72feJoJJec
ever replaces 7//c.#oJ%f altitudinally. The ranges of the two species overlap
widely in every area where we have found them together. As pointed out
above, there is almost complete overla pin the size range of the two species, and
so it might be expected that they might be in competition for food and other
limiting environmental resources. The largest known yo#frJo" are two fe-
males from Bat Cave (10l mm in body length, measured from the tip of the
snout to the posterior angle of the vent) (Adler and Denni.s, 1962), whl.1e

the largest recorded y/z/jz.oJwJ is a female (measuring 100 mm in body length

in life) we collected on Greenbrier Ridge, Sevier County, Tennessee, in the
Great Smoky Mountains National Park.
We have generally found that g/zJZz.coo..%J is more abundant than .vocaofJec.
At most of the places where we have taken the two together, more g/z/z.7zo"
were collected than yo7zcJfe/oJJcc. A great deal of further work is needed before
an understanding of the ecological interactions of J)07G4fe/oJfce with both 7.orcJo7zz.
and ./"Z!.#oJz/J will be obtained. We have found no evidence of any hybridiza-
tion between J/07z%/oJJcc and the other species.

INTERACTIoNS BETWEEN pJzJ}7.7zoJzJJ AND J.orc7c7#z..-Results of an eight-year

study of the genetic and ecological interactions of these species in the southwest-
ern part of the range of ;ordc#z' (west of the French Broad River) were reported
in Highton (1970). In this area, the range of jorda7zg. is subdivided into about
12 geographic isolates of varying size because it is largely restricted to higher
elevations in the various mountain ranges (Hairston and Pope, ]948 ; Hairston,
1949,1950,1951 ). Both the white-spotted and brassy-spotted g/#2.7GoJz/J occur
in this area, but the range of the brassy-spotted type is largely allopatric to
that of /.orJ47zz.. In some of the /.orJer7zz. isolates, g/z/g.#oJ#J occurs with /.orJcz7".
throughout the isolate, for we found the two together at every locality where
7.orJ%G. was collected, while in other isolates the two species replace each other
altitudinally with varying amounts of elevational overlap. In most areas where
the two species occur together there is no evidence of hybridization between
the two, but in certain isolates hybridization occurs with varying I requencies
ranging from occasional to very frequent. Since the ecological interactions
(measured by the degree of altitudinal overlap) and the genetic interactions
(measured by the amount of hybridization) are highly correlated with the
isolate of 7.ordc7zg., it is concluded that each isolate of /.orc7%z. has evolved its
own, somewhat independent, method of interacting with its very close relative.
In some isolates reproductive isolating mechanisms to protect its genetic integ-
rity have evolved more rapidly, resulting in less present hybridization than in
others,. while at three places, it appears as if earlier populations o /.orddr77J.
were unable to evolve reproductive isolating mechanisms rapidly enough to
prevent "swamping" by g/%f!.7!oJ%J. In some isolates the two species have
evolved mechanisms to permit cohabitation of large areas,. while in other areas,
competition must be so strong that they can coexist only in a very narrow
overlap zone.
During the last two years, a study of the interactions between J.orJ72z. and
gJ%Z2.ceoJaJJ in the remainder of the range o 7.orJar7zz. has been initiated. This
study is not yet complete, but I believe that we now have general outlines
of the isolates o /.ord72G. in the remainder of its range, although many of the are yet to be worked out. We have not found a single I.ndivi.dual

I rom any of the populations east of the French Broad River that can be clas-
sified as a hybrid between J.orda#g. and J#Jz.ceoJ%J. It would appear that the
two species have been in contact in this region long enough to evolve suffi-
cient reproductive isolating mechanisms to effectively prevent most interbreed-
ing between the two species.
The 12 southwestern isolates were discussed in detail in Highton (1970).
Little new information is available (other than a few additional localities for
7.orc7#cez. in some of the isolates), since little field work has been done in that
region since 1968. One new find is the discovery of a single individual of
J.ordc7#c. at Balsam Gap in the middle of the narrow (less than 1 mile) gap
between the Balsam and Blue Ridge isolates. We had previously collected at
this locality twice and had taken a total of 20 7J"f!.rooJ%J, but no 7.ordd!raG.. At
another locahty 0.4 miles north-northeast of Balsam Gap we had collected 17
y/%/z.zzoJ%J and no 7.orJ%,z.. The J.ord#G. at Balsam Gap was taken along with
l2 additional 7Jz/Zz.%oJ#J, providing a total of 32 g/z/Zz.#oJz/J and 1 jordo7z!. from
this locality. Table 6 shows the frequency of /.orcZceG. and #J%ZG.7GoJz/J from sev-
eral localities along the Blue Ridge Parkway in the area. Data I rom other
nearby localities (not along the Blue Ridge Parkway) are in general agree-
ment in respect to the relative I rcquencies of the two species at dierent alti-
tudes (although the elevation of the overlap zone varies considerably, depend-
ing on whether the transect is taken in a stream valley or on a ridge). It
would therefore appear that there is a good possibility that gene I low between
the Blue Ridge and Balsam isolates may still be possible in the vicinitv of
rrAB==_ 6=f lt.i,tudin,al variation in th_e frequency of P\ethodon glutinosus and P. 5ordanl

i.:A_C.0_l!_CtioThn,S!:?in the region b:tTeen i_h.a Brl_sam and flue Ridge i-;ol-;i;s--oi--i.
]or.:an\.i. Mileages are measured from Blue Ridge Parkcway mi;e'-os;;-.(~;;; ;: ;i.r


Locality Elevation Number Number

(mileage) (feet) g lutinosus jor darii
451.2 vi`t =: o`=?S= g=+O or c>o o o o o 0 \O-Cq-cqt- \OC>OhC`1+|`OC^\O
4.4-9. 0
444.2 12
443.0 (Balsam Gap)
0 12
0 14
0 28

Balsam Gap, although I suspect it might well be reduced due to the scarcity
of /.orJd}7ez. at the lowest elevations along the ridge in the vicinity of Balsam
Figure 19 shows all the localities where we have collecte
rdani. We
have been able to co;'irm almost all previous records, so this #g'iqj e essential-
1y represents the known distribution of the species. In the intervening areas
we have found only gJ%!c.72oJz(I (see fig. 20). In this area there are appar-
ently about nine additional isolates of /.ord%z', whose limits are approximately
outlined in figure 21 with suggested names for each.
The Blue Ridge isolate o J.orJ72; extends eastward in the mountains along
the North Carolina-South Carolina state line to the vicinity of Tryon, Polk
County, North Carolina, and then northward to southeastern Buncombe Coun-
ty. North Carolina. East of the French Broad River valley, g/%3.%oJ{/J is
always found with J.orc7cz%. throughout the isolate.
The Black Mountain isolate extends from northeastern Buncombe and north-
western McDowell Counties north through the Black and Green Mountains
to southern Unicoi County, Tennessee. In the Black Mountains, /.orJ#z.
appears to exclude g/z/fz.%oJz/J from its range (Hairston, 1949), but farther
north, y/zJZg.7zoJzJJ is also present (but our collecting has not yet been extensive
in this particular area).

`>- - FLAT TOP
cL`"CHt \




COWEE ` ==:,---
i "\z%
:_-:c---.-`` \ ---.--` -





FIGuRE 21-The names of the geographic isolates o P/efAodoce 7.a/c!4#2. in the south-
ern Appalachian Mountains.

The Bald Mountain isolate of jorcZc7aZ occurs along the Greene County,
Tennessee-Madison County, North Carolina state line. It is separated from
the Black Mountain isolate by perhaps a very narrow area in the vicinity of
Devils Fork Gap (where we have taken only g/c/fz.7zoJz"). At one of eight
collecting stations in the Bald Mountains (4480 feet elevation) we found
only /.orJcz#g., but at all of the other localities g/zJjz.7zoJzff was also taken.

We have not found jorcZaz. in the Nolichucky River Valley, but to the
immediate northeast, this species apparently occurs throughout a very exten-
sive area I rom the Unaka Mountains northeastward to the vicinity of Mount
Rogers, Virginia. This isolate is designated the Roan Mountain isolate after
the highest mountain of the area. In this isolate, g/z/fz.77oJc" and 7.orcJcz7z. occur
together over almost the entire area, except on the highest mountains. We
have not found g/a/2.72oJc# at higher elevations on Roan, Beech, Rich, Stone
and Grandfather Mountains, North Carolina, and Whitetop Mountain and
Mount Rogers, Virginia.
Hoffman and Hubricht (1954) reported /.orJ47,. from Burkemont Mouh-
tain in the South Mountains o Burke County, North Carolina. In spite of
the fact that these are quite low and isolated mountains, surrounded by the
low Piedmont Plateau (the highest point is only 2905 feet in elevation),
J.orJ7zz. is quite common throughout. We have taken it at seven localities in
the South Mountain isolate, but nowhere in the surrounding lowlands. P.
gJz/z.73ofz appears to occur at the highest elevcitions in the South Mountains,
so it probably is distributed throughout this isolate, although we have thus I ar
not taken it at all of the places where we have col_1ected /.orc7cz7zg.. The /.orJcz7zz.
of this isolate are extremely large and have very dark bellies. They appear
to be the population of /.orJdr#z. that resembles yJc/f.73oJz most closely in both
these usually distinctive differentiating characters.
Yet no color pattern inter-
mediates have been taken in the South Mountains; so there is, as yet, no con-
vincing evidence of hybridization in the area. It is surprising that jorJ7zz. has
not been found in another outlying chain of mountains just to the north, the
Brushy Mountains, which are even closer to the Blue Ridge, but are not quite
as high as the South Mountains.
In the Valley and Ridge province of southwestern Virginia there appear
to be five additional J.orcZcz7zz. isolates. Hoffman (1967) mentioned localities
in all of these isolates, but did not collect intensively in the surrounding areas,
so he was not aware of the number of separate isolated populations in that
region. The easternmost isolate, in western Giles County, is designated the
Flat Top isolate. We have so far found J.ord7zz. at only one place on this
mountain and then in company with g/"Zz.73oJzJJ.

The Buckhorn isolate is represented by ;orJz. from three localities at high

elevations on Buckhorn Mountain in Tazewell and Bland Counties, south-

west of Bluefield. At all three of these places g/2/Jc.ceoJztJ was also taken.
The thi.rd J.orda7zz. isolate occurs in the vicinity of Burkes Garden, We found
7.orJ#z. on the south and west rims o Burkes Garden and also took it on
Rich Mountain just southeast of Tazewell. At four o{ the five localities where
We have taken /.ord7c. in the Burkes Garden isolate wc also found gr/z/Zz.%oJ%J.
The one locality where 7/#Z!.oJz/J was not taken is the highest one, 1.2 air
miles north-northeast of the top o Beartown Mountain along the top of Gar-
den Mountain (4460 feet elevation). It is possible that 9/#Z.#oJztJ is absent
I rom the highest parts of this isolate in the vicinity o Beartown Mountain,
but at all the other of our highest localities of the five Virginia Vallev and
Ridge isolates the two species were found together.
The Clinch isolate of /.orc77zz. occupies the highest part of Clinch Mountain
in the area where the boundaries o RussellJ Smyth, Washington and Taze.
well Counties meet. We have not yet col.lected on the highest mountain of
this isolate (also called Beartown Mountain, elevation 4672 feet), so the
upper limit of g/z/f!.ceoJz/J in this area is not yet determined.
The Brumley isolate occupies an area along the Russell-Washington County
line east o Hansonville. We have taken g/2tJz.caoJztJ with 7.ordzzz. at all three
localities on Brumley Mountain where we have taken /.ordcez..
Figure 22 shows the areas of probable elevational overlap between /.07.J7cz.
and gJz/Z.7eoJ%J throughout the range o J.ordce3.. No doubt a number of cor.
reCtions will be made as a result o future field work, especiallv in the area
east of the French Broad River, but the general distributional pattern of the
two species is now probably clear. In the Unicoi, Great Smoky, Tusquitee,
Wayah Bald, Standing Indian, and in parts of the Balsam and Black Moun-
taI.n isolates, the two species generally replace each other altitudinally w;th
a mI.nimum of altitudinal overlap. In the two largest isolates, the Blue Ridge
and Roan Mountain, the two species overlap very extensively, but y/%f.7coJ%J
appears to be absent in several very high areas. In the Gregory Bald, Cheoah,
Cowee, Sandymush, Max Patch, South, Brumley, Buckhorn and Flat Top
MOuntain isolates, g/%Jz.73oJz/a appears to be sympatric with /.ord73. through-
out. The two species overlap extensively throughout most of the remaining
three 7.ord4rceG. isolates (Bald, Clinch and Burkes Garden) except Perhaps at
the very highest points of these isolates. As pointed out in Highton (1970),
the isolate itself , rather than the geographic location or the topography, seems
to be the major factor in determining the ecological interaction between these
two closely related species. It would thus appear that dierent methods O
sharing the available resources have independently evolved in at least some
of the /.a/d73z. isolates, and these dif ferences explain why the two can live
together extensi.vely in some areas but appear to completely exclude one an-
other I.n other regI.ons.


i rI

FlcoR_B ?2-I:t\e ecofogEcal .iTtelacton o Plpil3odon glutii.osus and Plethodon jordani.

Darkened areas are regions where the two species overlap altitudinally.



No obvious interactions between the large and small plethodons in the Appa-
lachians are apparent. It is quite likely that the large size differences between
members of the two groups, as well as the partial dierences in seasonal
activity patterns in at least some of the overlap zones, have reduced the com-
petition between the species of the two groups to the point where they are
minimal, Whether or not this is true for all stages of development is yet
to be determined. It is quite possible that the young of both groups (or the

young large plethodons and the adult small plethodcms) are more strongly
ln competition than are the adults of the two groups. Investigations of these
interactions are much to be desired. It should be emphasized that the nature
of the ecological interactions between species of P/c/feoJo" have just begun to
be studied, and experimental field and laboratory work of the type done by
Jaeger (1969) may in the future yield results that provide an understanding
of the real nature of the interspecific interactions between these species. The
distributional data presented in this paper are almost entirely descriptive, and
real understanding of the interactions described here remains for the I uture.
The pattern that emerges from this analysis is that closely related species seem
to exclude one another, both ecologically and geographically, more I requently
than distantlv related taxa.


I wish to thank all the many colleagues and students who have devoted
an enormous amount of time aiding in the field work and the National Sci-
ence Foundation for the successive grants (G-14026, GB-523, GB-3235, GB-
7299 and GB-19566X) that financiali.y supported this research program. The
General Research Board of the University o Maryland has also supported
the field work. The local Superintendents and Naturalists of the National
Park Service, the Supervisors of the National Forests and various state park
and forest ocials have cooperated by issuing appropriate permits to allow
us to do field work in their respective areas. Many museum curators and
other colleagues have provided information and extended manv other courtesies
to me during the course of these investigations.

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