Front Cover
 Table of Contents
 Specimens examined
 Literature cited
 Figures 1-36
 Back Cover

Title: Status of Desmognathus brimleyorum Stejneger and An Analysis of the Genus Desmognathus (Amphibia: Urodele) (FSM: Bulletin 18(1)
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00000975/00001
 Material Information
Title: Status of Desmognathus brimleyorum Stejneger and An Analysis of the Genus Desmognathus (Amphibia: Urodele) (FSM: Bulletin 18(1)
Series Title: Status of Desmognathus brimleyorum Stejneger and An Analysis of the Genus Desmognathus (Amphibia: Urodele) (FSM: Bulletin 18(1)
Physical Description: Book
Language: English
Creator: Means, D. Bruce
Publisher: Bulletin of the Florida State Museum of Biological Sciences, University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1974
 Record Information
Bibliographic ID: UF00000975
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: ltqf - AAA0350
ltuf - AAP2247
alephbibnum - 000126271
 Related Items
Other version: Alternate version (PALMM)
PALMM Version

Table of Contents
    Front Cover
        Front Cover 1
        Front Cover 2
        Page 1
    Table of Contents
        Page 2
        Page 2
        Page 3
        Page 4
        Page 5
        Page 6
        Page 7
        Page 8
        Page 9
        Page 10
        Page 11
        Page 12
        Page 13
        Page 14
        Page 15
        Page 16
        Page 17
        Page 18
        Page 19
        Page 20
        Page 21
        Page 22
        Page 23
        Page 24
        Page 25
        Page 26
        Page 27
        Page 28
        Page 29
        Page 30
        Page 31
        Page 32
        Page 33
        Page 34
        Page 35
        Page 36
    Specimens examined
        Page 37
        Page 38
        Page 39
        Page 40
        Page 41
    Literature cited
        Page 42
        Page 43
        Page 44
        Page 45
    Figures 1-36
        Page 46
        Page 47
        Page 48
        Page 49
        Page 50
        Page 51
        Page 52
        Page 53
        Page 54
        Page 55
        Page 56
        Page 57
        Page 58
        Page 59
        Page 60
        Page 61
        Page 62
        Page 63
        Page 64
        Page 65
        Page 66
        Page 67
        Page 68
        Page 69
        Page 70
        Page 71
        Page 72
        Page 73
        Page 74
        Page 75
        Page 76
        Page 77
        Page 78
        Page 79
        Page 80
        Page 81
        Page 82
        Page 83
        Page 84
        Page 85
        Page 86
        Page 87
        Page 88
        Page 89
        Page 90
        Page 91
        Page 92
        Page 93
        Page 94
        Page 95
        Page 96
        Page 97
        Page 98
        Page 99
        Page 100
    Back Cover
        Page 101
Full Text

of the
Biological Sciences


Number 1

THE STATUS OF Desmoanathus brimleyorum STEJNEGER



Volume 18


SCIENCES, are published at irregular intervals. Volumes contain about 300 pages and
are not necessarily completed in any one calendar year.

RHODA J. RYBAK, Managing Editor

Consultant for this issue:

Communications concerning purchase or exchange of the publications and all manu-
scripts should be addressed to the Managing Editor of the Bulletin, Florida State
Museum, Museum Road, University of Florida 32611.

Publication date: March 8, 1974

This public document was promulgated at an annual cost of $2403.52
or $2.404 per copy. It makes available to libraries, scholars, and all
interested persons the results of researches in the natural sciences,
emphasizing the Circum-Caribbean region.

Price: $2.45

THE STATUS OF Desmognathus brimleyorum STEJNEGER


SYNOPSIS: This is a systematic study of three externally similar species of plethodontid
salamanders (genus Desmognathus) previously considered to occur sympatrically in
northern Florida (D. auriculatus, D. brimleyorum, and D. fuscus). Larval mor-
phology, color pattern, relative size, and tail morphology were reinvestigated. The
morphology of teeth, jaw profile, premaxillary fontanelle, and prearticular spine were
studied in detail for the first time, and color changes due to ontogenetic melanization
and to metachrosis were also evaluated. All the above characteristics were found to
be taxonomically significant. In addition, the microhabitat selection and ecological
associates of D. auriculatus and D. fuscus were different, indicating ecological isola-
tion of the two species in the area of sympatric contact in northern Florida. Desmog-
nathus brimleyorum Stejneger is a species endemic to the Ouachita Mountains of
Arkansas and Oklahoma. The name D. fuscus carri Neill is a synonym of D. auri-
culatus Holbrook. Populations referred to D. fuscus in this study compared mor-
phologically more closely with D. ochrophaeus from the southern Appalachians than
with populations from near the type locality of D. fuscus Rafinesque.

1 The author is a Ph.D. candidate in the Department of Biological Science at Florida
State University, Tallahassee, Florida 32306, and a Research Associate at Tall Tim-
bers Research Station, Route 1, Box 160, Tallahassee, Florida 32301. This study
was submitted to the Florida State University, Department of Biological Science as
partial fulfillment of the requirements for the degree of Master of Science. Manu-
script accepted 2 April 1973. Ed.

Means, D. Bruce. 1974. The Status of Desmognathus brimleyorum Stejneger and
an analysis of the genus Desmognathus (Amphibia: Urodela) in Florida. Bull.
Florida State Mus., Biol. Sci., Vol. 18, No. 1, pp 1-100.

57 O.<2;
F636 1


INTRODUCTION ------- ...----------- -------------- --- ---------- 2
Taxonomic History of the Genus Desmognathus in Florida ---- -- 6
METHODS .--.......-------------------------------- -.---- 7
ACKNOWLEDGMENTS .......------------- ---------------------------- -- -- -------- 8
RESULTS ....------- ....----.......------- ------ --------- 9
Tooth Morphology .-..-.--- ----....---- --------------- 9
Jaw Profile ....----------..----------- ------ 11
Premaxillary Fontanelle .------------------ --- --------- 12
Prearticular Spine --...---_.-.---.-----------. ..------------------- 13
Tail Morphology ...---.. ..---- ----------------------- 14
Relative Size (SVL) ------------------------------------------ 14
Color Pattern .....---- ------------------------------- ------ 16
Melanization and Metachrosis --------- ---- -------20
Larval Morphology ----------_.-------------------- 23
Microhabitat Selection ------_--------------- --------------- 23
Ecological Associates ----------.------..--..------------------- 25
Summary of Differences between Taxa----- ------------------- 26
DISCUSSION ------------................--------------_--------- .--_.-------------------------------------- 27
Status of Desmognathus brimleyorum Stejneger --- ---------------- 27
Relation of auriculatus to fuscus in Florida---- ------------------- 28
Relation of Florida fuscus to ochrophaeus ------ ----------------35
CONCLUSIONS --_- ----------------..--------- -..-----..... -- 36
SPECIMENS EXAMINED ----- ---....................---- -----. 37
LITERATURE CITED -_. ...... -----------------------------. ------ -------------- .---------___..---- 42
FIGURES ....---- .- ..........-------------..--....... ...- 46

The plethodontid salamander genus Desmognathus Baird, 1849, has a
long and complicated taxonomic history (for partial reviews, see Dunn
1917, Folkerts 1968, Hinderstein 1969). Over 30 species and subspecies
were described between 1820 and 1958. Of these at least six species (D.
aeneus, fuscus, monticola, ochrophaeus, quadramaculatus and wright)
are presently recognized by most authors (Conant 1958, Wake 1966,
Brame 1966, Goin and Cochran 1970).
All of the above species, except D. wright, are polytypic. Of these
Desmognathus fuscus has been split at one time or another into 11 sub-
species (auriculatus, brimleyorum, carolinensis, carri, conanti, fuscus,
imitator, perlapsus, planiceps, ocoee, welteri). Desmognathus ochro-
phaeus, aeneus, monticola, and quadramaculatus have each been consid-
ered ditypic. None of the last three names has ever been synonomized
under D. fuscus, but ochrophaeus has been considered as a subspecies of
fuscus (Allen 1901, Fowler 1906). For purposes of communication some
of the above names will be grouped into three complexes, hereafter re-
ferred to as: (1) auriculatus complex (auriculatus, carri); (2) fuscus
complex (conanti, fuscus, welteri); and (3) ochrophaeus complex (caro-


57 O.<2;
F636 1


INTRODUCTION ------- ...----------- -------------- --- ---------- 2
Taxonomic History of the Genus Desmognathus in Florida ---- -- 6
METHODS .--.......-------------------------------- -.---- 7
ACKNOWLEDGMENTS .......------------- ---------------------------- -- -- -------- 8
RESULTS ....------- ....----.......------- ------ --------- 9
Tooth Morphology .-..-.--- ----....---- --------------- 9
Jaw Profile ....----------..----------- ------ 11
Premaxillary Fontanelle .------------------ --- --------- 12
Prearticular Spine --...---_.-.---.-----------. ..------------------- 13
Tail Morphology ...---.. ..---- ----------------------- 14
Relative Size (SVL) ------------------------------------------ 14
Color Pattern .....---- ------------------------------- ------ 16
Melanization and Metachrosis --------- ---- -------20
Larval Morphology ----------_.-------------------- 23
Microhabitat Selection ------_--------------- --------------- 23
Ecological Associates ----------.------..--..------------------- 25
Summary of Differences between Taxa----- ------------------- 26
DISCUSSION ------------................--------------_--------- .--_.-------------------------------------- 27
Status of Desmognathus brimleyorum Stejneger --- ---------------- 27
Relation of auriculatus to fuscus in Florida---- ------------------- 28
Relation of Florida fuscus to ochrophaeus ------ ----------------35
CONCLUSIONS --_- ----------------..--------- -..-----..... -- 36
SPECIMENS EXAMINED ----- ---....................---- -----. 37
LITERATURE CITED -_. ...... -----------------------------. ------ -------------- .---------___..---- 42
FIGURES ....---- .- ..........-------------..--....... ...- 46

The plethodontid salamander genus Desmognathus Baird, 1849, has a
long and complicated taxonomic history (for partial reviews, see Dunn
1917, Folkerts 1968, Hinderstein 1969). Over 30 species and subspecies
were described between 1820 and 1958. Of these at least six species (D.
aeneus, fuscus, monticola, ochrophaeus, quadramaculatus and wright)
are presently recognized by most authors (Conant 1958, Wake 1966,
Brame 1966, Goin and Cochran 1970).
All of the above species, except D. wright, are polytypic. Of these
Desmognathus fuscus has been split at one time or another into 11 sub-
species (auriculatus, brimleyorum, carolinensis, carri, conanti, fuscus,
imitator, perlapsus, planiceps, ocoee, welteri). Desmognathus ochro-
phaeus, aeneus, monticola, and quadramaculatus have each been consid-
ered ditypic. None of the last three names has ever been synonomized
under D. fuscus, but ochrophaeus has been considered as a subspecies of
fuscus (Allen 1901, Fowler 1906). For purposes of communication some
of the above names will be grouped into three complexes, hereafter re-
ferred to as: (1) auriculatus complex (auriculatus, carri); (2) fuscus
complex (conanti, fuscus, welteri); and (3) ochrophaeus complex (caro-



S linensis, imitator, perlapsus, planiceps, ochrophaeus, ocoee). D. brim-
leyorum will be treated as a full species.
Recently there has been disagreement on the number of subspecies in
D. fuscus in the Coastal Plain of the eastern United States. Some work-
ers have considered auriculatus and brimleyorum to be subspecies of D.
fuscus (cf. Chaney 1958, Folkerts 1968, Harima 1969) but others have ac-
corded them full specific rank (Valentine 1961, 1963, Wake 1966, Coin
and Cochran 1970).
The reasons for the different interpretations probably stem from three
major factors. First is the extensive geographic range of the forms in-
volved. D. f. fuscus (nominate race of the type species of the genus;
type locality, "northern New York") ranges northward to the St. Law-
rence River drainage in New England and southeastern Quebec and into
New Brunswick (Bleakney 1958). The fuscus and ochrophaeus com-
plexes occur southward throughout the Appalachians (where they are
represented by the most named forms: ochrophaeus, fuscus, ocoee, imi-
tator, planiceps), Piedmont (fuscus), and Coastal Plain physiographic
provinces to Florida (fuscus and auriculatus). Thence, members of the
fuscus or auriculatus complexes, or both, range westward to about the
western limit of the Austroriparian Biotic Province in eastern Texas
(Sanders and Smith 1949). The Ouachita Mountains of southeastern
Oklahoma and southwestern Arkansas appear to be the northwestern
S limit of the range of the subfamily Desmognathinae, where it is repre-
sented by brimleyorum. No previous worker has dealt with fuscus or
auriculatus throughout the entire range of either complex.
A second factor is the lack in salamanders of external, taxonomically
utilizable meristic characters. Salamanders are smooth-skinned, usually
lacking in epidermal accoutrements such as excrescenses, tubercles, and
scales. One of few such characters is costal groove number, but this is
not variable infragenerically in Desmognathus.
The third, and most important, factor contributing to the difficulties
encountered in determining systematic relationships in the genus is varia-
tion, both individual and populational. Mayr (1969:147) lists 17 types
of variation that can be found within a single population. Members of
Desmognathus under consideration in this study may be influenced by at
least ten of these.. Each is briefly discussed.
AGE VARIATION.-Members of the fuscus, auriculatus, and ochro-
phaeus complexes pass through a brief larval period of 2-6 months after
hatching from the egg (Wilder 1913, Chaney 1949, Hairston 1949, Eaton
1956, Organ 1961, Tilley 1968). The larval morphology differs from the
post-transformation morphology in several important respects: larvae
possess external gills, undivided monocuspid teeth, distinct palatal mor-



phology, tail fins, "suctorial" mouth, and lack eyelids. Post-larval indi-
viduals lack gills and fins, and possess pedicillate, bicuspid teeth, eyelids,
and reorganized mouthparts. After metamorphosis there is'a 2-3 year
juvenile state in which no dimorphism is apparent in any potential sec-
ondary sex character (Chaney 1949, Hairston 1949, Organ 1961, Tilley
1968). Sexual maturity is probably attained by most members of the
fuscus complex after 2-4 years of age for males and 3-4 for females
(Organ 1961, Tilley 1968). There is a very definite ontogenetic progres-
sion of color pattern from a dorsally spotted larval and juvenile pattern
in all forms to old males and sometimes old females that are entirely
dark. Males have a characteristic change in testicular lobing correlated
with age (Kingsbury 1902, Humphrey 1922, Noble 1927).
SEASONAL VARIATION.-Males and females show seasonal variation in
gonadal activity expressed as morphological change. Males undergo
slight changes in the size of the mental gland and in the lining of the vent
correlated with the advent of the breeding season (Noble 1931, Hays
1966). Gravid females become plump, and large white ova can be seen
through the abdominal wall.
HABITAT VARIATION.-It has been experimentally demonstrated that
individuals of D. auriculatus may undergo color change stimulated by
differently colored substrates (Grobman 1950). In southern Alabama
and northern Florida it has been suggested that auriculatus is an ecophe-
notype of the species Desmognathus fuscus found in mucky soils (Folk-
erts 1968).
DENSITY DEPENDENT VARIATION.-Although no studies are available
demonstrating that interdeme size differences exist in desmognathines,
some studies have shown that population densities, biomass production,
fecundity, and spatial relationships of individuals in the habitat are such
as to indicate that this type of variation is distinctly possible (Hairston
1949, Organ 1961, Spight 1967, Tilley and Tinkle 1968, Barbour et al.
1969). Mean size could vary from deme to deme, depending on age
structure. Males that can be classed to age, utilizing the criterion of tes-
ticular lobing (Kingsbury 1902, Humphrey 1922), may vary considerably
in snout-vent length between demes (pers. obs.). Several taxa in the
ochrophaeus and auriculatus complexes have been partly characterized
by their size differences (i.e., perlapsus, ocoee, planiceps, auriculatus,
ALLOMETRIC VARIATION.-Changes occur during ontogeny from larval
through adult stages in the proportions of the head-to-body and leg-to-
trunk lengths (Martof and Rose 1963, Rubenstein 1971).

Vol. 18 No. 1


color pattern caused by pigment dispersal in chromatophores is a common
phenomenon among amphibians. All the forms under consideration in
this study possess this variation. There may be a short-term change which
takes place in a few minutes to half an hour, or a longer term change in
response to a constant background color of the experimental substrate
(Grobman 1950, this study).
PARASITE INDUCED VARIATION.-This is especially common in D.
brimleyorum in which the digits of the manus and pes may be missing,
fused, or otherwise altered from secondary infection caused by trombi-
culid mites (Acarina, Trombiculidae). Sometimes the infestation is so
heavy that bumps are raised under the skin of the head and body (pers.
obs.). Occasionally other members of the fuscus complex have similar
infestations (Dunn 1926).
TERATOLOGICAL VARIATION.-Desmognathines have well-developed
regenerative ability. Frequently specimens are encountered whose partial
or entire appendage is undergoing regeneration. A leg may appear as a
half-sized miniature of its counterpart. The tail is so often in a state of
regeneration (sometimes for the second or third time), that it is of dubi-
ous value to include its length in any morphometric analysis.
SEX DIFFERENCES.-Aside from primary sex differences, variation is
present in the form of marked secondary sexual differences in all species
of the genus Desmognathus. Juvenile males are difficult to distinguish
from juvenile and adult females without gonadal inspection. Adult males
develop hedonic glands over the body surface (Noble 1931, Hays 1966),
which are clustered into a pad in the mental area of the mandible. Males
grow larger in length and in body bulk than do females. The depressor
mandibulae muscle in males is greatly enlarged, giving a "jowly" appear-
ance behind the angle of the commissure. Males often have a sinusoidal
commissure, whereas in females it is straight. In fuscus and ochrophaeus
both the structure of the dentary and the number of dentary teeth are
different from those in females (Noble 1927, Noble and Pope 1929). The
lining of the cloaca in all sexually mature males is papillaceous, and the
vent appears swollen in the breeding season. These modifications are im-
portant during the deposition of spermatophores. The females have an
invaginated sac in the dorsum of the cloaca called a spermatheca, which
serves as a sperm storage compartment (Kingsbury 1895b, Marynick
CONTINUOUS VARIATION.-This is exemplified by infrademe pattern
variation. Some demes contain individuals having all stages of dorsal pat-
terns from a perfectly maculate condition through a pattern representing
blotches that fuse ontogenetically into a median dorsal stripe. Colors may


also be variable on a more or less continuous basis within a given popula-

Lbnnberg (1894) was the first to list any species of the genus Des-
mognathus from Florida. He assigned a single specimen collected from
a wet hammock in Orange County to D. auriculata Holbrook. Brimley
(1910) listed three specimens from "Hastings, Florida," as D. fusca auri-
culata, apparently following Cope's (1869) referral of this form to a sub-
species of D. fuscus based on color pattern differences. Only D. f. auricu-
latus was reported from Florida (Dunn 1917, 1926, Van Hyning 1933)
until Carr (1940) listed populations from the Torreya State Park ravines
as D. f. fuscus. Although Stejneger and Barbour (1943) followed Carr,
Bishop (1943) apparently was not convinced of the presence of the latter
race in Florida, as he reported only D. f. auriculatus.
The first worker to do a specific study of the genus in Florida was
Grobman (1950). He concluded that, in addition to fuscus and auricula-
tus, a third subspecies (D. f. brimleyorum) was represented in the Flor-
ida panhandle whose range widely interdigitated with the other two.
Shortly thereafter Neill (1951) described a fourth subspecies (D. f. carri)
from spring habitats in central Florida. It was later placed in the syn-
onymy of D. f. auriculatus by Rossman (1959).
Schmidt (1953) followed Grobman, but Carr and Goin (1955) dif-
fered from Grobman slightly in that they considered D. f. brimleyorum
to be restricted to the Florida panhandle west of the Apalachicola River,
with auriculatus ranging eastward from that drainage into peninsular
Florida. They considered D. f. fuscus to range from the north only into
the Apalachicola River basin. Conant (1958), without explanation, listed
D. f. carri from central Florida and moved the eastern limit of brimley-
orum westward to the state of Mississippi.
Valentine (1963) restricted the name brimleyorum to populations in
the Ouachita Mountains of Arkansas and Oklahoma, but gave no reasons.
In his study of the genus Desmognathus in Mississippi, he concluded that
auriculatus was a valid species on the basis of larval morphology and
adult color pattern. This was the first study of desmognathines from the
Gulf Coastal Plain to utilize a new character, which was not based strict-
ly on adult color pattern and size differences. Wake (1966) considered
fuscus and auriculatus to be species in his osteological monograph of the
family Plethodontidae. Folkerts (1968) agreed with Valentine in consid-
ering brimleyorum to be restricted to the Ouachita Mountains. He did
not present data to support this.
Folkerts (1968), on the other hand, differed from Valentine in con-

Vol. 18 No. 1


sidering auriculatus to be a subspecies of fuscus, a conclusion subsequent-
ly followed by Harima (1969). Folkerts (op. cit.) proposed a band of
intergradation between the two subspecies approximately 100 miles wide.
Finally, Goin and Cochran (1970), without comment, listed all three
forms (auriculatus, brimleyorum, fuscus) as distinct species.
Thus, 134 years after Holbrook's (1838) original description of
auriculatus, the taxonomic status of desmognathine salamanders from the
Coastal Plain of the southeastern United States was more confused than
ever. The present study is an attempt to clarify that situation. This has
been done by 1) comparing populations of desmognathine salamanders
from the Ouachita Mountains of Arkansas with those from adjacent
Coastal Plain populations, in order to establish an objective morphologi-
cal basis for evaluating the brimleyorum-fuscus relationship; and 2) in-
vestigating intrademe variation in morphological and ecological charac-
ters of desmognathine salamanders from Florida and adjacent areas for
the purpose of evaluating the auriculatus-fuscus relationship. More con-
servative morphological characters than those used in the past were
sought for use as indicators of systematic relationships of the forms
found within the geographical limits of this study.


In the past many workers studying the genus Desmognathus have failed to ex-
amine variation in certain characters. Although it is well known that individuals of
most species of Desmognathus tend to become uniformly colored (dark brown or
black from melanophore invasion) with increasing age, most workers have relied
heavily on color pattern in their taxonomic analyses. Furthermore, because the
phenomenon of metachrosis is highly developed in these organisms (which also seems
not to be well known), it is very difficult to assess systematic relationships on the
basis of color pattern alone. Since age and sexual variation in the phenotype of an
individual were perceived to be important early in this study, I felt it necessary to
establish criteria by which age-classes could be delimited. This prevented attempts
to compare characters between recently transformed juveniles of a large species with
second-year, sexually mature individuals of a small species, for example.
All transformed males with a single testicular lobe were considered juveniles, even
though some may have been in their first season of sexual maturity. Males with two
or more testicular lobes were considered sexually mature (Kingsbury 1902, Humphrey
1922, Hairston 1949, Organ 1961, Martof and Rose 1963, Rubenstein 1969). Al-
though the number of lobes may not necessarily specify an individual's exact age,
males with the same number are probably closer in age than those with different
According to the state of their ovaries (determined by dissection and by season)
females were judged to be juveniles, sexually mature, or spent following criteria
utilized by Means and Longden (1970) and Tilley and Tinkle (1968).
Over 4,000 Florida specimens of the fuscus and auriculatus complexes used in this
study were collected from September, 1969 to December, 1971. Approximately
10,000 others were examined from museums in the southeastern United States, U. S.
National Museum, and Museum of Comparative Zoology. About 750 specimens of
Desmognathus brimleyorum were collected by me from the Ouachita Mountains.


Specimens were killed in diluted chloral hydrate and preserved in a straight position
in 10 percent commercial formalin. After one week most collections were soaked in
water for at least 24 hours, then transferred to, and stored in 40 percent isopropyl
alcohol. Skeletal preparations were obtained either by maceration in water or the
use of dermestid beetles. One hundred eighty-four desmognathine skulls were
cleaned by hand with jewelers forceps. Teeth were grossly examined with a Wild
M4A dissecting microscope at 50X magnification. Detailed examination was made
with a scanning electron microscope, up to 520X. All line drawings were made using
a camera lucida.
A technique for examining the interior of the mouth cavity was developed. The
atlas-mandibular ligament was severed near its insertion on the mandible by a scalpel
incision along a line beginning at the posterior corer of the eye and continuing to
just behind the angle of the gape. This eliminated damage to the crowns of the
teeth and destruction of the quadrate and squamosal bones, allowing the mandible to
be lowered to its full extent with ease.
Throughout the paper snout to vent measurements are indicated as "SVL."
STUDY AREA.-Specimens were collected personally from about 250 localities
throughout the southern range of the genus Desmognathus (Fig. 1 [not all plotted]).
The study region includes the southeastern United States from eastern Texas to
the Atlantic Ocean (Fig. 1). Within this area emphasis is placed on northern and
western Florida (Fig. 1B) and the Ouachita mountain system in Arkansas and
Oklahoma (Fig. 1A). Specimens have been examined from all areas peripheral to
the study region, with material examined from a total of about 700 localities through-
out the range of the subfamily. Only those specimens examined from the study
region and a few other localities are listed in the "Specimens Examined" section.


The following individuals, who have loaned study specimens under their care,
are gratefully acknowledged: Walter Auffenberg, Florida State Museum (UF/FSM);
Herbert T. Boschung, University of Alabama (UA); John Carpenter, University of
Oklahoma (OU); William Dopson and Joseph Hamilton, University of Georgia (UG);
Neil H. Douglas, Northeast Louisiana State College (NELSC); Harold Dundee, Tu-
lane University (TU); James N. Lane, McNeese State College (McNSC); Robert
Mount, Auburn University (AU); James S. Peters, United States National Museum
(USNM); Henry M. Stevenson, Florida State University (FSU); Richard Worthing-
ton, personal collection (RW); Sam Telford, personal collection (SRT); and E. E.
Williams, Museum of Comparative Zoology (MCZ).
To the members of my graduate supervisory committee, R. K. Godfrey, D. S.
Simberloff, and H. M. Stevenson, a special note of thanks is extended for their guid-
ance and especially for their infinite patience. Others who helped in many ways
were: W. Wilson Baker, Jeffrey Black, Steve P. Christman, Robert Crawford, Milton
Hopkins, P. E. Jinright, Byron C. Marshall, Richard McLean, Henry Nash, Ronald
Parker, William F. Vockell, and Ralph W. Yerger. I owe much to Storrs L. Olson
and Camm C. Swift, who provided me by personal example, models from which
most of my motivation was derived. Also, the able assistance in the field by Clive
J. Longden and James F. Berry is gratefully acknowledged. I thank Storrs L. Olson
for reading part of the manuscript.
Most of all, I thank my wife, Helen, who accompanied me in the field, silently
endured a houseful of specimens, typed each draft of the manuscript, and supported
my research in every way.
This study was supported mostly by Tall Timbers Research Station through a
Gerald Beadel Scholarship Grant. To Edwin and Roy Komarek, of that institution,
I extend my deepest appreciation for making it possible for me to carry out my re-
search activities to the fullest extent. The Department of Biological Science, Florida


Vol. 18 No. 1


S State University, provided transportation for three months and some of the materials.
Scanning electron microscopy was supported by the Sensory Biology Research Center
of Florida State University under USPHS Grant No. NS 07468-04. This paper was.
submitted in partial fulfillment of the requirements for the degree of Master of Sci-
ence, Florida State University, Tallahassee.
The Seal Salamander (Desmognathus monticola Dunn) was first dis-
covered in Florida during the course of this study. The range of this
species in the state is probably confined to northern Escambia County,
where small populations inhabit mesic, spring-fed ravines along Canoe
Creek, a tributary of Escambia-Conecuh River. Results of the discovery
were published elsewhere (Means and Longden 1970) and will not be
discussed further here.
The following characters were found to be of systematic value in this
study: 1) tooth morphology; 2) jaw profiles of sexually mature males; 3)
premaxillary fontanelle morphology; 4) prearticular spine morphology;
5) relative size (snout-vent length, SVL); 6) tail morphology; 7) color
pattern; 8) degree of melanization; and 9) larval morphology. In addi-
tion, two ecological aspects of the forms studied, microhabitat selection
and ecological associates, were found to be significant in assessing evolu-
tionary relationships. Each will be discussed in order of occurrence.

The gross morphology and structure of urodele teeth have been
studied by many workers (Hilton 1951, Kerr 1960, Parsons and Williams
1962, Parker and Dunn 1964, Means 1971, 1972). Adults of Recent
Amphibia are unique among vertebrates in possessing transversely di-
vided jaw teeth. These teeth consist of an enamel-capped orthodentine
crown attached to an orthodentine pedicel by a fibrous connection. The
teeth are pleurodont, the pedicel being attached to the underlying bone
along its base and labial side. In adult urodeles the crown is usually bi-
cuspid, and the axis of each cusp is parallel to the long axis of the bone
of attachment. The lingual cusp is usually apical and larger in mass
than the labial cusp. Palatal and occasionally coronoid teeth are present
in modern amphibians but will not be discussed in this study.
Analyses of tooth morphology have been attempted on a limited
basis within the subfamily Desmognathinae in studies of sexual dimorph-
ism (Noble 1927, Noble and Pope 1929). In his comparative osteology
of the family Plethodontidae, Wake (1966) commented only on the mor-
phology of pedicels in Phaeognathus hubrichti. Otherwise, he did not
compare the morphology of teeth nor of maxillary and dentary bones


In this study sexually mature males of the species of Desmognathus,
Leurognathus, and Phaeognathus were examined to determine interspe-
cific differences in detailed crown morphology of dentary and maxillary
teeth. Figure 2 illustrates the similarities between species of the three
genera. Basically the amphibian bicuspid crown is retained, but a trend
toward reduction and loss of the labial cusp is evident in the series A,
B, C, G, D, E, F, respectively. Desmognathus quadramaculatus, D.
aeneus, and D. wright have elongate crowns tapering to a more acute
apex than the other species studied. I have investigated variation in
crown morphology in this study only for desmognathines of the Gulf
Coastal Plain and Ouachita Mountain provinces.
Figure 3 illustrates the sharp distinction between the crown mor-
phology of D. brimleyorum (A-D) and all other forms in the subfamily
Desmognathinae. Of note particularly is the contrast between Ouachita
Mountain populations and the surrounding Coastal Plain forms (E-H).
The rounded, fungiform crowns of the Ouachita specimens were found
to be present on both maxilla and dentary throughout post-metamorphic
development to adulthood. There was no significant infraspecific varia-
tion noted in crown morphology in populations throughout the Ouachita
Mountain uplift.
Figure 4 compares the Ouachita crown morphology with the Coastal
Plain type more closely. The Coastal Plain type (A) is a piercing tooth
after the style of other desmognathines, which have a reduced labial cusp
and a pointed lingual cusp whose apex terminates above the top of the
former. In Ouachita Mountain specimens both cusps are reduced to thin
enamel ridges of equal height lying on top of the dome-shaped crown.
A conspicuous flat region lies between the two cusps, and the whole as-
pect of the tooth gives the impression that it serves a crushing function.
Whether these are functionally molariform, however, has not been investi-
Representatives of all described taxa of the subfamily Desmogna-
thinae were examined in order to assess the significance of the distinctive
crown morphology of the Ouachita Mountain populations (Means 1971).
Desmognathines can roughly be placed into three groups according to
the relation of the labial cusp to the lingual cusp (Table 1).
The first type is the least common in the subfamily; these teeth are
decidedly fungiform, with both cusps about equal. Species of Type II
(except Phaeognathus hubrichti) have the maxillary teeth crown shape
similar to Type I (usually the labial cusp is slightly reduced), but Type
II dentary teeth are uniquely different (the lingual cusp is strongly re-
curved posteriorly). This second type represents dimorphism within an
individual, but it is more strongly expressed in males than in females

_ _~_



Type I Type II Type III

Cusps Nearly Equal; Lingual Cusp Recurved Labial Cusp Lost or
Crown Fungiform, Posteriorly; Labial Greatly Reduced,
Non-Dimorphic Cusp Present, Non-Dimorphic

D. brimleyorum D. ochrophaeus D. aeneus
D. fuscus (NE U.S.) D. "fuscus" (SE U.S.) D. monticola
D. auriculatus D. quadramaculatus
D. ocoee D. wright
Phaeognathus hubrichti Leurognathus marmoratus

(Noble 1927, Noble and Pope 1929). This type of tooth morphology
commonly occurs with commissure sinuousity (an external character
sometimes used by others [Dunn 1917, 1926, Bishop 1943]). Type I
teeth are not variable in morphology between jaws and have both cusps
relatively well developed.
Type III teeth diverge from the typical amphibian bicuspid tooth
morphology by reduction of the labial cusp. The loss of the labial cusp
in some individuals of D. wright and D. aeneus is here judged to be a
secondary one, as reported in the frog genus Ceratophrys (Lehman 1968,
Schultze 1970).
The crowns of teeth in Ouachita Mountain populations are remark-
ably uniform morphologically in three ways: (1) maxillary and dentary
teeth are similar, (2) infrademe variation is small, and (3) interdeme
variation is small. Considering how little variation there is in the crown
morphology of brimleyorum, and how distinctive it is when compared
with that of most other members of the subfamily, I conclude that this
tooth type must have a genetic basis and is not environmentally induced.


Another osteological character, or complex of characters (apparently
best developed in the ochrophaeus complex of the southern Appala-
chians), is the jaw profile of sexually mature males. Cope (1869) was
the first to notice that males of D. ochrophaeus had the ". . posterior half
mandible concave and edentulous." Boulenger (1882) also mentioned the
concave and edentulous condition of the mandible of males in his descrip-
tion of D. ochrophaeus. Me made no comment concerning the mandible
of D. fuscus, therefore implying that it is not concave and edentulous.
Dunn (1916, 1917, 1926) seems to have ignored this character altogether,


but Noble (1927) and Noble and Pope (1929) carried out castration and
testicular transplant experiments which demonstrated that, once formed
in the adult, the concave and edentulous structure of the mandible in
males does not undergo any change induced by hormone treatment.
Figure 5 illustrates jaw profiles in 12 species of the subfamily Desmogna-
thinae. The line of the commissure is relatively straight between the
maxilla and dentary in A-E and G-J. In marked contrast to this is the
bowed condition of the maxilla and the notched (concave) dentary in
southern fuscus (K), ochrophaeus (L), and ocoee (F). It is significant
that no population of brimleyorum (I) or auriculatus (J) examined
showed any evidence whatever of this type of sexual dimorphism.
In Florida the marked sexual dimorphism in mandibular and maxil-
lary morphology of D. fuscus is geographically variable. This type of
dimorphism is constant and always present in populations in the Chatta-
hoochee-Flint-Apalachicola River drainage and the lower Ochlockonee-
Little River drainage of Florida (Fig. 6). Furthermore, specimens ex-
amined from ravine demes all the way to the headwaters of the Flint
and the Chattahoochee rivers in the southern Appalachians are nearly
identical in this condition. However, demes identified as fuscus from
Gulf Coast drainages (Escambia to Choctawhatchee rivers) on the basis *
of other characters may possess jaw dimorphism somewhere along a gradi-
ent of expression from none to fully developed (Fig. 6).
In a number of Florida localities both D. auriculatus and D. fuscus
can be collected within a few hundred meters of each other, and in two
localities where I have taken specimens of both under the same debris,
no intermediates have been found. Figure 7 illustrates the difference be-
tween both species in jaw profile dimorphism, viewed externally.
Within the geographical limits of this study, the significant facts
involving this character complex are: (1) D. brimleyorum has no sexual
dimorphism in jaw profile, whereas peripheral demes (northern Louisi-
ana parishes and Crowley's Ridge physiographic region, Arkansas), here
referred to D. fuscus, indicate slight to moderate dimorphism; (2) auri-
culatus has straight jaws in both sexes, whereas adjacent demes referred
to fuscus possess strong profile dimorphism; (3) in Florida strong profile
dimorphism is always present in fuscus demes throughout the Chatta-
hoochee-Flint-Apalachicola River drainage, but in more westerly Gulf
Coastal drainages (Escambia to Choctawhatchee rivers) demes referred
to fuscus are variable in this character.

Desmognathines have an axially oriented fontanelle between the two
nasal processes of the premaxilla. The fontanelle is closed by fusion of the

_ __


S two rami in Phaeognathus hubrichti, a terrestrial burrower, and in Leu-
rognathus marmoratus, a subaquatic burrower. Presumably the loss of
the fontanelle serves to add structural strength to the snout as a further
adaptation of a burrowing morphology already well developed in other
skull characters of the subfamily. This character has not previously been
used in studies of interspecific variation.
In all species examined in the genus Desmognathus, the fontanelle
is present and relatively similar among most forms (Fig. 8). It usually
occurs between the two premaxillary processes for about two-thirds of
their length, beginning at the mandibular portion and progressing well
beyond the nasal constriction, terminating about the level of the anterior
edge of the orbit. It is well developed in all skull preparations examined
of specimens from Florida ravines that meet other criteria for the fuscus
phenotype (Figs. 8, 9). However, swamp-muck forms of the auriculatus
phenotype have nearly lost the fontanelle, it being confined to a tiny
aperture between the nasal processes of the premaxilla just posterior to
the pars dentalis (Figs. 8, 10). This reduction of the fontanelle is not
subject to sexual dimorphism, and it is a constant character in all size
classes of these demes and in every population examined for this study
(Fig. 10).


In desmognathines the head is used as a wedge for burrowing in the
loose substrate. As a result, desmognathines have evolved a complex sys-
tem of muscles, bones, and ligaments specifically adapted to this behavior.
A large ligament (the atlas-mandibular ligament) has evolved in the
elevator mandibulae muscle which extends from a dorsal ridge on the atlas
to the mandible (Baird 1849, Wake 1966, Hinderstein 1971). It serves
to bring the skull down when the quadrato-pectoralis is contracted.
Hinderstein (1971) apparently followed Wake (1966) in assuming that
this ligament had its origin on the coronoid process (= spine) of the
prearticular bone of the mandible. Both are incorrect in this assumption,
however. The atlas-mandibular ligament actually is inserted on the den-
tary bone (Fig. 11C), with the dorsal spine lying along the lateral sur-
face of the ligament just posterior to the insertion.
The prearticular spine serves as an additional surface for insertion
of the elevator mandibulae muscles. The main surface for muscular inser-
tion on the mandible is the prearticular shelf. The muscles that insert
here close the mandible. The prearticular spine may have some fascia
attachment to the lateral side of the atlas-mandibular ligament, but ten-
sion on this ligament is transferred primarily to the surface of its insertion


on the posterior, lingual portion of the dentary immediately behind the
This spine is developed to a greater or lesser degree throughout the
subfamily. However, it is most strongly developed in Desmognathus
brimleyorum, and occurs as a high pinnacle pointing dorsally to the plane
of the prearticular shelf (Figs. 11, 12). Thus it may serve as a qualita-
tive character in partly delimiting D. brimleyorum from surrounding
coastal plain populations of desmognathines.

Much has been said about the tail of desmognathines as a character
serving to distinguish one form from another. Most authors (Cope 1889,
Dunn 1917, Bishop 1943) have considered the cross section at the base
of the tail to be important. Gross morphology of the whole tail and its
length are often unreliable as characters, because salamander tails are
commonly broken and undergoing regeneration. Usually a partially re-
generated tail is recognizable externally, but completely regenerated tails
often are not detectable.
In this study, tails examined using radiographs were found to be
significant as indicators of taxonomic relationships in Florida (Figs. 13,
14). The fleshy portion of the tails of swamp and river floodplain muck
inhabitants (auriculatus) is greater than those in ravine (fuscus) demes.
A small, but distinct, dorsal keel is present and usually evident for the
full length of the tail. The tails of swamp and floodplain, muck-inhabit-
ing auriculatus are decidedly compressed and blade-shaped when com-
pared to the slender, rounded aspect of ravine-inhabiting fuscus. Distal-
ly, on complete or fully regenerated tails, the blade aspect persists to the
tip in swamp and floodplain desmognathines. The tail of ravine speci-
mens tapers into a long, terete filament. The basal cross-section in
Florida fuscus is rounded and for its entire length compares exactly with
that of ochrophaeus. It is significant that the regenerating tails of ravine
salamanders and those inhabiting floodplains regain the particular mor-
phology of the original tail peculiar to the respective ecophenotype.

Although size differences have been used by most authors in diag-
noses of desmognathine taxa, no study has adequately compared the
three species investigated herein. Grobman (1950) listed all three from
Florida but did not comment on size differences.
Dunn (1917) presented data showing that D. brimleyorum (from
the Ouachita Mountains) was larger than fuscus (northeastern U. S.),

_ __

Vol. 18 No. 1


S but he gave no figures for auriculatus. In 1926 Dunn reported only the
largest known sizes for these three taxa. He indicated that brimleyorum
and fuscus reached approximately the same maximum length (65-68 mm
SVL), but implied that auriculatus was smaller (49 mm SVL). Bishop
(1941) presented more definite data on the size of adult D. fuscus from
New York: males, x=48.6 mm SVL, N=90; females, R=43.6 mm SVL,
The average of maxima for adult D. auriculatus from seven localities
in northern Florida and adjacent Georgia given by Rossman (1959,
Table 1) was 57.1 mm SVL. Neill (1951) and Valentine (1963) reported
that adult males of auriculatus ranged between 45 and 60 mm SVL.
Valentine (op. cit.) reported that adult males of fuscus in Mississippi
ranged from 30-58 mm SVL, thus showing that auriculatus was larger
than fuscus in that state.
Folkerts (1968) believed D. fuscus (35-65 mm SVL) was larger
than auriculatus (33-61 mm SVL) in Alabama and western Florida. In
this study auriculatus was found to barely enter southern Alabama along
floodplains of the major rivers. My examination of specimens of auricu-
latus seen by Folkerts in Auburn University collections indicates that
most were juveniles or small adults. Thus it became evident early in this
study that a method for determining relative age of specimens was im-
portant to avoid confusion resulting from mixture of measurements taken
S from several age classes. Whenever possible specimens were dissected,
and the gonads were examined when taking size measurements.
Figure 15 illustrates the size relationships between D. auriculatus,
brimleyorum, and Florida fuscus. Although these samples are not large,
I believe they are fairly representative and reflect the relative differences
between the same age classes of the three biotypes.
The sample of 126 D. brimleyorum from the top of Rich Mountain,
Arkansas (Fig. 15), is larger in all age classes than previously reported
for this form (Chaney 1958). I believe this is because the Rich Mountain
locality is at a very high altitude (ca. 2600 ft.) in the Ouachita Moun-
tains, and Chaney's sample (114 postlarval specimens) is near the lowest
altitude known for brimleyorum (ca. 400 ft.). Many ecothermic verte-
brates show an increase in size correlated with an increase in altitude
over the geographic range of the species.
It is readily seen that the mature male and female classes of D.
auriculatus (from the Ochlockonee River drainage) are larger than the
same classes of fuscus from the Torreya ravines in Liberty Co., Florida
(Fig. 15). Furthermore, at one locality where both species occur to-
gether (Sweetwater Creek, Liberty Co., Florida) in the western Florida
study area, the same relative size differences obtained (Fig. 16).


As an example of the problems that can arise from not comparing
similar age classes (determined by direct gonadal inspection), Figure 17
compares paratypes of Desmognathus fuscus carri Neill with topotypes
collected from the same Cabbage Palm hammock stream at Silver Glen
Springs by Mr. Steve Christman in 1970. Included for comparison is the
size-frequency distribution for D. auriculatus used in Figure 15. It is
apparent that most of Neill's original series was composed of juveniles.
Since relative size was diagnostic for D. f. carri, this character is invalid
(Rossman 1959). More intensive collecting has turned up adults, which
were missing from Neill's type series.
In this study D. auriculatus averaged larger than western Florida and
southern Alabama fuscus in all age classes, including larvae. This is in
agreement with Valentine (1963) and Rossman (1959), but not in accord
with Folkerts (1968).

Color Pattern
Color pattern has figured heavily in earlier attempts to systematize
salamanders of the genus Desmognathus. Largely because of the high
degree of variability of this character complex, along with a great deal
of overlap in gross appearance between variants of different forms, most
species have several synonyms. Difficulty in identification of species is
illustrated in a paper by Huheey (1966). Almost every study dealing
with desmognathines from the Great Smoky Mountains National Park has
listed D. fuscus as occurring there. Huheey reports that for every case
he reinvestigated, specimens in question turned out to be either D. monti-
cola, quadramaculatus, or ochrophaeus, and that in his opinion D. fuscus
is not in the park. Conversely, some species are not widely acknowl-
edged because of their gross similarity to a congener.
Cope (1896) assigned considerable taxonomic value to "mucous
pores" in the development of the lateral pattern of the adult. He recog-
nized two lateral series of pores, a superior and an inferior one. The
three species he listed in 1889 were partly distinguished by the presence
or absence of one or both series.
In his review of the genera Desmognathus and Leurognathus, Dunn
(1917) stated, "I have been unable to use the lateral pores as diagnostic
characters. This is largely because the distinctness of these pores is so
dependent on the preservation." However, nine years later in his mono-
graph of the family Plethodontidae, Dunn (1926) placed great importance
on ". . three rows of unpigmented areas on each side of the larva." His
source was Banta and McAtee (1909), who showed the development of
the color pattern of Eurycea bislineata to be dependent on these unpig-

_ __

Vol. 18 No. 1


S mented areas. Apparently neither Dunn nor Banta and McAtee realized
what the lateral unpigmented areas really were.
Kingsbury (1895a) was the first to recognize that the lateral rows of
unpigmented areas were actually lateral lines of the vertebrate acousti-
colateralis system. The "unpigmented areas" of Dunn and Banta and
McAtee surround free neuromast organs. These organs in Amphibia are
usually at the bottom of a shallow pit or groove (Dijkgraaf 1962). I
have examined every form in the subfamily Desmognathinae and find
that the only "pores" along the side of the larvae and transformlings cor-
respond to neuromast pits or their vestiges (Kingsbury 1895a, Hilton
1947). Thus, Cope's lateral rows of mucous pores correspond to the de-
pressions of free neuromast organs in desmognathines.
Three rows of lateral-line organs are found in the larvae of sala-
manders (Kingsbury 1895, Hilton 1947, Eaton 1956). The development
of adult pigment patterns in desmognathines is strongly influenced by the
presence of the lateral lines. Eaton (1956) very accurately described the
pigmentation of the larvae of Desmognathus quadramaculatus and
ochrophaeus (D. fuscus according to Eaton; I tentatively follow Martof
and Rose [1963] and Huheey [1966] in considering the form from Mt.
Mitchell, N. C., as ochrophaeus).
In this study, all species examined during the larval stage possessed
three lateral-line series that were usually obvious under magnification as
rounded, light spots. I follow Eaton's (1956) terminology in recognizing
dorsolateral (Row 1), lateral (Row 2), and ventrolateral (Row 3) lines
(Figs. 18a, 19a, 20a, and 21a). Row 1 begins immediately above or
just anterior to the insertion of the forelimbs on the trunk. It proceeds
down the entire length of the trunk and out onto the dorsal sides of the
tail for about two-thirds its length. Neuromast sites are located on ap-
proximately every second myomere. Row 2 begins in about the same
place as Row 1 and proceeds along the trunk just below the level of Row
1. Often both lines appear to form a single line because neuromast sites
of Row 2 usually alternate in position with those of Row 1. There occurs
about one neuromast site per myomere in Row 2. This row diverges
from Row 1, beginning above the insertion of the hind limbs, to form a
well-marked system of neuromast sites mid-laterally on the tail.
Row 3 is the shortest, occurring only from the axilla to the groin,
about one and one-half neuromast sites per two myomeres. In the above,
it is seen that the number of neuromast sites in any given line does not
exactly correspond to the number of myomeres (14 on the trunk, 20+- on
the tail). This is probably because the lateral line neuromast sites are
formed by nervous tissue that grows out from the acoustic tubercle in the
dorsolateral wall of the medulla oblongata (Dijkgraaf 1962), invading



the trunk region during embryogenesis. These sites thus form inde-
pendently of the metameric development of the trunk and the tail. Table
2 outlines the post-metamorphic fate of each lateral line (as each is ap-
parent on transformed specimens in the form of a line of lightly pig-
mented spots, portholes, dots, etc.) for the three desmognathines of this

brimleyorum, AND FLORIDA D. fuscus.

Taxon Row #1 Row #2 Row #3

forms small, dis- becomes obscured forms diagnostic
Desmognathus create, dorsal trunk by melanophore in- rows of white spots
auriculatus spots that do not vasion on trunk; between axilla and
coalesce; same for forms discrete port- groin throughout
tail. holes on sides of life; not on tail.
tail throughout life.

forms loose dorsal becomes obscured becomes obscured
trunk pattern dur- by melanophore in- by melanophore in-
ing juvenile stage, vasion on trunk; vasion on trunk
D. brimleyorum which dissipates to forms rounded port- following metamor-
uniform coloration holes on tail as ju- phosis; not on tail.
in adults of both veniles become in-
sexes; same for dor- vaded postmaturity
sal tail pattern, by melanophores.

forms strong dorsal barely discernible, becomes obscured
trunk pattern (be- even in larvae on by melanophore in-
Florida D. fuscus comes obscured trunk; forms round- vasion on trunk
only in old males); ed spots on tail, during early post-
forms strong dorsal which become ob- metamorphosis; not
tail stripe with scal- secured following on tail.
loped borders, transformation.

The color pattern of transformed Desmognathus brimleyorum devel-
ops from a typically desmognathine blotched condition present in most
juveniles to uniformly brown dorsally (and laterally) in mature adults
(Fig. 22). Some demes of brimleyorum have a high incidence of dorsal
blotching, especially at lower elevation localities. All Ouachita Mountain
collections examined contained a preponderance of patterned juveniles
and none or only a few uniformly-colored, mature adults. These collec-
tions could easily be identified as fuscus without determination of age
and close inspection of other characters (i.e., tooth and prearticular spine
morphology). The venter in younger juveniles of brimleyorum appears
immaculately white, but close inspection reveals a relatively uniform

Vol. 18 No. 1


suffusion of dark pigment cells, especially on larger juveniles. With in-
creasing age the intensity of pigment becomes greater, giving a faintly
soiled appearance. The venter is sometimes weakly mottled, but basi-
cally appears light tan to white when compared to the dark, uniformly
colored dorsum.
Laterally, D. brimleyorum is distinguished by a nearly bicolored ap-
pearance. The light ventral color meets the dark dorsal color about mid-
way along the side (between Row 3 of lateral-line light spots and a line
above the division between the underlying epaxial and hypaxial muscu-
lature). The meeting of dorsal and ventral colors is relatively well-
defined but not sharp. Considerable interdeme variation exists in the
lateral pattern, however, due to the effects of melanization.
The pattern of Desmognathus auriculatus is best illustrated by speci-
mens from Mississippi (Fig. 23). These are not so melanistic as indi-
viduals from Florida (Fig. 24) and southeastern Georgia (Fig. 25) demes,
and a pattern is evident. Note particularly that the neuromast organs
of lateral-line Row 1 do not repel melanophores nearly so strongly as
those in fuscus (Figs. 21 & 27). Because of this, fusion of pigmentless
areas surrounding the neuromast vestiges (so important in dorsal pattern
S development of fuscus) never occurs in auriculatus. Characteristically,
auriculatus has a parallel pair of thin, light lines on the neck region that
extend from the level of the insertion of the dorsalis trunci muscles on
the underlying skull, posteriorly to the insertion of the forelimbs (Figs.
23 & 24). Lateral-line Row 1 forms discrete, punctate spots on the dor-
solateral sides of the tail on either side of the fleshy dorsal fin (Figs. 23
& 24). The dorsal area between the rows of tail spots is usually less
intensely pigmented than any other part of many specimens and it often
appears deeply reddish (as do the trunk and tail "portholes" occasion-
ally in some individuals in demes generally lacking red color on other
parts of the body).
The ventral pattern of auriculatus is usually comprised of flecks of
iridiophores over a dark (usually black) ground color. Some demes may
lack the flecking entirely and others may have a reticulum of lighter and
darker areas (Figs. 23 & 24). The opposite extreme from the light pat-
tern of specimens pictured in Figure 23 is total absence of any pattern
due to intense black pigmentation in some demes in Florida and south-
eastern Georgia (Figs. 25 & 26). The typical color scheme of auriculatus,
when found in ravines in western Florida, is coal black with a wash of
intense reddish color over the dorsum (Fig. 26; red lost in preservation).
The red is usually obscured by black pigment and is only readily evident
where it occurs on the light neuromast sites, dorsal base of the tail, or on
the cheek. This color occasionally is so intense, however, that some in-


dividuals are quite red in gross appearance. The reddish color never
occurs on the venter. In my opinion, the spring margin ecophenotype of
central Florida described by Neill (1951) as Desmognathus fuscus carri
is a local variation similar to the ravine ecophenotype of auriculatus found
elsewhere during the course of this study.
The color pattern of transformed D. fuscus in the study area is also
highly variable. Basically, juveniles and females are strongly marked
dorsally by large light areas surrounding neuromast vestiges of lateral-
line Row 1 (Fig. 27). These light areas exist in all states, from discretely
isolated by fringing melanophores to broadly fused along the midline of
the dorsum as a smooth, light stripe. The light areas were often differ-
ently colored in different individuals with red, yellow, or lighter shades
of brown. Characteristically in fuscus the dorsal light spots are fringed
laterally by a dense accumulation of pigment that sets the dorsal pattern
off sharply from the sides of the specimen (Fig. 27a-c). Laterally, indi-
viduals are flecked with white iridiophores on a brown ground color.
Melanization, mostly present in older males, may or may not occur
(Fig. 27c, e, f). When it does, the quality of the dark color in fuscus is
decidedly brown as compared to black in auriculatus. This quality of
color is often diagnostic by itself in identifying specimens of both species
where they are found to occur together. The ventral color of fuscus also
contrasts with that of auriculatus. In fuscus the ventral ground color is
light but may be influenced by some degree of melanization (commonly
appears peppered or mottled by clusters of melanophores). In auri-
culatus the ground color of the venter is black and often is generously
sprinkled with white iridiophores.
Figure 28 illustrates a dorsal pattern variation in fuscus found in
some demes of the Yellow, Escambia, and Choctawhatchee River drain-
ages of western Florida. Dorsal light areas are not always present, and
the dorsal pattern consists of fine lines of melanocytes in the shape of
chevrons lying over myosepta. In a few localities where I found auricu-
latus and fuscus occupying the same habitat, mature males of fuscus were
extremely similar to equally sized auriculatus in color and in the absence
of a pattern (Fig. 29). However, inspection of tail morphology, jaw pro-
file dimorphism, and close scrutiny of the vestiges of the juvenile pattern
always effected a positive identification.

The desmognathincs investigated in this study exhibited a great deal
of variation in external color pattern, depending mainly on the amount
of melanization (deposition of melanin in the skin) occurring ontogenetic-
cally and geographically. Short-term (several hours to several days)

Vol. 18 No. 1


changes in individuals also occur, but are the result of another phenome-
non, metachrosis (change in dispersion of melanin in pigment cells). The
combination of the two darkening processes produces demes that are so
different in gross inspection of color patterns that different individuals
are hardly recognizable as the same species without reference to more
conservative characters.
Desmognathus auriculatus provides a good example to demonstrate
that the interaction of metachrosis and melanization leads to dramatic
differences in color patterns. Grobman (1950) reported that light-
colored individuals of auriculatus from Louisiana were experimentally in-
duced to become darker by metachrosis when left on dark substrates for
10 days. Dark Florida specimens became lighter when left on light sub-
strates. According to Grobman, Louisiana and Florida experimental
specimens converged on a similar color quality (although he reported
that they were still distinguishable).
Large series of specimens of D. auriculatus examined during this
study from Louisiana (Tulane University collections) were found to be
as lightly colored as some demes referred to fuscus from southern Missis-
sippi, although basic pattern differences existed. Living specimens col-
lected from southern Mississippi were also observed to be very light (Fig.
23). The reason for the decreased pigmentation of western Gulf drain-
age demes of auriculatus is not known precisely, but certain ecological
S correlations allow the following speculation. The very dark demes oc-
curring in Florida inhabit black, swampy mucks derived mainly from
cypresses (Taxodium spp.), gums (Nyssa spp.), and Magnolia spp. Muck
in western Gulf localities in general appears to be lighter in color, pos-
sibly because a greater percentage of its composition is due to contribu-
tions from hardwood litter (species of the genera Fagus, Quercus, Liquid-
ambar, Carya, Alnus, Ulmus, Acer, etc.). It is also noteworthy that, cor-
related with absence of black mucks west of the Pascagoula River system,
large deposits of Pleistocene loess are found in western Mississippi.
Whether the presence of these fine-grained aeolian sediments has some
effect on the ecology of desmognathine salamanders is not known, but it
is light in color and forms a pasty mud in the floodplains of streams in
which it is reworked (pers. obs.).
It was often noted in the field that the overall darkness of a specimen
was adjusted to that of the substrate upon which it was collected. Many
demes of both D. auriculatus and fuscus collected on dark substrates
were observed to have undergone a general decrease in intensity of dark
pigment if kept in white plastic containers several hours after collection.
Especially important in this regard are the observations made at Sweet-
water Creek (Sec. 31, T2NR7W, Liberty County, Florida), where both



fuscus and auriculatus were collected in a few instances under the same
debris in a mucky floodplain. Adults were very difficult to identify by
color pattern alone at the time of capture (Fig. 29). However, as little
as 30 minutes later, it was observed that, whereas auriculatus basically
retained an intense black color, fuscus always became lighter overall.
Even old males that had lost the juvenile blotching ontogenetically were
recognizable because a brownish quality of their ground color was ex-
posed by metachrosis (concentration of melanin particles in melano-
phores). These observations indicate that a considerable degree of the
quality of external appearance of desmognathines in this study is en-
vironmentally influenced. This single phenomenon alone probably ac-
counts for many of the different interpretations made by others in at-
tempting to distinguish species in the Coastal Plain.
Although D. fuscus primarily inhabits streamsides, where it is found
most abundantly in moist ravine situations in the area of this study, indi-
viduals nevertheless occupy mucky sites adjacent to ravine streams and
also seepage sites in the ravine floodplain or along valley walls. At the
headwaters of one upland stream this type of habitat was extensive and
appeared suitable for auriculatus (i.e., Beaverdam Creek steephead, Sec.
8, T1NR7W, north of Bristol, Liberty County, Florida), but was popu-
lated by fuscus. In these situations all size classes of fuscus usually take
on an overall dark wash due to the stellate condition of the melanophores.
A collection of fuscus (UF/FSM-16240 through 16259) from Kolomoki
Mounds State Park in southwest Georgia, and another from Calhoun
County, Florida (UF/FSM-17793 through 17821) are especially note-
worthy for this quality.
An important observation made in this study was a long-term (3-5
days) decrease in the intensity of black pigment in auriculatus, revealing
the persistence of a weak, underlying pattern in black Florida popula-
tions. The pattern is similar to that in demes from Louisiana and Missis-
sippi. On the basis of similarity of basal color pattern and of similarity
in other morphological characters, I conclude that Louisiana, Mississippi,
and Florida (swamp and floodplain muck) populations are conspecific.
There is no good evidence at present to indicate that taxonomic recog-
nition is warranted for light,. western populations and dark, eastern
Short-term (6-12 hours) color changes of dark individuals to a light-
er condition were frequently noted in D. fuscus from northern Florida
and southern Alabama (Figs. 30 & 31). Collections made in winter seem-
ingly had higher percentages of dark specimens. Often in the field I
initially identified intensely dark specimens of fuscus as auriculatus. Al-
though positive field identification of these dark individuals was possible

Vol. 18 No. 1


by examination of other morphological characters, dark fuscus invariably
became lighter if kept alive a few hours in collecting jars.

Valentine (1963) was the first student of desmognathine taxonomy
to use larval differences between Desmognathus auriculatus and fuscus
as diagnostic characters. He described and compared larvae from Missis-
sippi on the basis of body pattern, gill color and morphology, and relative
size at transformation. However, Goin (1951) had earlier noted differ-
ences between hatchlings of these taxa.
In Florida, D. auriculatus larvae were found to be larger at transfor-
mation (auriculatus: i= 21.4 mm, N=10; fuscus: = 13.5 mm; N=19);
to be more intensely pigmented (auriculatus: Fig. 20a, b; fuscus: Fig.
21a); to possess bushy gills, dark in color (Fig. 20a, b), as opposed to
tiny, iridiophore-populated gills of fuscus (Fig. 21a); and to be patterned
dorsally by much smaller light areas around dorsal neuromasts of Row 1
(Fig. 20b) than fuscus (Fig. 21a). In all these characters examined in
Florida material, the comparison between auriculatus and fuscus re-
vealed differences similar to those reported by Valentine (1963) for these
species in Mississippi.
Unquestionably, variation in the above characters exists. Drastich
(1927) performed experiments demonstrating that the number of gill fim-
briae decreases with availability of oxygen in larvae of Salamandra sala-
mandra. Because of time limitations, no experiments were carried out
on D. auriculatus during this study, but field observations established that
ecological isolation is greater between the larvae of the two species in
question than between transformed individuals from demes in the Apa-
lachicola and Ochlockonee river drainages. However, in ravine habitats
occupied by auriculatus in western Florida (which have not been open
to colonization by fuscus), the gills of larvae remain luxuriant with fim-
briae despite the presence of well-aereated water in which they were
found. Thus, it appears that large bushy gills are diagnostic in serving
to distinguish larvae of auriculatus from larvae of fuscus. At no time in
the study was it as difficult to identify larvae or adults as it was to iden-
tify juveniles.

Desmognathus brimleyorum is confined to rocky, gravelly streams in
the Ouachita Mountains. The adults were often difficult to collect be-
cause they were most often found under large rocks lying in the stream
or at its edge. Rock falls along upper portions of mountain streams were


also good collecting sites for adults. On removal of the rocks, adults
were found lying partially submerged in water. Rarely 'was one found
totally out of water, and invariably each specimen attempted to escape
into the stream. This species is one of the most aquatic in the genus.
The juveniles were readily collected by raking gravel along stream-
sides, but especially where seepage or a small freshet drains over gravel
and rock rubble. Thus, there appeared to be a microhabitat differentia-
tion between adults and juveniles. Larvae were found in small grained
gravel, under rocks in the stream, in small pools formed along stream
courses, and in moss covering rock faces where these occurred with water
flowing over them.
Desmognathus auriculatus was most often found under debris at the
edge of mucky, floodplain sloughs, at mucky edges of swampy lakes (Lake
lamonia, Lake Miccosukee, Florida) and in other sites of decomposing
muck associated with black waters. Individuals were often raked up
from below several inches of wet muck. In southeastern Georgia this
species was common in mucky streams draining the eastern flank of the
Tifton Uplands.
Considerable seasonal fluctuation of water levels occurs in the habi-
tat of D. auriculatus throughout the study area. When habitats are com-
pletely dry, the salamanders can be raked from under the surface of dried,
peaty muck at the lowest point of the depression where they aggregate.
Although D. auriculatus was rarely taken from situations other than
the above habitats, they were found in ravine streams in several places
in the study area (Econfina Creek steepheads, streams emptying into
the western end of Choctawhatchee Bay). These ravine environments,
however, yielded specimens only where organic debris had accumulated
along seepage sites. In such ravines, the basic muck microhabitat pref-
erence had not changed. There did not appear to be significant micro-
habitat differences between sex or age classes.
Those habitats primarily inhabited by Desmognathus fuscus in the
study area are best described as sandy-bottomed, wooded, upland ravine
streams. Usually the floor of a ravine valley is covered with multi-
colored leaf litter (hardwood origin), and the streamside has mossy
banks with accumulated organic debris supporting a large arthropod and
oligochaete fauna. Numerous seepage sources occur near the stream
head and along the downstream course, which maintains a relatively
permanent flow.
Of the three species studied, D. fuscus appeared to be the most
ubiquitous in its microhabitat selection. It was found in wet, mucky sites
along lower reaches of upland streams where seepage areas were not well
drained and partially decomposed hardwood litter had accumulated.

Vol. 18 No. 1


Although fuscus was not often found in river floodplain slough or swampy
terrace stream habitats within the geographical range of auriculatus, it
(fuscus) was taken from such habitats in southern Alabama, Louisiana,
Mississippi, and from localities in Florida west of the Choctawhatchee
River drainage. In Florida, these localities were always near or immedi-
ately adjacent to ravine habitats. Field observations indicated that at no
time was auriculatus found occupying a ravine habitat if populated by
fuscus, but as indicated above, fuscus could be found in some river flood-
plain habitats occupied by auriculatus. D. fuscus was never found in
coastal flatwoods swamps and streams, or in the center of extensive flood-
plain swamps of larger rivers. Relative dispersal ability into a greater
variety of habitats seemed to be higher for fuscus than for auriculatus.

Just as there are differences in microhabitat preferences between D.
fuscus and auriculatus in Florida, there exists a similar relationship in the
microhabitat preferences of two species of the genus Pseudotriton. Each
species is an ecological associate of one of the desmognathines. Invar-
iably, a species of Desmognathus and a species of Pseudotriton not only
occurred together in the same habitat, but occupied similar microhabitats
as well (differences in food habits and demography were at least two
parameters that prevented these microhabitat cohabitants from sharing
an identical ecological niche).
Pseudotriton montanus was found to be a muck dweller along
swampy streams of floodplain terraces, coastal lowlands, and in upland
mucky streams along the eastern flank of the Tifton Uplands. Pseudotri-
ton ruber was found exclusively in deep, upland ravines. Whereas D.
fuscus and auriculatus have higher population densities and wider dis-
persion throughout their respective habitats, the two species of Pseudo-
triton were comparatively less abundant in post-larval age classes and
more highly localized in the different environments as far as it was pos-
sible to determine.
Desmognathus fuscus and Pseudotriton ruber are ecologically sym-
patric, forming a species pair largely confined to upland stream habitats.
D. auriculatus and P. montanus form another pair, ecologically sympatric
in river floodplains, coastal flatwoods streams, and in mucky streams
draining the eastern slopes of the Tifton Uplands. It was found during
the course of this study that these two pairs are also geographically sym-
patric in the study area, the geographic range of each member of both
pairs coinciding almost exactly with the range of the other member.


Desmognathus brimleyorum Stejneger
1. large-sized (adults greater than 60 mm SVL)
2. fungiform crown morphology
3. no dimorphism between dentary and maxillary teeth
4. no sexual dimorphism in jaw profile
5. adults with no distinct pattern
6. large prearticular spine
7. premaxillary fontanelle about 50 percent of length of premaxillary spines (well
8. neuromast vestiges of Row 2 and 3 surrounded by iridiophores, usually obvious
as lateral lines of dots
9. tail cross-section round at base; tail keeled, compressed posteriorly
10. found in Ouachita Mountains of Arkansas and Oklahoma

Desmognathus auriculatus Holbrook
1. medium-sized (adults 45-60 mm SVL)
2. piercing, recurved dentary teeth
3. dimorphism between dentary and maxillary teeth
4. no sexual dimorphism in jaw profile
5. adults with a weak, but distinctive pattern in west (Louisiana, Mississippi), ob-
secured in east (Florida, Georgia), but inducible by metachrosis
6. prearticular spine not well developed
7. premaxillary fontanelle nearly closed, spines fused posteriorly
8. neuromast vestiges of Row 2 (on tail) and Row 3 usually surrounded by iri-
diophores, usually obvious as lateral lines of dots in adults
9. tail cross-section round to trigonal at base; tail keeled; compressed to tip
10. range incompletely known to the northeast and west of Florida, but distinctly
Coastal Plain

Desmognathus fuscus (Rafinesque)
from Florida
1. small-sized (adults 40-50 mm SVL)
2. piercing, recurved dentary teeth
3. strong dimorphism between dentary and maxillary teeth
4. extreme sexual dimorphism in jaw profile
5. adults usually with an elaborate pattern derived from juvenile blotched condition
6. prearticular spine moderately developed
7. premaxillary fontanelle well developed
8. neuromast vestiges usually not set off from lateral color pattern by iridiophores
as lines of dots
9. tail cross-section round; tail not keeled, tapering to a terete filament at tip
10. ranges from at least Piedmont of Georgia to Florida, then westward at least to
the Alabama River drainage (possibly to the Mississippi River)

Desmognathus fuscus (Rafinesque)
from New York
1. medium-sized (adults 45-60 mm SVL)
2. fungiform crown morphology
3. no dimorphism between dentary and maxillary teeth
4. no sexual dimorphism in jaw profile
5. adults usually with vestige of juvenile pattern

_ ___ __ ____ _ _


6. prearticular spine moderately developed
7. premaxillary fontanelle well developed
8. neuromast vestiges of Row 2 usually present on tail as line of dots
9. tail cross-section at base trigonal; tail compressed to tip
10. ranges from Ontario to at least the Piedmont of North Carolina (inaccurately

Desmognathus ochrophaeus Cope
from southern Appalachians
1. small-sized (adults 40-50 mm SVL)
2. piercing, recurved dentary teeth
3. strong dimorphism between dentary and maxillary teeth
4. extreme sexual dimorphism in jaw profile
5. adults usually with an elaborate pattern derived from juvenile blotched condition
6. prearticular spine moderately developed
7. premaxillary fontanelle well developed
8. neuromast vestiges usually not expressed in adult as lateral lines of dots
9. tail cross-section round; tail not keeled, tapering to a terete filament at tip
10. ranges throughout the southern Appalachians


STATUS OF Desmognathus brimleyorum STEJNEGER

In this study, Ouachita Mountain desmognathines were found to be
distinguishable from adjacent Arkansas and Louisiana Coastal Plain pop-
ulations on the basis of the following characters: tooth morphology, rela-
tive size, color pattern, prearticular spine morphology, larval morphology,
and jaw profiles. Unpublished dissertation research of Chaney (1958,
p. 84) indicates that Ouachita Mountain populations near Russellville,
Arkansas are significantly different from Louisiana desmognathines in:
"(1) head length/head width ratio; (2) size at hatching; (3) growth
rate; (4) maximum size attained; (5) tail length; (6) time of attainment
of sexual maturity; (7) number of eggs deposited; (8) color patterns
possessed by the larvae and the adults." Thus, the results of both studies
clearly indicate that populations in the Ouachita Mountains are strongly
different from surrounding desmognathines in at least eleven morphologi-
cal and developmental characters.
In the original description of D. brimleyorum, Stejneger (1895)
pointed out that maxillary and mandibular teeth were "all very blunt."
Other characters he mentioned are shared with one or more other species
except one, "the anterior glandular prolongation of the lower lip," which
Stejneger claimed was absent. All adult males I examined possessed a
mental gland that is characteristic for all species in the genus Desmogna-
* thus and is responsible for an anteriorly directed protrusion of the fleshy
portion of the mandible. However, Stejneger's observation that a "pro-
longation . is absent," is essentially correct. The mental gland is



proportionately smaller in D. brimleyorum males than in other species
and is difficult to see without microscopic examination. Furthermore,
the elongate, monocuspid premaxillary teeth (which occur in direct physi-
cal contact with the mental gland when the mouth is closed) of male D.
brimleyorum are also proportionately less exaggerated.
In summary, populations of Desmognathus from the Ouachita Moun-
tains area 1) differ from their congeners in the quality of at least 12 mor-
phological and developmental characters, and 2) occur in rocky-bottomed
streams in a unique geological and physiographic region on the periphery
of the range of the subfamily Desmognathinae. I believe this constitutes
sufficient evidence that these populations represent a valid species, in
agreement with Stejneger's original evaluation. The approximate geo-
graphic range of Desmognathus brimleyorum is mapped in Figure 32.

THE RELATION OF D. auriculatus TO D. fuscus IN FLORIDA
MORPHOLOGICAL DIFFERENCES.-In Florida, desmognathines (except
Desmognathus monticola) can be separated into two categories on the
basis of differences in osteological and external morphological characters.
D. auriculatus shows no sexual dimorphism in jaw profile, has a reduced
premaxillary fontanelle, is larger than fuscus in comparisons between the
same age-classes, possesses a compressed tail that retains a vestige of a
dorsal fin in post-larval life, differs from fuscus in larval morphology, and
attains an overall darker pigmentation that obscures an underlying pat-
tern also different from that of fuscus.
D. fuscus may be characterized by having strong sexual dimorphism
in jaw profile; a well-developed premaxillary fontanelle; small relative
size; a terete, finless tail tapering to a round point; distinctive larval mor-
phology; and strong dorsal pattern in post-larval life (except as some-
times obscured by melanization in old males).
All the above characters taken collectively allow positive identifica-
tion of specimens as fuscus or auriculatus. However, jaw profile dimor-
phism, which is strongly developed and always present in fuscus in the
Apalachicola and Ochlockonee River drainages, becomes less marked in
western Florida drainages of the Choctawhatchee, Yellow, Blackwater,
and Escambia Rivers. In some demes from western Florida, dimorphism
is well developed, whereas it is nearly absent in others. This character
alone serves to identify mature males of fuscus in the Apalachicola and
Ochlockonee River drainages, but is less reliable when considered by it-
self in western Florida.
It is significant to note that in the entire subfamily the character of
jaw profile dimorphism is most abundantly and consistently present in

_ ___ __ __ ~__

Vol. 18 No. 1


populations of D. fuscus and ochrophaeus from the Piedmont and south-
ern Appalachians, respectively. The only watershed in the area of this
study that has its source in either the Piedmont or the southern Appala-
chians is the Apalachicola ( = Chattahoochee) River. Variation in jaw
profile dimorphism is negligible in Florida demes along this watershed,
suggesting no recent barriers to gene flow. Equally significant is the fact
that, while virtually no reduction of jaw profile dimorphism was observed
from the southern Appalachians, variation in this character becomes
greater as one moves westward in the upper reaches of the Alabama-
Tombigbee River drainage, crossing drainage divides. Hence, it is
logical that gene flow occurs more readily within drainage systems than
between them. This assumption underlies all conclusions reached in this
study concerning genetic relationship between demes of fuscus based on
morphological similarity.
The other five characters mentioned that are significant in identifica-
tion of D. auriculatus and fuscus do not exhibit consistent interdeme
variation. Tail shape may be affected by the state of health or diet of
individuals. Variation in relative size is possibly affected by density-
dependent factors (no direct evidence of this was gathered). Occasion-
ally individuals and populations were collected which were smaller than
average for the species, but in general fuscus was smaller than auriculatus
at any given age. Specimens occasionally were examined that fell into
the range of variation in morphology of the premaxillary fontanelle of the
other species, but in general this character is reliable. Color pattern is
probably the one aspect of the phenotype most influenced by the local
environment, and because of this it is the least reliable when taken by it-
self. Color pattern is so variable, depending on the substrate, that speci-
mens of both species collected from the same locality were often nearly
identical. However, metachrosis usually revealed basic differences if spe-
cimens were maintained alive several hours after capture on selected
substrates. Larval morphology is also somewhat affected by the environ-
ment, but in this study larvae were always recognizable as either auricu-
latus or fuscus. The number of gill fimbriae may vary with oxygen and
water available in the larval environment (Drastich 1927), but the pres-
ence or absence of iridiophores on gill rami and larval size, color, and
color pattern still serve to distinguish fuscus from auriculatus.
ECOLOGICAL DIFFERENCES.-The most striking differences between
Desmognathus fuscus and D. auriculatus within the geographical limits
of this study were habitat and microhabitat preferences. In those drain-
age systems where both forms occurred, auriculatus was exclusively found
in floodplain and coastal flatwoods swampy environments, where it was
most often collected from sites of decomposing organic litter and from


along the margins of standing or slow-moving water. Usually specimens
were discovered under the larger debris such as logs and leaf piles in
muck. Often individuals were raked from under the surface of stagnant,
shallow muck sites. Individuals invariably attempted to retreat into fluid
muck or shallow, standing water.
D. fuscus was most commonly found in shaded, hillside ravines such
as occur along the eastern escarpment of the Tallahassee Red Hills-
Tifton Uplands physiographic region in Florida and adjacent Georgia.
This type of environment offers a variety of microhabitats which were
occupied by fuscus. The borders of ravine streams, especially where fri-
able materials were present for burrowing, were teeming with fuscus.
Density was highest, however, in hillside seepage sites where ground
water and burrowing arthropods provided a wealth of anastomosing sub-
terranean channels supplied with abundant moisture. D. fuscus was not
often encountered more than a few inches from streamsides or seepage
water. Some seepage sites (especially downstream from ravine heads
where the gradient lessens and the ravine floor becomes wider) are some-
what mucky where leaf litter accumulates faster than it decomposes.
These were also inhabited by fuscus.
In parts of the study area outside the geographical range of D. auri-
culatus, fuscus colonizes floodplain and other habitats where one would
expect to find auriculatus (parts of western Florida and Coastal Plain
Alabama). Metachrosis and possibly local selective pressures produce
demes that resemble auriculatus in the degree of melanization of color
pattern. Proper identification is dependent on other characters, especial-
ly the morphology of jaw profile, premaxillary fontanelle, and the tail.
There was greater latitude in habitat selection in D. auriculatus
where this species occurred outside the geographical range of fuscus (Fig.
33) in southeastern Georgia and northeastern Florida. In the headwaters
of tributaries of the Ochlockonee, St. Marks, Aucilla, Suwannee, and
Satilla rivers, auriculatus was found in abundance (Fig. 34). D. auricu-
latus was also collected from steephead ravines of the Econfina River and
streams of similar aspect draining into western Choctawhatchee Bay.
However, in none of the upland stream environments were microhabitat
preferences significantly different from those where auriculatus and fuscus
were geographically sympatric. Such upland streams are the headwater
tributaries of the Ochlockonee, St. Marks, Aucilla, Suwannee, and Satilla
rivers, which drain a gently seaward sloping, ancient plain (Tifton Up-
lands). These headwater streams do not dissect this plain deeply and
most have wide, low-gradient floodplains. These are characteristically
very swampy, containing large accumulations of decomposing organic
litter derived mostly from gums (Nyssa spp.), cypress (Taxodium sp.),

___ ____ __

Vol. 18 No. 1


Magnolia virginiana, titi (Cyrilla racemiflora), and other hydrophilic
plants. True ravines occur along eastern escarpments of the larger rivers
here, but these are notably depauperate in salamanders except where oc-
casional mucky sites have developed.
The ravine steephead habitats along the Econfina River and lower
Choctawhatchee Bay are peculiar in their geological development (Sharp
1938), and differ significantly in biology from most ravine habitats along
the eastern escarpment of the Apalachicola River. Ravines in the latter
area develop from surface runoff over relatively impermeable elastic sedi-
ments of the Miocene Hawthorne Formation. These ravines are V-like
in cross-section and have a fairly steep gradient in their upper reaches.
These ravines contain a diverse hardwood forest comprised of many
northern species, wide-ranging southern species, and a number of ende-
mics. The unique northern aspect of these ravines is well known botan-
ically. On the other hand, steephead ravines along the Econfina River
and lower Chochawhatchee Bay develop in loose sands of younger age
which are relatively permeable to water. Surface runoff is much reduced.
Steepheads form from lateral sapping of the water table (Sellards and
Gunter 1918, Sharp 1938, Vernon 1942). Valley formation is U-shaped
S since erosion is headward along the base of valley walls. The plant com-
munities common to these ravines differ in many ways from those along
the Apalachicola River eastern escarpment. Two of the dominants are
titis (Cyrilla racemiflora and Cliftonia monophylla). The ravine sides
are very dry upslope from the stream bottom, and support a number of
xerophilic plants. The muck and soil of these ravines are derived from
different plant species than those dominants found in the Apalachicola-
Ochlockonee ravines. This type of ravine contains a great many mucky
seepage sites, and the gradient near the ravine steephead is not too differ-
ent from downstream. This particular type of habitat seems equally suit-
able to colonization by muck-'loving" auriculatus or by ubiquitous fuscus.
In fact, this study has indicated that fuscus competitively displaces auri-
culatus from these ravines when both species have the opportunity to
colonize them (Fig. 35).
The northward flowing ravine steephead tributaries of the Yellow
River, whose watershed divide is common to southward flowing steep-
head ravines along Choctawhatchee Bay, are nearly identical to the latter
geologically and biologically. Figure 35 shows that D. auriculatus occurs
exclusively in these habitats at the western end of the bay. D. auriculatus
is also found in the swampy floodplain of the Yellow River at the con-
fluences of ravine steephead streams there and thus is capable of coloniz-
ing the upper reaches of these tributaries. However, the headwaters of
these ravines are instead populated by D. fuscus (Fig. 35).


It is my belief that fuscus has not yet invaded the ravines occupied
by D. auriculatus that drain into western Choctawhatchee Bay because
physical barriers exist to the dispersal of fuscus (which ranges northward
throughout the Choctawhatchee River drainage to its headwaters). One
such barrier may be the discontinuous connection of the ravines between
the eastern and western ends of the bay. During Pleistocene glacial
maxima all tributaries of the lower Choctawhatchee River may have
been ravines, but the presence in western ravine streams of the endemic
darter, Etheostoma okaloosae, and the absence of its close relative, E.
edwini, in these same drainages (Collette and Yerger 1962) indicates that
there may never have been a connection between streams at both ends
of the present bay. Although fuscus is not present in ravine streams
along the western end of Choctawhatchee Bay, it occurs in those draining
into the eastern end. A similar distribution is recorded for Etheostoma
edwini (Collette and Yerger 1962).
As already mentioned, D. fuscus is assumed to disperse within drain-
ages more readily than between them. This is because stream divides
become well-defined barriers of inhospitably dry habitats for salamanders
in the area of this study (usually pine-turkey oak, sand ridge commun-
ities). Swampy habitats of low elevation along the Gulf Coast also act as
effective barriers to the dispersal of fuscus, inasmuch as aquatic, larval
individuals appear to be more strictly adapted to the ecology of ravine
habitats than transformed, terrestrial specimens; in contrast, however,
such habitats are virtual highways for auriculatus. Hence auriculatus
is capable of colonizing tributary ravine streams by immigration upriver
from alluvial plains and coastal swamps that are continuously connected
along the Gulf Coast of Florida from the peninsula to Pensacola Bay (i.e.,
ravines and steepheads along Econfina Creek, which has probably never
been connected with either the Apalachicola or the Choctawhatchee
Rivers; fuscus presently occurs on both sides of this small, independent
drainage system, but not in it).
Desmognathus fuscus disperses downstream from upland population
centers initially, and displaces auriculatus whenever fuscus is able to sur-
mount ecological barriers to its dispersal. Furthermore, in the few local-
ities found in this study where optimum habitats for both species were
immediately adjacent (i.e., Sweetwater Creek and Ocklawaha Creek lo-
calities), the two species did not hybridize and were mutually exclusive
in their respective microhabitats except along the zone of contact.
I consider the above to constitute evidence for the validity of D.
auriculatus and D. fuscus as full species within the study area. This is
in agreement with Valentine's (1963) conclusions regarding the relation-
ship between auriculatus and fuscus in Mississippi. Also, Chaney (1949,


1958) reported that his studies of desmognathines from southern Louisi-
ana and Mississippi revealed two larval and adult morphotypes. He did
not venture a speculation as to the taxonomic identity of the morphs.
During examination of his extensive collections, I concluded that fuscus
and auriculatus both are present in the Florida parishes of Louisiana.
However, it is important to note that Mississippi and Louisiana specimens
are the most difficult of all to distinguish morphologically in the entire
range of sympatry between auriculatus and fuscus. All the characters in-
vestigated in this study and found to be significant in separating the two
species in Florida converge in specimens of the two forms from those
GEOGRAPHICAL DISTRIBUTION.-Both Desmognathus fuscus and auri-
culatus range widely beyond the geographical boundaries of this study.
The evolutionary relationships between these species in other parts of
their ranges may be different from those postulated herein. Hence, Fig-
ures 33 and 35 represent the distribution of fuscus and auriculatus strict-
ly as determined for specimens directly examined (and in most cases
collected) by me. Extralimital studies are presently under way.
D. fuscus ranges throughout the Apalachicola-Flint-Chattahoochee
* river system from the Piedmont of Georgia to the ravines dissecting the
western escarpment of the Tallahassee Red Hills physiographic region.
It is found eastward to ravines along Lake Talquin (an artificial im-
poundment of the Ochlockonee River) and throughout the Little River,
a tributary of the Ochlockonee. All efforts to collect fuscus in the Och-
lockonee above the confluence of the Little River were unsuccessful. I
do not believe fuscus occurs above this confluence for the following
reasons: the Ochlockonee River formerly flowed southward to the coast
through the Lake Bradford chain and the abandoned valley on which the
old Tallahassee Municipal Airport was built. The Ochlockonee has been
captured by a tributary of the Little River (Hendry and Sproul 1966)
so recently that fuscus has not yet dispersed northward through the wide,
swampy floodplain above the site of stream capture. As evidence of this,
only auriculatus is commonly collected in stream habitats above the level
of capture in the Ochlockonee River, but only fuscus is found in similar
situations in northern tributaries of the Little River immediately to the
west (Fig. 33).
Desmognathus fuscus has entered the lower Chipola River drainage
where that stream breaches the remnant uplands that used to form a con-
tinuous connection between the Western Highlands and the Tallahassee
S Red Hills physiographic regions (Fig. 33). In general, a hiatus in the
range of fuscus seems to occur between the Choctawhatchee and the
Apalachicola rivers in Florida (probably due both to inadequate collect-


ing and scarcity of ravines). A few fuscus were found in western ravines
of the Holmes Valley Escarpment, but none on other remnant highlands
between these two rivers. Geologists (Puri and Vernon 1964) speculate
that the Apalachicola River once flowed westward across the Marianna
Lowlands, joining the ancestral Choctawhatchee to breach the highlands
along the present course of the latter river. The occurrence of fuscus in
the Choctawhatchee River drainage may be due to colonization during
this past connection with the Chattahoochee-Apalachicola River. Head-
water stream capture or fortuitous migration over drainage divides shared
with the Alabama or Chattahoochee systems, or both, also may explain
the presence of fuscus throughout the Choctawhatchee drainage. It may
be significant that fuscus demes from this system are variable in some
morphological characters that do not vary in fuscus in the Apalachicola-
Chattahoochee system. It is possible that after colonization of west
Florida river systems, reduced or discontinued gene flow with the parent
population, coupled with a change in selective pressures, allowed gene
frequencies to change. This would account for increased variation in
otherwise strongly conservative characters.
Desmognathus fuscus occurs throughout the Escambia-Conecuh
River drainage in Florida and Alabama. Demes were sampled in this
study from ravines emptying directly into Escambia Bay below the mouth
of the Escambia River. Also, fuscus was collected in the floodplain of
the Escambia and Conecuh tributaries where auriculatus was expected.
Much variation in size, melanization, jaw profile dimorphism and other
characters was noted between demes, especially in Florida localities.
Demes sampled from deep ravines in the Red Hills of southern Alabama
(headwaters of the Escambia-Conecuh system) were less variable rela-
tive to each other and more closely approximated the phenotype of fuscus
from the Apalachicola-Chattahoochee drainage. I feel this is due to the
closer proximity of the Escambia-Conecuh headwater populations to the
Apalachicola-Chattahoochee River populations (Fig. 36), where gene
flow may still occur. An even more convincing argument, however, may
be made for the possibility that because these headwater populations in
Alabama inhabit the deepest and most continuous ravines in the system,
selective pressures still favor strong expression of character states.
Probably no greater interdeme variation in fuscus was observed than
that from the Blackwater and Yellow River drainages. I hypothesize
that fuscus has met with somewhat different selective pressures in adapt-
ing to the microhabitats available to it in these two river systems. The
ravine habitats are different because the underlying geology of permeable
Plio-Pleistocene elastic sediments has allowed steephead types of ravines
to develop near the coast. Shallow, young valleys have developed in


headwater tributaries. The vegetation is unique in these streams because
of the presence of both titis (Cliftonia monophylla and Cyrilla racemi-
flora), pitcher plants, sphagnum, and a host of other plants most com-
monly found in flatwood bays and not in upland stream valleys.
Desmognathus fuscus probably entered the Blackwater and Yellow
River drainages via past connections of their floodplains with that of the
Escambia River during lower sea levels in the Pleistocene. Examination
of drainage patterns in Figure 36 reveals that the Escambia and Chocta-
whatchee drainages surround those of the Blackwater and Yellow rivers.
Because of the youth of the latter systems, headwater stream capture has
probably not occurred. D. fuscus demes in the Blackwater and Yellow
rivers not only have become adapted to somewhat different ecological
conditions from those of ancestral Escambia River stock, but have prob-
ably been protected from the homeostatic effect of gene flow with the
Escambia gene pool during the high stand of seas in Recent times.

APPALACHIAN ochrophaeus
Skulls and preserved series of specimens from the type locality of
Desmognathus fuscus (northern New York) were examined to determine
the characteristics of the nominate race. These states were used as
standards with which the morphology of populations of Piedmont and
Coastal Plain desmognathines was compared to assess taxonomic related-
It is now apparent that the category of D. fuscus is a heterogeneous
one if all southern populations of desmognathines heretofore referred to
this species remain lumped under this name. All the Florida, Georgia, and
Alabama Piedmont and Coastal Plain specimens examined in this study
agreed on a morphological basis more closely with ochrophaeus from the
southern Appalachians than with fuscus from New York. The southern
limit of specimens examined that agree with New York fuscus was about
Uwharrie National Forest in the Piedmont of North Carolina. The exact
delimitation of the southern limit of the range of fuscus has yet to be de-
termined. A study of what happens in the zone of contact of northern
fuscus, southern fuscus, and auriculatus in North and South Carolina
would go far to elucidate evolutionary relationships between these three
taxonomic entities.
The most readily observed features of D. fuscus (New York) are
fungiform tooth morphology, compressed tail, and absence of sexual di-
morphism in jaw profiles. All literature accounts of fuscus clearly indi-
cate that little or no jaw sinuosity (external commissure somewhat re-



fleets the profile between jaw bones) and a compressed, keeled tail (Cope
1889, Dunn 1917, Bishop 1943) are present in specimens from New York
and adjacent northeastern states.
Specimens examined by me that previously were assigned to D. fuscus
from Piedmont localities of Georgia typically have jaw profile dimorph-
ism strongly developed in adults, teretely tapering tails, and teeth di-
morphic between maxilla and mandible. These character states are also
found in adjacent populations of ochrophaeus from the southern Appala-
chians. This may partly explain why other workers have been unable
to distinguish some populations of fuscus from ochrophaeus in this region
(Martof and Rose 1963).
The major conclusion obtained from comparisons of New York D.
fuscus with southern material previously referred to by that name is that
Florida and Georgia "fuscus" very strongly resemble ochrophaeus from
the southern Appalachians of Georgia and not topotypic specimens of
fuscus. It is notable that relative size and color pattern is also more
closely alike between southern fuscus and ochrophaeus than between
southern and northern fuscus.


1. Desmognathus brimleyorum Stejneger is a valid species confined
to the Ouachita Mountains of Arkansas and Oklahoma.
2. Desmognathus auriculatus in Florida is considered to be a valid
species; D. fuscus carri is considered to be an ecophenotype of D. auri-
3. Desmognathus "fuscus" ranges throughout Florida west of the
Ochlockonee River.
4. Desmognathus "fuscus" in the Apalachicola and Ochlockonee
river drainages is closely similar morphologically to demes of this species
upstream in the Flint-Chattahoochee rivers. These resemble D. ochro-
phaeus more closely than the nominate race of D. fuscus from northern
New York and adjacent states.
5. Intensive study of qualitative characters indicates strong, localized
variation in D. fuscus. This variation, and peripheral populations ex-
amined, indicates a critical need for further study of southern "fuscus"
and ochrophaeus, using a multidisciplinary approach.


Desmognathus aeneus

Georgia-Dawson Co.: UF/FSM*-27271 (1m).

Desmognathus auriculatus

Florida-Bay Co.: DBM-1488 (1m, If), DBM-1620 (2m, ij); Columbia Co.: UF/
FSM-4076-1 (Im); Gadsden Co.: DBM-1660 (1m); Jefferson Co.: no # (Im);
Leon Co.: FSU-598 (1m), DBM-1229 (Im, 2j), DBM-1511 (1m), no # (1m,
If, 1j); Liberty Co.: DBM-1527 (1m); Marion Co.: DBM-1649 (1m), no #
(lm); Okaloosa Co.: FSU-645 (1m), DBM-1255 (1m), DBM-1563 (1m),
DBM-1725 (Im); Wakulla Co.: FSU-785 (1m); Washington Co.: DBM-1519
Georgia-Charlton Co.: DBM-1551 (1m, If); Grady Co.: DBM-1281 (Im); Irwin
Co.: DBM-1626 (1m); Liberty Co.: DBM-1545 (1m); Thomas Co.: DBM-1634
Louisiana-Washington Parish: TU-11832 (Im, If).
Mississippi-Forrest Co.: DBM-1537 (1m).
South Carolina-Berkeley Co.: DBM-1546 (1m).

Desmognathus brimleyorum
Arkansas--Clarke Co.: DBM-1295 (12 juv ad); Garland Co.: NELSC-2350 (1m);
Polk Co.: DBM-1245 (1m, If), DBM-1246 (2m, If, Ij), no # (Im); ?Co.:
USNM-118476 (1m).
Oklahoma-LeFlore Co.: DBM-1476 (Im), DBM-1711 (Im, Ij).

Desmognathus cf. fuscus
Alabama-Baldwin Co.: DBM-1219 (1m); Butler Co.: DBM-1232 (1m), DBM-1428
(2m); Clay Co.: DBM-1607 (Im); Cleburne Co.: DBM-1186 (im); Conecuh
Co.: DBM-1427 (2m); Dale Co.: DBM-1189 (im); Houston Co.: UF/FSM
9316-2 (1m); Tuscaloosa Co.: UAIC 49-1241 (1m).
Arkansas-Cross Co.: DBM-1589 (1m).
Florida-Escambia Co.: DBM-1240 (im), DBM-1241 (1m), DBM-1242 (1m),
no # (1m); Gadsden Co.: DBM-1171 (Im, lj); Leon Co.: DBM-1490 (1m);
Liberty Co.: DBM-69CC (2m, If, 1j), DBM-1154 (1m), DBM-1527 (2m);
Okaloosa Co.: DBM-1263 (1m), DBM-1558 (1m); Santa Rosa Co.: DBM-1220
(lm), DBM-1259 (2m, 2f), DBM-1260 (1m), DBM-1667 (1m); Walton Co.:
DBM-1222 (1m), DBM-1235 (1m), DBM-1236 (1m).
Georgia-Bibb Co.: UF/FSM 14019-2; Clarke Co.: UG-60 (1m); Fulton Co.: no #
(lm); Randolph Co.: DBM-1291 (2m).
Louisiana-Ouachita Par.: NELSC-17147 (1m); St. Tammany Par.: TU-2870 (lm);
Union Par.: NELSC-24002 (1m).
Kentucky-Harlan Co.: NELSC- (no #) (1m).
Massachusetts-Franklin Co.: MCZ A-82583.
Mississippi-Benton Co.: DBM-1592 (1m); Forrest Co.: DBM-1537 (1m); Lawrence
Co.: TU-14136 (1m); Tishomingo Co.: DBM-1188 (1m), TU-14635 (1m);
Walthall Co.: DBM-1534 (1m).

*Collection abbreviations are explained in the Acknowledgments (p. 8); DBM-
numbers are collections made by the author expressly for this study.


North Carolina-Bladen Co.: DBM-1547 (2m); Caswell Co.: DBM-no # (1m);
Montgomery Co.: DBM-1601 (im); Orange Co.: DBM-1704 (1m).
New York-Orange Co.: USNM-23222 (1m), USNM-23227 (If); Warren Co.:
USNM-80232 (1m), USNM-80234 (1m), USNM-80237 (If).
Tennessee-Giles Co.: JWR (1m); Montgomery Co.: NELSC-15658 (1m), NELSC-
19503 (1m); Sevier Co.: NELSC-10588 (1m), NELSC-11812 (1m), NELSC-
12001 (lm), NELSC-12154 (1m).
Texas-Tyler Co.: RW-627 (lm).

Desmognathus monticola
Alabama-Butler Co.: DBM-1232 (1m), DBM-1575 (1m); Clay Co.: DBM-1607
(lm); Randolph Co.: DBM-1185 (Im, If, Ij).
Georgia-Lumpkin Co.: UAIC 68-(179-220) (Im, If).
North Carolina-Avery Co.: MCZ-6463 (1m).
Tennessee-Sevier Co.: NELSC-11809 (1m), NELSC-12154 (1m).
Virginia-Grayson (?) Co.: MCZ-6558 (1m).
West Virginia-Greenbriar Co.: no # (1m, If).

Desmognathus ochrophaeus
Georgia-Rabun Co.: TTRS-10 (Im), USNM-147576 (Im), USNM-147577 (1m),
USNM-147579 (If), USNM-147597 (If); Towns Co.: USNM-147668 (1m),
USNM-147683 (If).
Maryland-Garrett Co.: USNM-101903 (1m), USNM-101905 (If).
New York-Cattaraugus Co.: MCZ-33834 (1m), MCZ-33868 (If), MCZ-65690
North Carolina-Buncombe Co.: UC-1314-4 (1m); Cherokee Co.: USNM-147688
(lm); Swain Co.: DBM-1398 (1m).
Tennessee-Polk Co.: USNM-147802 (1m), USNM-147804 (If).

Desmognathus "ocoee"
Tennessee-Polk Co.: USNM-147802 (1m), USNM-147804 (If).

Desmognathus quadramaculatus
Georgia-Fannin Co.: UG-677 (lm); Habersham Co.: UG-96 (1m), no data (1m);
Haywood Co.: DBM-1400 (1m), no data (1m).

Desmognathus wright
North Carolina-Avery Co.: UF/FSM-8241-1 (im).

Leurognathus marmoratus
Georgia-Rabun Co.: USNM-155979 (1m), USNM-155985 (If), USNM-156484
(lm), USNM-156508 (If).
North Carolina-Haywood Co.: DBM-1399 (1m).
South Carolina-Oconee Co.: AUM-12109 (1m).

Phaeognathus hubrichti

Alabama-Butler Co.: DBM-1232 (Im, 2f), DBM-1413 (2m, If), no # (1).



Desmognathus auriculatus

Alabama-Baldwin Co.: AU-10543, AU-13008, USNM-57226, USNM-57227; Hous-
ton Co.: AU-2173?; Mobile Co.: USNM-57228, -57232 to -57236, UF/FSM-
3039 (4).
Florida-Alachua Co.: UF/FSM-14111, -333, -600 (23), -1460 (4), -1522 (2),
2031 (1), -3086 (6), -3675, -8073, -9218, -55 (6), -173 (4), -1094 (8), -14030,
-14230 (2), -16046, -17178, -18311 to -13, -25869, -25872 to -75, SRT-572;
Baker Co.: UF/FSM-256 (2), -2958, -2368 (12), -26076, -26078, -26080,
-26082, -26084, -26084, -26087, -26077 to -86, USNM-136950 to -136954; Cal-
houn Co.: UF/FSM-27249 to -55, DBM-1179 (3), FSU-35 (3), FSU-154 (5),
FSU-440 (2), FSU-167 (8), RW-3061 (1); Bay Co.: DBM-1488 (25), DBM-
1499 (2), DBM-1517 (ca. 5), DBM-1620 (23), DBM-1621 (9), DBM-1662
(ca. 10), DBM-1663 (1), DBM-1664 (1), DBM-1685 (1), DBM-1696 (5);
Columbia Co.: UF/FSM-4076 (17); Clay Co.: UF/FSM-1078 to -80; Dixie Co.:
UF/FSM-387 (6); Duval Co.: Vockell (1); Escambia Co.: TU-16572 (20);
Franklin Co.: UF/FSM-9258 (4); Gadsden Co.: DBM-1153 (13), DBM-1283
(ca. 10), DBM-1340 (ca. 15), DBM-1538 (ca. 10), DBM-1539 (1), DBM-
1544 (1), DBM-1622 (9), DBM-1635 (7), DBM-1654 (5), DBM-1660 (4),
DBM-1700 (2); Gulf Co.: UF/FSM no # (4), -7833 (1), -9147 (2), -2676
(7), 9143 (7); Hillsborough Co.: UF/FSM-3036 (3), -18316, SRT-1126 (2),
SRT-1127 (3), SRT-1174 (1), SRT-1176 (1); Holmes Co.: UF/FSM-2680 (2),
AUM-10531, -2, -3; Jackson Co.: DBM-1595 (6), DBM-1596 (ca. 15), DBM-
1656 (10), UF/FSM-6566, -6760 (6), -6761, -7761, -7762 (25), -7763, -9395
(3), -391 (4), -704 (3), -2646 (2), -1233 (19), -26068 to -74, TU-13631 (8),
RW-2997 (1), TU-13395 (73), TU-14909 (1), TU-13320 (1), TU-13374 (1),
TU-13424 (13); Jefferson Co.: DBM-1305 (ca. 20), DBM-1541 (4); Lake Co.:
UF/FSM-18292 to -18310, SRT-512, SRT-278; Leon Co.; DBM-1152 (2),
DBM-1206 (ca. 8), DBM-1221 (ca. 10), DBM-1229 (ca. 20), DBM-1276
(10), DBM-1391 (ca. 10), DBM-1494 (1), DBM-1511 (1), DBM-1512 (1),
DBM-1560 (43), DBM-1619 (10), FSU-38 (7), FSU-818 (8), FSU-473 (6),
FSU-193 (8), FSU-240 (1), FSU-152 (2), FSU-40 (1), FSU-643 (10), FSU-
508 (2), FSU-827 (2), FSU-205 (4), FSU-681 (4), FSU-525 (1), FSU-662
(7), FSU-339 (9), FSU-276 (1), FSU-428 (3), FSU-816 (1), FSU-511 (1),
FSU-600 (2), FSU-444 (1), FSU-602 (2), FSU-168 (2), FSU-39 (2), FSU-39
(2), FSU-37 (2), FSU-598 (6), FSU-53 (1); Liberty Co.: DBM-1233 (ca. 5),
DBM-1234 (3), DBM-1354 (7), DBM-1484 (1), DBM-1527 (ca. 5), DBM-
1528 (ca. 6), DBM-1611 (11), DBM-1657 (28), DBM-1659 (2), UF/FSM-
10111 (7),-10157 (6), -10159 (3), -10166 (3), -10138, -10116, -17784, -17787,
-9141 (4), 9144 (4), -9138 (3), -6837, -6900 (3), -7760 (3), FSU-817 (19),
FSU-235 (3), FSU-424 (3); Marion Co.: UF/FSM-18292 to -18310, S. Christ-
man (78), -3076 (15), -3077 (10), -7133 (6), -17446 to -17463, DBM-1648
(8), DBM-1649 (9), DBM-1650 (10); Okaloosa Co.: DBM-1223 (10), DBM-
1255 (ca. 50), DBM-1562 (2), DBM-1563 (ca. 6), DBM-1567 (1), DBM-1568
(2), DBM-1699 (23), DBM-1716 (17), DBM-1717 (4), DBM-1723 (3), DBM-
1724 (5), DBM-1725 (44), DBM-1726 (8), DBM-1729 (ca. 20), NELSC-
19952 to -19956, NELSC-17175 to -17177, UAHC-68-1193 to 68-1195, FSU-
645 (10); Polk Co.: UF/FSM-7378, -26068 to -26074, -2962 (2), -2963, SRT-
28 (6), SRT-317 (3), SRT-278 (3), SRT-1177 (4); Putnam Co.: UF/FSM-
1079; Santa Rosa Co.: DBM-1666 (8), TU-15839 (3); Taylor Co.: DBM-1651
(2); Volusia Co.: UF/FSM-18317; Wakulla Co.: DBM-1553 (6), DBM-1554
(8), DBM-1570 (9), DBM-1613 (1), DBM-1671 (2), FSU-785 (10), FSU-
767 (8), FSU-36 (6), FSU-33 (29); Walton Co.: DBM-1653 (11), DBM-1665


(1), UAHC-68-646 to 68-675, FSU-87 (3), UAHC-68-774, -68-775; Washington
Co.: DBM-1519 (2), UF/FSM no # (8), AUM-15703 to -15705 (3).
Georgia-Berrien Co.: DBM-1645 (14), USNM-104554 (2), USNM-62097 to
-62102, USNM-62181; Bullock Co.: UF/FSM-1569 (4); Camden Co.: DBM-
1550 (5), UF/FSM-2062 (1), UF/FSM-7813 (1); Charlton Co.: DBM-1551
(1), USNM-129943 to -129946; Chatham Co.: USNM-21379 to -80; Coffee Co.:
DBM-1644 (10); Decatur Co.: UF/FSM-1407-2, -1410 (3); Glynn Co.: USNM-
92160 (1); Grady Co.: DBM-1281 (4), DBM-1565 (1), DBM-1623 (3), DBM-
1672 (9); Liberty Co.: DBM-1545 (ca. 15), USNM-3901 (12), UF/FSM-8072
(1); Irwin Co.: DBM-1626 (15), DBM-1642 (11), UG-1331, -2, -3, -7, UG-
1079; Mitchell Co.: DBM-1674 (12); Thomas Co.: DBM-1634 (16), DBM-
1673 (4), TTRS-12, -13; Ware Co.: AUM-10012 (1), USNM-92247 (1); Wil-
cox Co.: TU-14890 (6); Worth Co.: DBM-1676 (11), DBM-1678 (6), DBM-
1680 (5).
Louisiana-Jefferson Par.: TU-13671 (23); St. Charles Par.: DBM-1688 (11), TU-
18518 (5); St. Tammany Par.: DBM-1691 (2), USNM-8874 (1), USNM-
113252, -3; Tangipahoa Par.: DBM-1692 (4), USNM-115964 (1); Washington
Par.: TU-11832 (17).
Mississippi-Forrest Co.: DBM-1537 (ca. 20); Harrison Co.: USNM-51142 to -51151;
Jones Co.: DBM-1535 (ca. 25); Perry Co.: DBM-1583 (ca. 8); Walthall Co.:
DBM-1533 (1).
North Carolina-Brunswick Co.: DBM-1548 (1).
South Carolina-Jasper Co.: RW-2626 (1); Berkeley Co.: DBM-1546 (72).
Texas-Wood Co.: DBM-1684 (19).

Desmognathus brimleyorum
Arkansas-Clark Co.: DBM-1295 (ca. 15); Combs Co.: USNM-118475 to -118483;
Garland Co.: TU-16801 (4), USNM-22157 holotypee), USNM-22158 to -22169
(paratypes), MCZ-2598 (2 paratypes), USNM-57214 to -57225; Hot Springs
Co.: DBM-1296 (36), TU-16795 (10), TU-16803 (18); Howard Co.: Nash
(1); Mortgomery Co.: DBM-1297 (ca. 20), DBM-1298 (27); Pike Co.: TU-
16794 (5); Polk Co.: DBM-1245 (122), DBM-1246 (55), DBM-1481 (2),
DBM-1707 (ca. 40), DBM-1708 (ca. 150), TU-18299 (7), TU-17993 (36),
TU-18291 to -18297, TU-18287 to -18289, NELSC-7350 to -7352, NELSC-
16445 to -16451, Nash (7), OU-25550 to -25571; Pope Co.: AM-A60759 to
-A60762; Scott Co.: DBM-1479 (9), TU-16793 (1); Yell Co.: DBM-1480 (4).
Oklahoma-LeFlore Co.: DBM-1476 (ca. 15), DBM-1477 (2), DBM-1478 (5),
DBM-1709 (ca. 10), DBM-1710 (ca. 10), DBM-1711 (ca. 300), OU-6601, -3,
-4, OU-6609 to -6617, OU-6874 to -6877, OU-6881, -2, OU-6972, USNM-99409
to -99436; McCurtain Co.: DBM-1475 (1), OU-27233 to -27256, OU-31414 to
-31419; Pushmataha Co.: DBM-1473 (ca. 20).

Desmognathus cf. fuscus
Alabama-Baldwin Co.: DBM-1219 (ca. 60), DBM-1581 (ca. 10), DBM-1582 (1),
DBM-1686 (11), DBM-1693 (55); Barbour Co.: AUM-6434, -5, AUM-13009
to -18, AUM-13101, -2, AUM-16796 to -800; Butler Co.: DBM-1413 (ca. 10),
DBM-1428 (ca. 25), DBM-1575 (10), AUM-5062 to -8, AUM-5727, AUM-9524;
Clarke Co.: DBM-1482 (ca. 10), AUM-11136, -37, AUM-12702 to -25, AUM-
14816 to -21, AUM-15698, -99, AUM-15706 to -10; Clay Co.: DBM-1607 (2);
Cleburne Co.: DBM-1186 (ca. 12); Conecuh Co.: DBM-1232 (ca. 5), DBM-
1414 (5), DBM-1427 (ca. 30), AUM-16784 to -95; Covington Co.: AUM-5010,
-5061, AUM-14735 to -44; Crenshaw Co.: AUM-6459 to -64, AUM-7530 to -3;
Dale Co.: DBM-1189 (ca. 10), AUM-14868 to -76, AUM-15118 to -75, AUM-
15278 to -80, AUM-15295 to -306; Geneva Co.: AUM-77, AUM-245; Henry Co.:

____ ___ __


S DBM-1322 (ca. 10), AUM-7765 to -83, AUM-10013 to -15, AUM-10029 to -31,
AUM-15190, AUM-15236 to -40, AUM-16933 to -35; Houston Co.: AUM-1464,
AUM-15292 to -94, UF/FSM-9316; Monroe Co.: AUM-6943, -6946, -6961,
-6977, -7028, -7196, -7234, -7239, -7248, -7249, -7254 -7257, AUM-15202 to
-12, AUM-18750 to -53; Pike Co.: AUM-15155 to -72; Randolph Co.: DBM-
1184 (ca. 12); Russell Co.: AUM-5372, -5449, -5470, -11469, -15265 to -70,
-15272 to -77.
Arkansas-Cross Co.: DBM-1588 (3), DBM-1589 (ca. 35), DBM-1590 (8), DBM-
1591 (24), UF/FSM-16091 to -16112.
Florida-Calhoun Co.: DBM-1265 (ca. 20), DBM-1661 (11), UF/FSM-11793 to
-17821; Escambia Co.: DBM-1193 (ca. 25), DBM-1218 (ca. 20), DBM-1239
(ca. 50), DBM-1240 (ca. 50), DBM-1241 (53), DBM-1242 (ca. 20), DBM-
1248 (ca. 30), DBM-1258 (1); Gadsden Co.: DBM-1162 (ca. 50), DBM-1171
(ca. 35), DBM-1227 (ca. 10), DBM-1228 (ca. 15), DBM-1230 (ca. 30), DBM-
1231 (ca. 15), DBM-1384 (ca. 30), DBM-1515 (ca. 15), DBM-1538 (1),
DBM-1654 (1), DBM-1706; Leon Co.: DBM-1152 (6), DBM-1160 (ca. 15),
DBM-1168 (ca. 15), DBM-1210 (ca. 20), DBM-1211 (6), DBM-1226 (ca. 40),
DBM-1282.5 (9), DBM-1490 (1); Liberty Co.: DBM-1154 (7), DBM-1161 (2),
DBM-1177 (ca. 20), DBM-1203 (ca. 15),DBM-1207 (27), DBM-1212 (ca. 50),
DBM-1213 (ca. 30), DBM-1214 (ca. 30), DBM-1215 (ca. 20), DBM-1233 (ca.
15), DBM-1234 (3), DBM-1253 (ca. 105), DBM-1264 (50), DBM-69-CA
(ca. 35), DBM-69-CB (ca. 15), DBM-69-CC (ca. 30), DBM-69-CD (ca. 25),
DBM-1334 (ca. 35), DBM-1341 (ca. 20), DBM-1388 (ca. 30), DBM-1527
(ca. 10), DBM-1528 (ca. 6), DBM-1611 (9), DBM-1658 (5); Okaloosa Co.:
DBM-1224 (6), DBM-1257 (5), DBM-1262 (3), DBM-1263 (ca. 70), DBM-
1558 (15), DBM-1606 (1), DBM-1718 (ca. 20), DBM-1727 (ca. 50), DBM-
1728 (6); Santa Rosa Co.: DBM-1220 (ca. 30), DBM-1225 (ca. 20), DBM-
1259 (ca. 50), DBM-1260 (ca. 75), DBM-1261 (ca. 20), DBM-1667 (25),
DBM-1670 (11), DBM-1695 (15), DBM-1698 (22); Walton Co.: DBM-1216
(ca. 15), DBM-1217 (ca. 10), DBM-1222 (ca. 15), DBM-1235 (ca. 20), DBM-
1236 (ca. 20), DBM-1243 (ca. 15), DBM-1715 (26), DBM-1719 (4), DBM-
1720 (3), DBM-1721 (5), DBM-1722 (4); Washington Co.: DBM-1238 (4),
DBM-1594 (4), DBM-1652 (10).
Georgia-Bibb Co.: UF/FSM-1408 (7), -14019 (6) -14020 (2), -14021 (3);
Clarke Co.: UG-204 (72), UG-60 (50); Clay Co.: DBM-1183 (ca. 60), UF/
FSM-16240 to -16259; Cobb Co.: UG-207 (6); Decatur Co.: DBM-1170 (ca.
25), DBM-1180 (ca. 40), DBM-1403 (1); Fulton Co.: no # (15); Harris Co.:
UG-280, UG-887; Pike Co.: UG-1158 (12); Randolph Co.: DBM-1291 (ca. 15);
Twiggs Co.: DBM-1641 (ca. 20); White Co.: UG-154 (2).
Louisiana-Natchitoches Par.: TU-16797 (11); Sabine Par.: TU-13291 (35), TU-
13732 (146), TU-14105 (20); St. Tammany Par.: TU-13672 (111), TU-16441
(71), TU-2642 to -2860 (ca. 200), TU-2861 to -2975 (114) (possibly mixed
with auriculatus); Union Par.: NELSC-23995 to -24003.
Mississippi-Benton Co.: DBM-1592 (18), DBM-1187 (ca. 10); Forrest Co.: DBM-
1536 (ca. 15), DBM-1537 (14); Jones Co.: DBM-1535 (22); Lawrence Co.:
TU-14136 (35); Tishomingo Co.: DBM-1188 (4), TU-14635 (31); Walthall
Co.: DBM-1534 (38).
North Carolina-Bladen Co.: DBM-1547 (73); Durham Co.: DBM-1702 (1), DBM-
1703 (11); Gaston Co.: DBM-1604 (4); Montgomery Co.: DBM-1597 (6),
DBM-1598 (8), DBM-1599 (1), DBM-1600 (12), DBM-1601 (ca. 30); Orange
Co.: DBM-1704 (12).
South Carolina-York Co.: DBM-1605 (4).
New York-Cattaraugus Co.: MCZ-65702 to -12; Erie Co.: MCZ-33133 to -51;
Orange Co.: USNM series; Warren Co.: USNM series.


Desmognathus ochrophaeus

Most of the approximately 5500 specimens examined by Martof and Rose (1963)
are now in my care and were referred to in this study. Also, voluminous material in
the Museum of Comparative Zoology, Harvard University, and in the U. S. National
Museum, Washington, D. C., was examined for this study from the entire range of
this form.


Allen, G. M. 1901. Notes on the reptiles and batrachians of Intervale, New Hamp-
shire. Proc. Boston Soc. Nat. Hist., 29: 63-75.
Baird, S. F. 1849. Revision of the North American tailed batrachians with descrip-
tions of new genera and species. Jour. Acad. Nat. Sci. Philadelphia (n.s.): 281-
Banta, A. M. and W. L. McAtee. 1909. The life history of the cave salamander,
Spelerpes maculicaudas (Cope). Proc. U. S. Nat. Mus., 30: 67-83.
Barbour, R. W. 1950. A new subspecies of the salamander, Desmognathus fuscus.
Copeia, 1950 (4): 277-278.
Barbour, R. W., J. W. Hardin, J. P. Schafer, and M. J. Harvey. 1969. Home range,
movements, and activity of the Dusky Salamander, Desmognathus fuscus.
Copeia, 1969 (2); 293-297.
Bishop, S. C. 1941. The salamanders of New York. N. Y. State Mus. Bull., 324: 1-
1943. Handbook of salamanders. Cornell Univ. Press, Ithaca.
Bleakney, J. S. 1958. A zoogeographical study of the amphibians and reptiles of
eastern Canada. Nat. Mus. of Canada Bull., 155: 1-119.
Boulenger, G. A. 1882. Catalogue of the Batrachia Gradientia s. Caudata and
Batrachia Apoda in the collection of the British Museum. British Museum,
Brame, A. H., Jr. 1967. A list of the world's recent and fossil salamanders. Herpe-
ton, 2(1): 1-26.
Brimley, C. S. 1910. Records of some reptiles and batrachians from the southeastern
United States. Proc. Biol. Soc. Washington, 23: 9-18.
Burger, J. W. 1937. The relation of germ cell degeneration to modifications of the
testicular structure of plethodontid salamanders. J. Morph., 60: 459-487.
Carr, A. F., Jr. 1940. A contribution to the herpetology of Florida. Biol. Sci. Ser.,
Univ. Fla. Publ., 3(1): 1-118.
Carr, A. and C. J. Coin. 1955. Guide to the reptiles, amphibians and fresh-water
fishes of Florida. Univ. Florida Press.
Chaney, A. H. 1949. The life history of Desmognathus fuscus auriculatus. M. S.
thesis, Tulane Univ., New Orleans, La.
-- 1958. A comparison of Louisiana and Arkansas populations of Desmognathus
fuscus. Ph.D. thesis, Tulane Univ., New Orleans, La.
Collette, B. B. and R. W. Yerger. 1962. The American percid fishes of the sub-
genus Villora. Tulane Stud. Zool., 9(4): 213-230.
Conant, R. 1958. A field guide to the reptiles and amphibians of the United States
and Canada east of the 100th Meredian. Houghton-Mifflin, Boston.
Cope, E. D. 1859. On the primary divisions of the Salamandridae, with descriptions
of two new species. Proc. Acad. Nat. Sci. Philadelphia, 11: 122-128.
1869. A review of the species of Plethodontidae and Desmognathidae.
Proc. Acad. Nat. Sci. Philadelphia, 21: 93-118.
1889. The Batrachia of North America. Bull. U. S. Nat. Mus., 34: 1-525.
Dijkgraaf, S. 1962. The functioning and significance of the lateral line organs.
Biol. Rev., 38: 51-105.

_ ___~I __ 1~~__ __ _ ~~

Vol. 18 No. 1


Drastich, L. 1927. Uber das Leben der Salamanderlarvan bei hohem und niedrigein
Sauerstoffpartialdruck. Z. verg. Physiologie, 2: 632-657.
Dunn, E. R. 1916. Two new salamanders of the genus Desmognathus. Proc. Biol.
Soc. Washington, 29: 73-76.
1917. The salamanders of the genera Desmognathus and Leurognathus.
Proc. U. S. Nat. Mus., 53: 393-433.
1926. The salamanders of the family Plethodontidae. Smith College
Fiftieth Anniversary Publ., Northampton, Mass., 441 pp.
Eaton, T. H., Jr. 1956. Larvae of some Appalachian plethodontid salamanders.
Herpetologica, 12(4): 303-311.
Folkerts, George W. 1968. The genus Desmognathus Baird (Amphibia: Plethodon-
tidae) in Alabama. Ph.D. thesis, Auburn Univ., Auburn, Ala.
Fowler, H. W. 1906. Note on the dusky salamander. Proc. Acad. Nat. Sci. Phila-
delphia, 58: 356-357.
Goin, Coleman J. 1951. Notes on the eggs and early larvae of three more Florida
salamanders. Ann. Carnegie Mus., 32(2): 253-263.
Coin, C. J. and D. M. Cochran. 1970. The new field book of reptiles and amphibians.
G. P. Putnam's Sons, N. Y., vi-xxii, 359 pp.
Grobman, A. B. 1950. The distribution of the races of Desmognathus fuscus in the
southern states. Chicago Acad. Sci. Nat. Hist. Misc., 70: 1-8, 2 figs.
Hairston, Nelson G. 1949. Local distribution and ecology of the plethodontid sala-
manders of the southern Appalachians. Ecol. Monog., 19(1): 47-73.
Harima, H. 1969. A survey of the herpetofauna of northwestern Florida. M. S.
thesis, Univ. Alabama, Tuscaloosa, Ala.
Hays, R. M., Jr. 1966. The mental hedonic gland-cluster of the male salamander,
Desmognathus fuscus. Ph.D. thesis, Univ. Cincinnati, Cincinnati, Ohio.
Hendry, C. W., Jr. and C. R. Sproul. 1966. Geology and ground-water resources of
Leon County, Florida. Bull., Fla. State Geol. Survey, 47: i-xii, 178 pp., 1 pl.
Hilton, W. A. 1947. Lateral line sense organs in salamanders. Bull So. Calif. Acad.
Sci., 46(3): 97-110.
1951. Teeth of salamanders. Herpetologica, 7(3): 133-136.
Hinderstein, B. 1969. Studies in the comparative biochemistry and morphology of
the salamander genus Desmognathus (Amphibia: Caudata). Ph.D. Thesis, City
Univ. of New York, N. Y.
.1971. The desmognathine jaw mechanism (Amphibia: Caudata: Pletho-
dontidae). Herpetologica, 27(4): 467-476.
Holbrook, J. E. 1838. North American Herpetology, Ed. 1, Vol. 3: 115-116.
Huheey, J. E. 1966. The desmognathine salamanders of the Great Smoky Mountains
National Park. J. Ohio Herpetol. Soc., 5(3): 63-72.
Humphrey, R. R. 1922. The multiple testis in urodeles. Biol. Bull., 43(1): 45-67.
Kerr, T. 1960. Development and structure of some actinopterygian and urodele
teeth. Proc. Zool. Soc. London, 133: 401-422.
Kingsbury, B. F. 1895a. The lateral line system of sense organs in some American
Amphibia and comparison with the dipnoans. Proc. Amer. Microscop. Soc., 17:
.1895b. The spermatheca and methods of fertilization in some American
newts and salamanders. Proc. Amer. Microscop. Soc., 17: 261-304.
- 1902. The spermatogenesis of Desmognathus fuscus. Amer. J. Anat., 1(2):
Lehman, J. P. 1968. Remarques concernant la phylog6nie des amphibiens. In Orvig,
T. (ed.), Nobel Symposium 4: Current Problems of Lower Vertebrate Phylogeny.
Stockholm, 307-315.
L6nnberg, E. 1894. Notes on reptiles and batrachians collected in Florida in 1892-3.
Proc. U. S. Nat. Mus., 17: 317-339.


Martof, B. S. and F. L. Rose. 1963. Geographic variation in southern populations of
Desmognathus ochrophaeus. Amer. Midi. Nat., 69(2): 376-425.
Marynick, S. P. 1971. Long-term storage of sperm in Desmognathus fuscus from
Louisiana. Copeia, 1971(2): 345-347.
Mayr, E. 1969. Principles of systematic zoology. McGraw-Hill, New York.
Means, D. B. 1971. Dentitional morphology in desmognathine salamanders (Am-
phibia: Plethodontidae). ASB Bull., 18(2): 45. (Abstr.).
S1972. Comments on undivided teeth in urodeles. Copeia, 1972(3): 586-
Means, D. B. and C. J. Longden. 1970. Observations on the occurrence of Desmog-
nathus monticola in Florida. Herpetologica, 26(4): 396-399.
Neill, W. T. 1951. A new subspecies of dusky salamander, genus Desmognathus,
from south-central Florida. Publ. Res. Div. Ross Allen's Reptile Inst., 1(3):
Noble, G. K. 1927. The plethodontid salamanders: some aspects of their evolution.
Amer. Mus. Novit., No. 249: 1-26.
1931. The hedonic glands of the plethodontid salamanders and their rela-
tion to sex hormones. Anat. Record, 48: 57-58.
Noble, G. K. and S. H. Pope. 1929. The modification of the cloaca and teeth of the
adult salamander, Desmognathus, by testicular transplants and by castration.
J. Exp. Biol., 6: 399-411.
Organ, J. A. 1961. Studies of the local distribution, life history, and population
dynamics of the salamander genus Desmognathus in Virginia. Ecol. Monogr.,
31: 189-220.
Parker, H. W. and E. R. Dunn. 1964. Dentitional metamorphosis in the Amphibia.
Copeia, 1964(1): 75-86.
Parsons, T. S. and E. E. Williams. 1962. The teeth of Amphibia and their relation
to amphibian phylogeny. J. Morph., 110: 375-389.
Puri, H. S. and R. O. Vernon. 1964. Summary of the geology of Florida and a guide-
book to the classic exposures. Fla. Geol. Surv. Spec. Publ. 5: i-ix, 312 pp.
Rafinesque, C. S. 1820. Annals of Nature. Vol. 1: 4.
Rossman, D. 1959. Ecosystematic relationships of the salamanders Desmognathus
fuscus auriculatus Holbrook and Desmognathus fuscus carri Neill. Herpetologica,
15(3): 149-155.
Rubenstein, N. 1969. A study of the salamanders of Mt. Cheaha, Clebume County,
Alabama. Jour. Herpetol., 3(1-2): 33-47.
1971. Ontogenetic allometry in the salamander genus Desmognathus.
Amer. Midl. Nat., 85(2): 329-348.
Sanders, D. and H. M. Smith. 1949. Some noteworthy records of amphibians in
Texas. Trans. Kansas Acad. Sci., 52(1): 28-29.
Schmidt, K. P. 1953. A check list of North American amphibians and reptiles.
Amer. Soc. Ichthyologists and Herpetologists, Chicago viii, 280 pp.
Schultze, H. P. 1970. Folded teeth and the monophyletic origin of Tetrapods.
Amer. Mus. Novit., 2408: 1-10.
Sellards, E. H. and H. Gunter. 1918. Geology between the Apalachicola and Och-
lockonee Rivers in Florida. Fla. Geol. Surv., 10th-llth Ann. Rept.: 9-56.
Sharp, H. S. 1938. Steepheads and spring sapping in Florida-Holt and Niceville
quadrangles, Florida. J. Geomorphol., 1: 247-248.
Spight, T. M. 1967. Population structure and biomass production by a stream sala-
mander. Amer. Midl. Nat., 78(2): 437-447.
Stejneger, L. 1895. A new salamander from Arkansas with notes on Ambystoma
annulatum. Proc. U. S. Nat. Mus. (1894), 17: 597-599.
Stejneger, L. and T. Barbour. 1943. A check list of North American amphibians
and reptiles. Fifth ed., Bull. Mus. Comp. Zool. 93(1): v-xix, 1-260.

____ ______ __ _

Vol. 18 No. 1


Tilley, S. G. 1968. Size-fecundity relationships and their evolutionary implications
in five desmognathine salamanders. Evolution, 22: 806-816.
Tilley, S. G. and D. W. Tinkle. 1968. A reinterpretation of the reproductive cycle
and demography of the salamander Desmognathus ochrophaeus. Copeia, 1968(2):
Valentine, B. D. 1961. Variation and distribution of Desmognathus ocoee Nicholls
(Amphibia: Plethodontidae). Copeia, 1961(3): 315-322.
- 1963. The salamander genus Desmognathus in Mississippi. Copeia, 1963(1):
Van Hyning, O. C. 1933. Batrachia and Reptilia of Alachua County, Florida.
Copeia, 1933(1): 3-7.
Vernon, R. 0. 1942. Geology of Holmes and Washington Counties, Florida. Bull. Fla.
Geol. Surv., 21: i-ix, 161 pp., 2 maps.
Wake, D. B. 1966. Comparative osteology and evolution of the lungless salamanders,
family Plethodontidae. Mem. So. Calif. Acad. Sci., 4:1-111.
Wilder, I. W. 1913. The life history of Desmognathus fusca. Biol. Bull., 24(5): 251-


Vol. 18 No. 1

FIGURE 1.-Map of study areas. A) Ouachita Mountain Uplift of Arkan-
sas and Oklahoma, and peripheral portions of Coastal Plain Arkansas and
Louisiana. B) Florida west of the Suwannee River drainage. Dots rep-
resent localities from which specimens were examined during this study.



EIrEER 2.CIown morphology in the subfamily Desmognathinae. A)
Phweognaths hubrichti, B) LMeuognathus marmoratusw C) DesmagmSthus
monticola; D) D. quadramacularts; E) D. oeneus; F)D. wright; G)

FIunRE 3.-Crown morphology of Ouachita Mountain and northern
Louisiana desmognathines. A-D) Desmognathus brimleyorum Stejneger
dentary teeth of 2-lobed male from Rich Mountain, Polk Co., Arkansas;
E-H) Desmognathus cf. fuscus dentary teeth of 2-lobed male from Union
Par., Louisiana. A, E) (20X, 24X) lingual view; B, F) (20X, 24X) labial
view; C, G) (both 20X) occlusal view; D, H) (124X, 120X) labial view.



Fmooo 4. -Detailed occlusal crown morphology. A) Desmognathus of.
fusous from Union Pao.n La (32OX), 2-lobed male; B) D. brimleyoorm
from Polk Co., Ark. (l1OX), 2-lobed male. Both dentary teeth.

Fmonc5,-Jaw profile in sexually mature males of species in the sub-

Co., Ga.; C) Desmogothuso quadroooculatus (DBM 1400), Haywood
Co., N. C.; D) D. monticola (DBM oIoQ- r 1 1.0 Ala.; E) D.
fuscus topotypo (USNM-8O 32), 0) D. oooo
(USNM 147802. 0 .,. .
Co. N. C. H) "'' -
boitoloyorum (DBM 1476), Polk Co., Ark.; J) D. aurioulatus (DBM-
1551), Charlton. i V ,F ,. 1 1 j ., Yancey Co.,
N.C.;L)D.oc I, -. -,I -.. -- - ', Allare 2
lobed males drawn in same potion with camera lucida (scale =lO ma).








4-I -




Vol. 18 No. 1


FIGURE 6.-Variation in jaw profile dimorphism of Florida Desmognathus
cf. fuscus. Skulls from each locality depicted were rated from 1 to 5 in
profile dimorphism as compared to standards (1=D. auriculatus males
from Florida which show no dimorphism; 5 = D. ochrophaeus males from
Swain Co., N. C. which possess profile dimorphism in the extreme).


FIGURE 7.-External jaw profile in adult Florida males. A) Desmogna-
thus cf. fuscus; B) D. auriculatus. Both 2-lobed males from Sweetwater
Creek, Liberty Co., Florida where they are sympatric in the same habitat
(but not in the same microhabitats).


Vol. 18 No. 1

FIGURE 8.-Premaxillary fontanelle morphology. A) Desmognathus
bremleyorum adult female, B) 3-lobed male from Rich Mountain, Ar-
kansas; C,D) Desmognathus cf. fuscus 2-lobed males, E) adult female
from Florida, F.G) D. auriculatus 2-lobed males, H) adult female from









FIGURE 9.-Premaxillary fontanelle morphology. These and the follow-
ing figure show wide variation in bone shape along suture lines, but con-
servatism both in the shape of the skull as a whole and in the degree of
fusion of the posterodorsal projections of the premaxilla which form the
premaxillary fontanelle. Desmognathus auriculatus: A) female; C) 2-
lobed male. D. cf. fuscus: B) female; D) 2-lobed male. All are from

Vol. 18 No. 1


FIGURE 10.-Premaxillary fontanelle morphology. Variation in Desmog-
nathus auriculatus along suture lines of bones of the skull is great, but
size of the premaxillary fontanelle and shape of skull remain constant.
A) 2-lobed male from Marion Co., Fla.; B) 3-lobed male from Bay Co.,
Fla.; C) 3-lobed male from Wakulla Co., Fla.; D) 2-lobed male from
Grady Co., Ga.


A ^ ^ --- ^ fc



FIGURE 11.-Prearticular spine morphology. Mandibular bones of an
adult male Desmognathus brimleyorum Stejneger. A) Occlusal view of
macerated dentary; B) occlusal view of prearticular; C) A and B com-
bined; D-F) lingual, occlusal, and labial views respectively of prearticu-
lar bone illustrating the high dorsally pointing spine. A female is repre-
sented in G, H. Arrow indicates surface of insertion of atlas-mandibular
ligament on dentary (crosshatched).

Vol. 18 No. 1






FIGURE 12.-Prearticular spine morphology. Comparison of Desmogna-
thus brimleyorum with D. cf. fuscus from the adjacent Coastal Plain of
Arkansas and Louisiana. A) D. cf. fuscus male from Union Par., La.; B)
D. cf. fuscus male from Cross Co., Arkansas; C) D. brimleyorum male
from Polk Co., Ark.; D) D. brimleyorum female from Polk Co., Ark.
(Scale= 10 mm).



FLUES 13-Tail morphology. X-ys ofEDesmognathusauriculatus males
on the left, D. cf. fuscs males on the right. ArroNs mark the beginning
of regenerated tail tips.



FIGUM 14-Tai morphology. Xssays of Desmsgnthms, auricultm fe
mHls on thIe lft;D. f.fuu fmals onrgt Arross indicat be-
ginngo tdtltip


FIGURE 15.-Size comparison between equivalent age classes of repre-
sentative populations of Desmognathus auriculatus, D. brimleyorum, and
D. fuscus (Fla.). Squares with numbers mature males, numbers indi-
cate lobes per testis; squares with slash=undissccted juveniles of either
sex (arbitrarily divided equally on diagram); open circles=spent or
juvenile females (ova less than 1.5 mm diameter in ovaries); filled circles
=gravid females (ovarian ova greater than 1.5 mm diameter). Arrows
indicate mean snout-vent length of sexually mature males (2 lobes or
more per testis) and gravid females. D. brimleyorum collected Decem-
ber, 1969 from top of Rich Mountain (2600 ft. elev.), Polk Co., Arkansas;
D. auriculatus collected spring, 1971, 1972 from Ochlockonee River
drainage, Leon Co., Florida; D. cf. fuscus collected spring, 1968 from
ravine in Gadsden Co., Florida.

Vol. 18 No. 1







c 60

0) 30
c 1

O 2
o 000 00 0

N 115 N 81 N: 88
S 00 0000

20 1 4 21 4 5 3 1 i 2
Number of Individu

N= 115 N= 81 N= 88
20 543211234 343211 2345 5432 1 2245

Number of Individuals


S9 d 9


SE **o mm2
o so oe

00 2
S0 [0 0
0 3o B o
N 40 =7 N

cNu ber of hIdividuals
IGURE 16.-Comparison of equient age classes of Desmognathus

0 30 CK

20 N=37 N=63


Number of Individuals

FIGURE 16.-Comparison of equivalent age classes of Desmognathus
auriculatus (A) and D. cf. fuscus (B) collected at the same time from
the same locality. Stars indicate undissectcd juveniles of either sex.
Floodplain and escarpment toe of Sweetwater Creek in See. 21 T2N R7W,
Liberty Co., Florida. (Symbols as in Fig. 15.)

Vol. 18 No. 1



E3 2 22

2 3
2 3 2

2 2 x
2 23221122


2 4 5 X43
2 x
0 xxx 7

Number of Individuals

FIGURE 17.-Size-frequency distribution of Desmognathus fuscus carri
Neill. A representative size-frequency distribution for D. auriculatus
from Ochlockonee River drainage (A) is compared with D. f. carri topo-
types collected from Silver Glen Springs by S. Christman (B) and para-
types (C). Males are designated by squares with numbers signifying
lobes per testis; squares with X indicate undissected juveniles; filled
circles = gravid females; open circles = spent or juvenile females (ovarian
ova less than 1.5 mm diameter); squares with M indicate 1-lobed mature


Vol. 18 No. 1

FIGURE 18.-Development of color pattern in Desmognathus brimleyorum
Stejneger. A) larva; B) juvenile of previous year's larva; C, D) 1+
year old juveniles. Lateral line rows 1-3 become obscured on the trunk
by melanophore invasion after transformation (B-D); row 2 (on the tail)
persists in D. brimleyorum longer than any lateral line row, but is eventu-
ally lost in sexually mature specimens.


if* 0 1


Vol. 18 No. 1

FIGURE 19.-Development of color pattern in Desmognathus auriculatus
from Mississippi. A, B) larvae; C) recent transformling. Note unco-
alesccd, pigmentless areas surrounding neuromast sites of lateral line
rows, especially on dorsum of transformling. Collected 26 March 1971
from floodplain of Leaf River, Forrest Co., Miss. (DBM-1537).





Vol. 18 No. 1

FIGURE 20.-Development of Color Pattern in Desmognathus auriculatus
from Florida. A, B) larvae; C) transformling. Note extensive dark pig-
ment surrounding neuromast sites, and swamped-out pattern in transform-
ling. Collected 28 March 1971 from mucky depression in Ochlockonee
River tributary, Gadsden Co., Fla. (DBM 1538).






FIGURE 21.-Development of color pattern in Desmognathus fuscus from
Florida. A) Larva; note obvious presence of lateral line rows 1 and 3,
but absence of row 2 on trunk. Row 2 is present on tail. B-D) first year
transformlings; B and D illustrate variation in fusion of light dorsal
blotches; C illustrates a faint persistence of row 3 on the trunk and row 2
on the tail (these become obscured by melanophore invasion with in-
creasing age). Collected 16 December 1969 from Apalachicola escarp-
ment ravine in Liberty Co., Fla. (DBM-1253). Scale= 1 mm.

Vol. 18 No. 1




Vol. 18 No. 1

FIGURE 22.-Ontogenctic fate of dorsal pattern in Desmognathus brimley-
orum. A) larva; B-D juveniles; E) small adult female; F) adult male.
Collected 28 November 1969 from top of Rich Mountain, Polk Co., Ark.



oW 1WI ~-~
c t 63f, Ik~ ~~-:




Vol. 18 No. 1

FIGURE 23.-Color pattern in adult Desmognathus auriculatus from Mis-
sissippi. A, B) juveniles; D) adult female; C, E, F) adult males. Note
dorsal pattern of uncoalesced light spots surrounding neuromast sites on
the trunk and tail; also, note presence of faint lines on "neck." Collected
25 March 1971 from ox-bow lake of Leaf River, Jones Co., Miss. (DBM-

c D


Vol. 18 No. 1

FIGURE 24.-Color pattern of adult Desmognatthus auriculatus from Leon
Co., Fla. Note overall dark color and presence of lighter neuromast sites
on B; faint "neck" lines are present on B. Dorsum of tail is lighter me-
dially and has light spots surrounding neuromast vestiges in some. A, B)
juveniles; C) adult male; D-F) adult females. Collected 1 May 1971
from mucky margin of cypress pond in Leon Co., Fla. (DBM-1560).



Vol. 18 No. 1

FIGURE 25.-Color pattern of adult Desmognathus auriculatus from Liber-
ty Co., Georgia. Note overall darkness of topotypes. A) juveniles; B,
C, E) adult females; D) adult male. Collected 13 April 1971 from Rice-
boro, Liberty Co., Ga. (DBM-1545).

A B C D E.


Vol. 18 No. 1

FIGURE 26.-Color pattern of postlarval Desmognathus auriculatus from
ravine in Okaloosa Co., Florida. These specimens have lost a strong wash
of brick red color due to preservation. The red color is evident on the
dorsum on close examination of living specimens; but where it is super-
imposed on "portholes" (neuromast sites of row 3 on trunk and row 2 on
tail) and other less intensely pigmented areas, it shows up brightly.
Collected 17 December 1969 from Rogue Creek head, Okaloosa Co., Fla.

_ ~___



Vol. 18 No. 1

FIGURE 27.-Color pattern of postlarval Desmognathus fuscus from Liber-
ty Co., Florida. Note scalloped-edge pattern on dorsum of juvenile males
and adult female (A, B, D) and effect of melanization on adult males
(E, F). One adult male (C) retains the juvenile pattern of coalesced
light spots. Collected 17 December 1969 from head of Beaverdam Creek,
Liberty Co., Fla. (DBM-1215).

__ ___ __





Vol. 18 No. 1

FIGURE 28.-Color pattern of postlarval Desmognathus fuscus from Wal-
ton Co., Florida. Note the thin lines of melanophore concentration on
dorsa. These appear to result from ontogenetic breakup of dorsal blotch-
es (e.g., in the series A-D-F). Collected 14 November 1969 from ravine
tributary of Bruce Creek, Walton Co., Florida (DBM-1236).





Vol. 18 No. 1

FIGURE 29.-Color pattern of Desmognathus auriculatus and D. fuscus
collected at the same time from the same locality. Top row, D. fuscus;
bottom row, D. auriculatus. Note the close similarity of adult males
(largest specimens) of fuscus to auriculatus at first glance. Closer in-
spection reveals externally visible differences in tail morphology, head
shape, qualitative differences in color. Collected 31 August 1971 from
floodplain of Sweetwater Creek, Liberty Co., Florida (DBM-1611).

_ __ __ _~_ _




F-aTnc 30-Color change in Demsogrothts focus from Liberty Co.,

fi' .
acid hillside seepage site in ravine dissecting Apalachicola escarpment,
Liberty Co., Florida (DBM-1658). Differences in proportions due to Se
fraction of light (B photographed through water).

IF- 3E 1.--Color change in Destoognathus twusn from Baldwin Co.,

identical in color at the time of collection. Collected 6 April 1972 (DBM-
1686) and 10 April (DBM-1693) from mucky ravine draining into east
side of Mobile Bay, Baldwin Co., Alabarna.


I ME V6 ;i.7lL f m N I E r F A r


Vol. 18 No. 1

FIGURE 32.-Geographical range of Desmognathus brimleyorum Stejneger.
Solid circles represent localities from which specimens of D. brimleyorum
were examined for this study. Triangles represent the closest localities
from which other species were examined.



Vol. 18 No. 1

FIGURE 33.-Geographic range of Desmognathus fuscus in Florida drain-
ages. Solid line represents probable boundary of range to the south and



Vol. 18 No. 1

FIGURE 34.-Geographic range of Desmognathus auriculatus in Florida
drainages. Solid line represents probable boundary of range to the north.



University of Florida Home Page
© 2004 - 2010 University of Florida George A. Smathers Libraries.
All rights reserved.

Acceptable Use, Copyright, and Disclaimer Statement
Last updated October 10, 2010 - - mvs