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Group Title: Bulletin Florida state museums
Title: Bulletin of the Florida State Museum ; Volume 10, Number 7
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 Material Information
Title: Bulletin of the Florida State Museum ; Volume 10, Number 7
Series Title: Bulletin of the Florida State Museum
Alternate Title: Variation among the Southeastern crowned snakes, genus tantilla
Physical Description: Serial
Language: English
Creator: Telford, Sam Rountree Jr.
Publisher: Bulletin of the Florida State Museum, University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1966
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Volume ID: VID00001
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Holding Location: University of Florida
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UNIVERSITY

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BULLET


IN


OF THE

FLORIDA STATE MUSEUM


BIOLOGICAL


SCIENCES


Volume 10


Number 7


VARIATION AMONG THE SOUTHEASTERN
CROWNED SNAKES, GENUS TANTILLA


Sam Rountree Telford. Jr.





Of~


UNIVERSITY


OF FLORIDA


Gainesville


L











Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM are pub-
lished at irregular intervals. Volumes contain about 300 pages and are not nec-
essarily completed in any one calendar year.


WALTER AUFFENBERG, Managing Editor
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Consultants for this issue:

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HOBART M. SMITH


Communications concerning purchase or exchange of the
manuscripts should be addressed to the Managing Editor of
State Museum, Seagle Building, Gainesville, Florida. 32601


publication and all
the Bulletin, Florida


Published May 25, 1966


C


Price for this issue $.65













VARIATION AMONG THE SOUTHEASTERN
CROWNED SNAKES, GENUS TANTILLA


SAM ROUNTREE TELFORD, JR.1








SYNOPSIS: The coronata group of crowned snakes in southeastern United States
includes three species: Tantil!a coronata Baird and Girard, 1853, and two spe-
cies herein described, T. relicta and T. oolitica. T. coronata ranges through
out southeastern United States, but does not enter peninsular Florida south of
the Suwannee River; T. oolitica is restricted to Dade and Monroe counties in
southeastern Florida; T. relicta inhabits peninsular Florida.
Variation in characters of pattern, scutellation, body proportions, and habitat
support recognition of three subspecies in T. relicta: T. r. neilli occurs in sand-
hills and mesic hammocks of northeentral Florida; T. r. relicta of central Florida
is restricted to scrub habitat where syntopic with neilli, but also occurs in sand-
hills where allopatric in southeentral Florida; T. r. pamlica inhabits the narrow
strip of coastal dunes and scrub in southeastern Florida. The little evidence of
gene flow among subspecies is suggestive of secondary intergradation.
Two named subspecies of T. coronata, wagneri (Jan) and mitrifer Schwartz,
arc synonymized with T. coronata.
The three southeastern species of Tantilla probably were derived from a com-
mon precursor of the early Pleistocene, diverging during the successive fluctua-
tions of sea level which characterized the glacial-interglacial stages.









1 The author completed most of this study while a arlnt" student at the
University of Florida, Gainesville and the University of .1. ..... Los Angeles,
and presented preliminary conclusions at the annual meetings of the American
Society of Ichthyologists and Herpetologists at Austin, Texas, in April 1961. He
is currently National Institutes of Health Post-doctoral Fellow in the Department
of Parasitology, The Institute for Infections Diseases, University of Tokyo, Japan
Manuscript received 21 June 1965.-ED.



Telford, Sam Rountree, Jr. 1966. Variation among the southeastern crowned
snakes, genus Tantilla. Bull. Florida State Mus., vol. 10, no. 7, pp. 261-304.









262 BULLETIN FLORIDA STATE MUSEUM Vol. 10




TABLE OF CONTENTS
Introduction -.. ..---. -- 262 T. oolitica ....... 281
Acknowledgements and T. coronata ... 285
Abbreviations --------.. 263 Gross Ecology ......... 291
Materials and Methods 264 Size and Sexual Maturity .... 294
Taxonomy .. ...... 264 Discussion .. ... 295
Tantilla relicla. --- 270 Summary 301
T. r. relicta ...-- 278 Key to southeastern Tantilla 302
T. r. neilli ...... 279 Literature cited -..... 303
T. r. pamlica ------- -- 280




INTRODUCrION
Baird and Girard (1853: 131) described Tantilla coronata from a
specimen collected in Kemper County, Mississippi. The holotype,
United States National Museum 1876, serves also as the type of the
genus Tantilla, which Baird and Girard defined at the same time.
In 1862 Jan described Homalocranion wagneri from a specimen col-
lected in "Florida" by a Professor Wagner. Jan and Sordelli (1866:
livr. 15, pl. 2, fig. 3) figure what is apparently the type specimen of
H. wagneri. Garman (1883: 163) treated H. wagneri as Tantilla coro-
nata var. wagneri. Boulenger (1896: 218) referred H. wagneri to
the synonymy of Homalocranium coronatum, adopting the generic
designation of Dum6ril and Bibron (1854). Cope (1900: 1111) men-
tioned briefly that H. wagneri had not been observed by American
naturalists, and since the holotype of wagneri had an entire anal
plate, it could not belong to the genus Tantilla, in which the anal is
divided. In the same work (1900: 1114), he listed H. wagneri as a
synonym of Tantilla coronata. Until Blanchard's review (1938) of
the United States members of the genus, the name wagneri was con-
sidered a synonym of T. coronata. Blanchard resurrected wagneri
for peninsular Florida Tantilla, justifying subspecific relationship to
T. coronata on grounds of overlap in scale characters. Most workers
since 1938 have followed Blanchard's arrangement. Wright and
Wright (1957: 733) commented on the polymorphism of Florida speci-
mens, in which all possible variations of banded and non-banded head
patterns occur in no proved correlation, and speculated that perhaps
there is but one highly variable form of Tantila in southeastern









VARIATION AMONG SOUTHEASTERN SNAKES


United States. In the meantime Schwartz (1953) designated montane
populations of Tantilla in Tennessee and North and South Carolina
as T. coronata mitrifer.
In 1959 I was impressed by the fact that all Tantilla I collected
from rosemary scrub habitat (Carr, 1940; Laessle, 1958; Telford, 1959)
in southcentral Florida had a prominent light band across the parietal
area, separating the black head cap from the black nuchal collar,
while all that I collected from sandhills (Laessle, 1958) or mesic ham-
mock (Carr, 1940) habitats in northcentral Florida had a predom-
inantly black head and nuchal collar with no distinct light parietal
band. This prompted me to review relationships of southeastern
Tantilla in the light of additional material accumulated since Blanch-
ard's 1938 review. In the last 5 years I have examined 215 Tantilla
coronata from sites throughout the range exclusive of peninsular
Florida, and 166 specimens from the peninsula, of populations pre-
viously called Tantilla coronata wagneri.
Useful taxonomic characters in scutellation are restricted to ven-
trals and subcaudals. Examination of 83 maxillae did not prove
particularly fruitful in clarifying the status of the populations. But
by comparing structure of the hemipenis with pattern, size, head
shape, scutellation, body proportions, and ecology, I am able to in-
terpret the polymorphism of peninsular populations.

ACKNOWLEGEMENTS AND ABBREVIATIONS

For the loan of specimens I should like to thank: W. J. Riemer,
University of Florida Collections (UF); B. Martof, University of
Georgia (UG); E. M. Burton, Charleston Museum (CHM); D. M.
Cochran, United States National Museum (USNM); C. M. Bogert,
American Museum of Natural History (AMNH); E. Williams, Museum
of Comparative Zoology (MCZ); N. D. Richmond, Carnegie Museum
(CM); R. F. Inger and H. Marx, Chicago Natural History Museum
(CNHM); the late N. Hartweg, University of Michigan Museum of
Zoology (UMMZ); H. M. Smith, University of Illinois (UI); W. E.
Duellman, University of Kansas (UK); H. Dundee, Tulane University
(TU); and O. T. Owre, University of Miami (UM); K. L. Hansen,
Stetson University; C. D. Wilder, Memphis State University; E. L.
Modder, Frostproof High School, Frostproof, Florida; and R. L.
Pyke and B. Mansell, Jacksonville, Florida. For the contributions of
specimens I would like to thank D. R. Paulson, R. H. Mount, W. T.
Neill, and A. Beckenbach. The drawings were prepared by M.
Fukushima and E. C. Bovee, to whom I am most grateful. Finally,









BULLETIN FLORIDA STATE MUSEUM


I should like to thank my colleagues F. W. King, D. A. Rossman,
C. W. Myers, J. McCrone, A. Arata, H. W. Campbell, M. J. Fou-
quette, Jr., W. T. Neill, and especially R. H. Mount for many stimu-
lating discussions and much useful advice.

MATERIALS AND METHODS
All scale counts were made under a dissecting microscope as
follows: ventrals as proposed by Dowling (1951); subcaudals from the
tail tip to the vent, excluding the terminal scale; dorsals from the
left diagonally across the body at the level of the first ventral; at one
head-length posterior to the head; at approximately midbody; and
at one head-length anterior to the vent. Head patterns were recorded
by shading the appropriate portions of a dorsal and left lateral outline
drawing of a Tantilla head. Color descriptions of living specimens
were based upon Maerz and Paul (1950). All specimens were meas-
ured to the nearest half millimeter with a clear plastic ruler. Left
maxillae were removed, cleaned gently with a needle, and dried
under a light bulb before examination. In every case where extruded
hemipenes were not in evidence, sex was determined by examining
the reproductive tract. Sexual maturity of males was established on
the basis of enlarged, contorted, or turgid vasa deferentia in larger
specimens, and by positive sperm smears in the case of smaller in-
dividuals. The presence of ovarian follicles of varying size or of
oviductal eggs was considered indicative of sexual maturity in fe-
males, in contrast to uniformly small follicles in immature individuals.

TAXONOMY
The characters found to be of taxonomic importance among the
species of Tantilla in the southeastern United States are: (1) ventrals,
(2) subcaudals, (3) tail to total length ratio, (4) degree of develop-
ment of the light parietal band, (5) width of the black nuchal collar
posterior to the parietal area, and (6) number of basal hooks of the
hemipenis. Other characters recorded that either do not vary or do
so without demonstrable geographic correlation are the following:
(7) number of dorsal scale rows, (8) preoculars, (9) postoculars, (10)
temporals, (11) supralabials, (12) infralabials, (13) number of supra
labials entering the orbit, (14) number of infralabials contacting the
anterior chin shields, (15) condition of the anal plate, (16) contact be-
tween the mental and anterior chin shields, (17) number of maxillary
teeth, (18) length of retracted hemipenis, (19) degree of pigmentation
of the postoculars and ultimate supralabial, (20) lateral extent of the


Vol. 10









VARIATION AMONG SOUTHEASTERN SNAKES


black nuchal collar, and (21) initial and terminal vertebral scales con-
tacted by the black nuchal collar. Characters 1 to 21 were recorded
for all populations.

THE PENINSULAR POPULATIONS.
Populations of Tantilla from peninsular Florida are presently des-
ignated as Tantilla coronata wagneri (Jan).
The holotype of Homalocranion wagneri was originally located at
Munich, according to Jan and Sordelli (1866). Dr. Walter Hellmich
indicated (in litt.) that it is not there and was perhaps destroyed dur-
ing World War II. Dr. M. Torchio of the Museo Civico di Storia
Natural, Milan, which had many of Jan's types, has informed me
that the Museum was destroyed during World War II and that if the
type was there rather than at Munich, it has not survived. In lieu
of examining the holotype, it is necessary to rely upon the descrip-
tion, and the fortunate illustration of the presumed type of Jan and
Sordelli. I have compared Blanchard's translation of Jan's descrip-
tion with the original and find no discrepancies. Consequently refer-
ences to the type description refer to Blanchard's translation.
The holotype had "Behind two or three gular scutes . 138
ventrals, an entire anal, and 45 double caudals." Disregarding the
anal for the moment, when 138 ventrals are plotted against 45 cau-
dals in scatter diagrams as in figure 1, it is obvious that the specimen
lies well outside the range of variation of peninsular populations (with
the possible exception of the Dade County population), and well with-
in the distribution of Tantilla coronata. As sex of the type specimen
is not given, it is plotted on scatter diagrams of both sexes. On this
basis I conclude that Jan had at hand a specimen of Tantilla coronata
from a non-peninsular population. Though the type locality is given
as "Florida", it should be pointed out that Florida once extended from
the Atlantic Ocean to the Mississippi River along the Gulf Coast.
Even today those Louisiana parishes east of the Mississippi are called
the "Florida parishes." As Jan named the specimen in 1862, it was
probably not collected much later than 1859. By that time very few
American naturalists had collected in peninsular Florida, and I doubt
seriously that Professor Wagner numbered among those who did.
Tantilla coronata is common in the area between the Appalachicola
and Mississippi rivers, and I suspect that Jan's specimen was from this
area. Nothing in the type description settles the issue. Presence
of a single anal is noteworthy, however, for I found but two speci-
mens among the 381 Tantilla I examined that had an undivided anal.


1966














67 le
Males



6q-


.. ,
*.

** **
** * *
"


A Tanllll. coroneta
Peninfsulr Populalions
A
o B
v 0 Se.,se,. K.y
AxB hybrids
C
V7
T.coronot. woagner
holotype


o o
0 *

o Ae A



SAA AA

A

A A


S120 ventrals
110 120 ventralls


0o io
140 150


Females


0
*o * *



0* 60 0 *


It 'L A 'L
A AV A
AA A



A A



A A


1to ventrals


Fig. 1. Scatter diagrams plotting ventrals against subcaudals. Enlarged
symbols represent population means, except where otherwise indicated.









VARIATION AMONG SOUTHEASTERN SNAKES


The illustration in Jan and Sordelli of H. wagneri is that of a typical
Tantilla coronata with a prominent parietal band. On the basis of
the evidence cited, I place the name Homalocranion wagneri Jan,
1862 in synonymy of Tantilla coronata Baird and Girard, 1853.
While Blanchard was correct in stating that overlap in scale
characters between peninsular Tantilla and T. coronata exists, this
overlap is evident only when ventrals or subcaudals are considered
separately. If viewed as a combination of characters, as in the scat-
ter diagrams (figure 1), specimens from the peninsula, except for
those from Dade and Monroe counties on the extreme southern
periphery of the range, show no appreciable overlap with T. coro-
nata: 4.1 percent of males and 6.1 percent of females from peninsular
populations lie within the distribution of T. coronata on the scatter
diagrams. Conversely, only 1 percent of males and 2.6 percent of
females of T. coronata lie within the distribution of peninsular Tan-
lilla in figure 1, again with the exception of Dade and Monroe Coun-
ties. I interpreted this to mean that no intergradation exists between
T. coronata and peninsular populations of Tantilla, which the distri-
bution map (figure 2) substantiates. No specimens of Tantilla are
available from a broad area between the Appalachicola and Suwan-
nee rivers extending north to Chattahoochee and Irwin Counties,
Georgia. Though the hiatus may be real, it may also reflect lack of
collecting, for apparently suitable habitat occurs sporadically through-
out this area.
The scatter diagrams also reveal substantial scutcllation differ-
ences between Dade County Tantilla and other peninsular popula-
tions, and affinity between Dade County Tantilla and T. coronata.
A hiatus of approximately 50 miles exists between the northernmost
locality for Dade County specimens at Miami and the southernmost
site for Tantilla at Boynton Beach, Palm Beach County. There is no
evidence of intergradation between these two populations.
Hemipenial characters of all the main populations of southeastern
Tantilla lend support to the conclusions drawn from characters of
escalation. The hemipenis of Tantilla (figure 3) is single, bearing a
simple sulcus spermaticus, and ornamented with undifferentiated
spines. In Tantilla coronata the basal half is naked, the distal half
spinose, with spine size decreasing toward the apex, which is covered
with small spines. There are two basal hooks, a larger postero-lateral
hook adjacent to the sulcus in the basal third, and a slightly smaller
antero-medial hook in the mesal third of the organ. The hooks are
well differentiated from the larger spines, being a good third longer
and half wider than the basal spines in the apical half. Although


1966










BULLETIN FLORIDA STATE MUSEUM


the spines show minor differences in extent and size, the hemipenes
of peninsular populations from the Suwannee River south through
Palm Beach County generally resemble that of T. coronata-with
one important difference: only the large postero-lateral basal hook,
adjacent to the sulcus, is present. The antero-medial hook has ap-
parently been lost. Two basal hooks at approximately the same size
are present in the Dade County population.


- Lake Wales Ridge


Fig. 2. Distribution of Tantilla in relation to availability of scrub and sand-
hills habitat in southeastern United States. Triangles T. corncata; circles main
peninsular population; squares Dade County population.


Vol. 10























a b














c d

Fig. 3. Hemipenes of southeastern Tantilla: a, T. coronata; b, peninsular
population A; c, peninsular population D; d, peninsular population C.









BULLETIN FLORIDA STATE MUSEUM


Other differences between these three populations exist, and will
be discussed below, along with variation within the divisions. I be-
lieve that the characters discussed so far, in conjunction with the real
or apparent hiatuses in distribution, justify division of the southeast-
ern Tantilla into three distinct species. Tantilla coronala extends
from Florida north and west of the Suwannee River to Virginia, the
Mississippi River, and extreme southern Indiana. A second species
extends from the Suwannee River south in Florida to the Peace
River on the west coast, and to Broward County along the east coast.
The third is restricted to Dade and northern Monroe counties in
extreme southern Florida.
In recognition of its probable origin during fluctuations of Ple-
istocene sea levels, I designate the populations of Tantilla within
peninsular Florida south of the Suwannee River and north of Dade
County and the outlet of the Peace River into the Gulf of Mexico as

Tantilla relicta new species
HOLOTYPE: UF 12421, an adult female collected 26 December
1960 on the south side of Babson Park, Polk County, Florida by
Sam R. Telford, Jr.
PARATYPES: Polk County: UF 12429-1&2, 12420, 12419; Palm
Beach County: UF 12430-1&2; Alachua County: UF 12406, 7973-
1&2, 3586.
DIAGNOSIS: A small Tantilla, characterized by the presence of a
single basal hook on the hemipenis, postero-lateral in position and
adjacent to the sulcus spermaticus in the basal third of the organ.
The species is composed of several populations varying somewhat
among themselves in details of pattern and scutellation, divisible into
three distinct subspecies. Ventrals range between 115 and 142; sub-
caudals from 39 to 67. Males have fewer ventrals and more subcau-
dais than females, and are slightly smaller. Although constant with-
in subspecies, head pattern varies from completely black with no
light parietal band, to a prominent parietal band separating the
black head cap from the black nuchal collar, with extensive unpig-
mented areas extending forward along outer edges of parietals. Ratio
of tail to total length varies between 18.5 and 29.1 percent, males
having longer tails on the average. Maxillary teeth 14 to 16, usually
15. Head pointed in outline with countersunk lower jaw, to nar-
rowly rounded with noncountersunk lower jaw, varying somewhat
among populations.
DESCRIPTION OF HOLOTYPE: Adult female, 136.5 mm snout-vent


Vol. 10









VARIATION AMONG SOUTHEASTERN SNAKES


and 34.5 mm tail length. Ventrals 131; subcaudals 50; dorsal scales
smooth, in 15 rows throughout; on each side are 1 preocular, 2 post-
oculars, 1 anterior and 1 posterior temporal, 7 supralabials, and 6
infralabials. Supralabials III and IV enter the orbit; infralabials
I to IV contact the anterior chin shields; mental plate in contact with
anterior chin shields; posterior chin shields are two-thirds length of
anterior; rostal slightly scarred; anal divided. In preservative the
head dorsum is black. A light parietal band, about one and one-half
dorsal scales in width, extends from the rear fourth of parietals across
the distal half of the secondary temporals; the light band is broken
by a dark streak extending along the mesal edges of the parietals
and the parietal suture to unite with the black first vertebral scale.
The black nuchal collar is three and one-half vertebral scales in width
and extends laterally on to scale row 2; supralabials I to IV and VI,
the loreal region, and superior postocular are black; the distal fourth
of inferior postocular, anterior fourth of primary temporal, and dor-
sal edge of supralabial V lack pigment; the distal half of supralabial
VII lacks pigment, the light area merging with the parietal band.
The mental, infralabials I to V, and adjacent edges of the anterior
chin shields are suffused with black; infralabial VI has dark flecking.
The dorsum is light tan, and the venter tannish white. In life, the
dorsum was light tan (army brown, 6-A-10) throughout, becoming
lighter laterally to pinkish tan (rosestone, 6-C-9). Tail venter pink-
ish gray (cobwebs, 5-B-7), becoming white along body venter. Parie-
tal band tan (French beige, 13-A-7); head dorsum dark brown (sepia,
8-A-10); nuchal collar black.

VARIATION IN TAXONOMIC CHARACTERS.
The correlation and distribution of variation within Tantilla relicta
show the total sample to be composed of several distinct groups which
should be discussed individually. The three most distinct groups of
specimens are treated below as subspecies, although no strong evi-
dence is available to establish continuity of gene flow among them.
For the moment they are designated populations A, B, and C.
Population A extends from the Suwannee River south to the Hills-
borough River along the west coast of Florida, and in the central por-
tion of the state, west of the St. Johns River to northern Polk County.
It is apparently found only in sandhills and mesic hammock habitats.
Population B occurs in disjunct demes from the vicinity of Lake
George south along the central ridge to southern Polk County (and
presumably Highlands, as well), and in insular type situations along











TABLE 1. VARIATION OF VENTRALS AND SUBCAUDALS AMONG SOUTHEASTERN Tantilla POPULATIONS.


Ventrals Subcaudals

N range mean S' N range mean S2

T. coronate
males 109 123-140 132.5 10.8 97 34-53 46.7 9.5
females 86 133-147 140.0 10.2 77 35-50 43.1 9.0
T. oolitica
males 6 137-143 138.3 5.4 4 51-63 55.3 28.3
females 5 135-146 140.2 19.8 3 45-48 46.3 2.5

T. relicta

Pop. A males 50 123-135 127.5 6.9 45 51-67 59.1 10.7
females 28 129-142 134.2 9.6 20 46-60 53.3 13.1

Pop. B males 8 117-131 123.6 22.9 8 44-59 50.8 26.1
females 14 120-134 128.0 19.2 10 40-55 48.1 22.3

Pop. C males 7 115-118 116.3 2.8 5 45-51 47.8 7.8
females 10 119-129 123.2 8.0 5 45-51 48.2 7.3

Pop. D males 8 121-134 126.8 18.7 7 48-52 50.6 2.0
females 11 123-139 133.2 40.4 8 45-54 50.3 10.1

Seahorse
males 6 119-125 122.5 4.6 6 45-55 50.2 13.8
females 5 128-134 130.6 5.8 4 50-51 50.5 0.3









VARIATION AMONG SOUTHEASTERN SNAKES


the west coast from Seahorse Key in Levy County to Charlotte Coun-
ty north of the mouth of the Peace River. In areas of sympatry with
population A along the northern ridge, B is restricted to scrub habi-
tat, but in allopatric areas of the southern ridge, it also occurs in
sandhills.
Population C is found in coastal dunes and scrub from central
Brevard County to southern Palm Beach County. The southern-
most record is some 50 miles north of the distinctive Dade County
population.
Population D is comprised of 19 specimens that cannot be as-
signed satisfactorily to either A or B, although they were collected
from scattered localities within and on the periphery of the ranges
of both A and B. These specimens show an overlap of characters
from both A and B and may represent intergrades. They are the prin-
cipal reason for considering A and B conspecific. The Seahorse Key
population of T. relicta resembles population B in characters of scutel-
lation and body proportions, but differs somewhat in patterns; it is
therefore treated separately in the tables and figures. Variation in
ventrals and subcaudals is presented in Table 1.
The Dade County population evidently has the highest number
of ventrals among these populations, while the T. relicta population
closest to it geographically, C, has the lowest. T. coronata has the
fewest subcaudals, and the T. relicta population geographically
closest, A, has the highest number. It is tempting to speculate that
these are examples of character displacement.
Variation in proportionate tail length is considerably less than in
scale characters. Table 2 presents the ratio of tail length to total
length among the various samples, while figure 4 depicts regression
lines for this character, calculated by the method of Sncdecor (1946:
103).
In correlation with subcaudal number, population A again has
the proportionately longest tail, while T. coronata has the shortest.
As the sexes showed no significant differences in color pattern,
the data were lumped for analysis. Sorting specimens according
to the distribution of pigment in the parietal and temporal regions
permitted dividing them into ten different categories (figure 5) rang-
ing from specimens with a completely black head cap which fused
with the nuchal collar (category 1) to specimens in which only the
anterior, central portion of the parietal was pigmented (category 10).
This procedure provided a simple method of quantifying the pat-
tern data (table 8).


I


















TABLE 2. TAIL LENGTH IN SOUTHEASTERN Tantilla EXPRESSED


AS PERCENT OF TOTAL LENGTH.


Males

N range mean S2

94 17.2-23.0 20.6 1.4

4 20.8-22.4 21.9 0.6


21.7-28.7

20.9-23.5

21.7-24.0

20.7-23.8

21.5-25.2


Females


7. coronata

T. oolitica

T. relicta

Pop. A

Pop. B

Pop. C

Pop. D

Seahorse


range

15.9-21.2

19.1-19.6



19.3-29.1

18.5-23.1

18.7-23.2

19.0-22.7

20.6-21.6


S2

5.4

0.1



4,6

2.1

2.8

1.9

1.0










VARIATION AMONG SOUTHEASTERN SNAKES


Moles


Females


snout-vent length (m


m) 192


Fig. 4. Regression lines depicting the relationship of tail length to snout-
vent length: a, peninsular population A; b, peninsular population B; c, T. coro-
nata; d, peninsular population D.


1966






BULLETIN FLORIDA STATE MUSEUM


I I


I T


a b c d






e f g h






i j
Fig. 5. Variation in parietal pattern among southeastern Tantilla popula-
tions. Categories a-d are non-banded, e-j banded.


Vol. 10









VARIATION AMONG SOUTHEASTERN SNAKES


TABLE 3. PARIETAL PATTERN* IN SOUTHEASTERN Tantilla POPULATIONS,
EXPRESSED AS PERCENT OF SAMPLE

Pattern Types

N a b e d e f g h i ] mean

T. coronata 117 3 16 37 23 9 9 3 6.6
T. oolitica 10 80 10 10 1.7
T. relicta
Pop. A 77 20 27 39 14 2.5
Pop. B 23 14 56 22 4 4 5.4
Pop. C 18 33 6 22 22 17 7.5
Pop. D 19 16 42 32 5 5 3.4
Seahorse 11 9 18 9 36 18 9 3.7

SSee Fig. 5.

Somewhat arbitrarily, the dividing line between banded and non-
banded patterns was set between categories 4 and 5, with pattern
types 5 and above considered banded. According to this classifi-
cation population A is clearly 100 percent nonbanded, while 87
percent of B, 100 percent of C, and 96.6 percent of T. coronata are
banded. The small sample from Dade County (9) are all non-banded;
a single specimen from Key Largo has a broken band. Of the 11
snakes in the small Seahorse Key sample, 8 are nonbanded, as are
2 of 4 T. relicta from Charlotte County. The series of possibly inte
gradient T. relicta, D, whose body proportions and scutellation char-
acters are perhaps closer to population B, are 89.5 percent non-
banded.
Another pattern character of taxonomic use is the width of the
black nuchal collar at midline. The data are readily quantified by
expressing the width in terms of number of vertebral scales pig-
mented (table 4).
The banded populations, T. coronata, B, and C, have nuchal col-
lars predominantly four scales or less in width, the percentages of
the respective samples being 99.5, 91.5, and 67.0. Of the nonbanded
populations, A and Dade County, 66 percent of A have collars five
scales or more in width, while 4 of the 10 Dade County specimens
have collars of this width. Eight of the 11 Seahorse Key Tantilla
relicta and 52 percent of the possibly intergradient population, D,
have collars five scales or more in width. Again, with respect to
both pattern components discussed, the greatest contrast is between
T. coronata and the T. relicta population closest to it geographically,
population A.









BULLETIN FLORIDA STATE MUSEUM


TABLE 4. WIDTH OF NUCHAL COLLAR IN SOUTHEASTERN Tantilla POPULATIONS,
EXPRESSED AS PERCENT OF EACH SAMPLE

Width in Vertebral Scales

N 1 2 3 4 5 6 7 8 9 mean

T. coronala 183 13 63 23.5 0.5 3.0
T. oolitica 10 10 20 30 20 20 4.2
T. relicta
Pop. A 76 16 18 46 9 8 3 4.8
Pop. B. 23 4 17 26 44 9 3.4
Pop. C 18 11 39 17 33 3.7
Pop. D 19 11 5 16 16 47 5 4.1
Seahorse 11 18 10 27 27 18 5.8


In summary of variation in the five characters discussed above,
it is evident that the three main populations of T. relicta are better
distinguished among themselves, as well as from T. coronata and the
Dade County population, by parietal pattern and width of the nuchal
collar than by characters of scutellation and body proportion. These
latter, however, are useful in some instances, i.e., the longer tail and
greater number of subcaudals of population A in comparison to other
populations, and the lower number of ventrals in an admittedly in-
adequate sample of population C.
I believe that sufficient grounds exist to designate populations
A, B, and C as subspecies of Tantilla relicta, representing geograph-
ically and ecologically delimited populations that possess within
themselves common and distinctive morphological features.
In accordance with this concept of the subspecies, I designate
population B

Tantilla relicta relicta new subspecies
HOLOTYPE: UF 12421, an adult female collected 26 December
1960 on the south side of Babson Park, Polk County, Florida by Sam
R. Telford, Jr.
PARATYPES: UF 12429-1&2, 12420, 12419, 12418, 12424; USNM
23429, 23430; CNHM 29592, 29595.
DIAGNOSIS: A population of Tantilla relicta composed of many
more or less isolated demes, restricted in northcentral Florida to scrub
habitat, but occurring also in sandhills of southcentral Florida,
where syntopic to other subspecies of T. relicta. Ventrals range
from 117 to 134; subcaudals between 40 and 59. Tail length varies


Vol. 10









VARIATION AMONG SOUTHEASTERN SNAKES


from 18.5 to 23.5 percent of total length. Parietal pattern is predom-
inantly banded (87 percent). The nuchal collar is usually (43.5 per-
,cent) 4 scales wide at midline, but ranges from 1 to 5 (mean, 3.4
scales). The head is pointed in outline, with countersunk lower jaw.
GEOGRAPHICAL RANGE: Peninsular Florida from the vicinity of
Lake George, Marion County, south along the central ridge to south-
ern Polk County. Presumably this subspecies occurs in Highlands
County as well. Disjunct populations occur in coastal scrubs of
Charlotte, Sarasota, and Pinellas counties, and on Seahorse Key,
Levy County.
DESCRIPTION OF HOLOTYPE: The holotype is described above as
holotype of Tantilla relicta sp. nov.
In recognition of Wilfred T. Neill's contributions to the biography
of Florida, I take pleasure in designating the nonbanded population
of Tantilla relicta in northcentral Florida (A) as

Tantilla relicta neilli new subspecies
HOLOTYPE: UF 12406, an adult male collected 12 July 1961 on
the University of Florida campus, Gainesville, Alachua County, Flor-
ida, by John Funk.
PARATYPES: UF 7973-1&2, 3586, 12404, 8930, 1047, 9586; CNHM
8558; UMMZ 44967; MCZ 43130.
DIAGNOSIS: A population of Tantilla relicta relatively continuous
in distribution in habitats of sandhills and mesic hammock of north-
central Florida. Ventrals range from 123 to 142; subcaudals between
46 and 67. Tail length varies from 19.3 to 29.1 percent of total length,
and is proportionately longer in this subspecies than in any other
population of Tantilla in southeastern United States. The parietal
pattern is nonbanded in all specimens examined, with only 15 per-
cent having unpigmented areas on the parietals larger than a dorsal
scale in size. The black nuchal collar, which fuses with the black
parietal area, is usually (46 percent) 5 scales wide at midline, but
varies from 3 to 8 (mean, 4.8 scales). The head is less sharply pointed
in outline than that of T. relicta relicta, and the lower jaw is less
noticeably countersunk.
GEOGRAPHICAL RANGE: Peninsular Florida from the Suwannee
River south to the Hillsborough River and northern Polk County, and
east to the St. Johns River.
DESCRIPTION OF HOLOTYPE: Adult male, 168.0 mm snout-vent
and 52.0 umm tail length. Ventrals 131; subcaudals 60; dorsal scales
smooth, in 15 rows throughout; on each side are one preocular, two


1966








BULLETIN FLORIDA STATE MUSEUM


postoculars, one anterior and one posterior temporal, seven suprala-
bials, and six infralabials. Supralahials III and IV enter the orbit;
infralabials I to IV contact anterior chin shields; mental plate sep-
arated from anterior chin shields by infralabials I; posterior chin
shields are two-thirds length of the anterior; anal plate divided. In
preservative the head dorsum is black with no light parietal band;
an area the size of one dorsal scale on the posterior tip of the parie-
tals lacks pigment. The black nuchal collar, fused with the black
of the parietals, extends onto the anterior tip of vertebral scale IV,
and laterally onto scale row 2 of each side. The mental, infralabials
I to V, and lateral half of the anterior chin shields are heavily suf-
fused with black; infralabial VI and the gular scales have some black
flecking. The entire loreal and postocular regions and supralabials
I to VI are black; supralabial VII is black except for a small light
area on the rear ventral third. The dorsum is dark tan, venter white.
In life the dorsum was dark tan (argus brown, 7-A-12) becoming
lighter (Natal brown, 7-A-10) on the sides. The tail venter was
yellowish cream (amber white, 11-C-1), and the body venter pinkish
gray (greystone, 12-A-2). The head dorsum and gulars were black,
with the gulars a slightly lighter shade. The parietal spots were
dark tan (Vandyke brown, 7-A-ll).
The third population of T. relicta, C, inhabits coastal dunes and
scrub along a narrow strip of Pamlico terrace in southeastern Florida.
In reference to the geological origin of its habitat, I designate this
population
Tantilla relicta pamlica new subspecies
HOLOTYPE: UF 12430-2, an adult female collected 4 February
1960 approximately 1 mile south of Boynton Beach, Palm Beach
County, Florida, by Robert H. Mount.
PARAiYPE : UF 12430-1, 12423; MCZ 19146, 16272, 16273, 12802;
UMMZ 55779-1&2, 84193.
DIAGNOSIS: A population of Tantilla relicta occurring in relatively
isolated coastal dunes and scrub of southeastern Florida. Ventrals
range from 115 to 129 and are fewer in this subspecies than in any
other southeastern Tantilla population. Subcaudals range between 45
and 51, and tail length varies from 20.7 to 24.0 percent of total length
The parietal pattern is banded in all specimens examined; over 60
percent show a trend toward total absence of pigment on the parie-
tal, supraocular, and temporal regions, a reduction in pigmentation
found in less than 10 percent of T. r. relicta. The nuchal collar is
usually (40 percent) 3 scales in width, and ranges from 2 to 5 (mean,


Vol. 10









VARIATION AMONG SOUTHEASTERN SNAKES


3.7 scales). The head is prominently pointed in outline, with counter-
sunk lower jaw. In life the dorsum is reddish brown, a color seldom
observed in T. r. relicta.
GEOGRAPHICAL RANGE: The east coast of Florida from the vicin-
ity of Cape Canaveral south to southern Palm Beach County, and
limited to the west by the Kissimmee River valley.
DESCRIPTION OF HOLOTYPE: Adult female, 166.0 mm snout-vent
length, with an incomplete tail 31.0 mm long. Ventrals 120; subcau-
dals, incomplete) 30; scales smooth, in 15 rows throughout; on each
side are one preocular, two postoculars, one anterior and one posterior
temporal, seven supralabials, and six infralabials. Supralabials III
and IV enter the orbit; infralabials I to IV contact the anterior chin
shields; mental plate is in contact with the anterior chin shields,
which are one-third longer than the posterior chin shields; anal plate
divided. In preservative the frontal, supraoculars, and anterior two-
thirds of the parietals are black; the unpigmented posterior third
forms a parietal band, which is narrowed slightly at midline by the
black vertebral scale I; the nuchal collar is black and extends onto
the anterior edge of vertebral scale V, and laterally on both sides
onto scale row 2. The medial anterior corners of the secondary tem-
poral are black, the remainder unpigmented. The loreal region, su-
pralabials I to IV, and superior postocular are black; pigment is lack-
ing from the rostral, three-fourths of the internasals, and nasal half
of the prefrontals, anterior edge of the frontal, antero-ventral two-
thirds of the primary temporals, distal edge of the inferior postocu-
lar, supralabials V to VII, and infralabials V and VI. Infralabials
I to IV have a light suffusion or flecking of black. The dorsum is
light pinkish tan, and the venter pinkish white. In life the dorsum
was light reddish brown (cedarwood, 6-G-10), becoming lighter some-
what (plantation, 6-D-11) in the anal region. Sides were pinkish tan
(roseglow, 5-D-9). The tail venter was pinkish gray (cobweb, 5-B-7),
becoming lighter (iris mauve 3-B-7) on the body venter. The parietal
band was a prominent tan (suntan, 13-B-7), while the head dorsum
was dark brown (bracken, 8-C-12), and the nuchal collar black.
I designate the distinctive species of Tantilla restricted to Dade
and Monroe counties, Florida, after its association with the dominant
geological formation of the area, the Miami oolite, as

Tantilla oolitica new species
HOLOTYPE: UF 17326, an adult male collected in April 1955 in
Miami, Dade County, Florida in a vacant lot on southwest 27th


1966








BULLETIN FLORIDA STATE MUSEUM


Avenue near 24th Street. Collector unknown, received through the
courtesy of Dennis R. Paulson.
PARATYPEs: Dade County, UF 17325, MCZ 37399; CM 20077;
UM 55-137, 55-636; Monroe County: UMMZ 103174.
DIAGNOSIS: A species of Tantilla resembling Tantilla coronata
in details of hemipenial structure and scutellation. The hemipenis
has both basal hooks, the postero-lateral hook adjacent to the sulcus
spermaticus in the basal third, and the antero-medial hook, approxi-
mately equal in size to the postero-lateral, in the mesal third of the
hemipenis. Ventrals range from 135 to 146; subcaudals between
45 and 63. Males appear to have more subcaudals than females;
too few specimens are available to demonstrate a trend in ventrals.
Females exceed males in size. Two types of head pattern are evi-
dent; all specimens from Dade County exhibit a completely black
head and neck with no light parietal band; a single specimen from
Key Largo, Monroe County, has a prominent but broken band sep-
arating the black head cap from the black nuchal collar. Males have
slightly longer tails than females, with tail length ranging (both sexes)
from 19.1 to 22.4 percent. Maxillary teeth 14, 15 or 16. Head broad-
ly rounded, with noncountersunk lower jaw.
DESCRIPTION OF HOLOTYPE: Adult male, 166.5 mm snout-vent
and 48.0 mm tail length. Ventrals 137; subcaudals 63; dorsal scales
smooth, in 15 rows throughout; 1 preocular, 2 postoculars, 1 anterior
and 1 posterior temporal, on each side. There are 7 supralabials on
the left and 6 on the right, with III and TV entering the orbit on both
sides. Infralabials are 5 on the left and 6 on the right; I to III contact
the anterior chin shields on the left, and I to TV on the right side
Mental plate contacts the anterior chin shields; posterior chin shields
two-thirds length of the anterior. In preservative the head dorsum
is black; no light parietal collar, black of the head extending along
the neck to the middle of vertebral scale TV and to scale row 2 lat-
erally. Mental and infralabials heavily suffused with black, as is
the rear half of infralabial VI at angle of the jaws. Snpralabials
I to IV and postoculars completely black, as is loreal region. Supra-
labials V and adjacent portions of VI and primary temporal lack
pigment; supralabial VII largely black, lacking pigment in most of
rear half. Dorsum is light tan, and venter white. No color notes
from life available.

THE NON-PENINSULAR POPULATIONS.
Schwartz (1953) based his diagnosis of Tantilla coronata mitrifer
upon three main points:


Vol. 10









VARIATION AMONG SOUTHEASTERN SNAKES


(1) "posterior ventrad extension of the black cap tends not to
reach the upper labial border . "

(2) "black collar does not extend farther posteriorly than the
fourth scale row in the midline . ."

(3) "with more ventral and less subcaudal scutes on the aver-
age ...

I have examined the holotype from Caesar's Head, Greenville
County, South Carolina (CHM 53-92.2), and six paratypes from
Roundtop Mountain, Rutherford County, North Carolina (AMNH
66207-66211). Other paratypes were not made available to me.
The holotype certainly agrees with the main points of the subspe-
cies diagnosis quoted above, but of the six paratypes I examined,
only one agrees with the holotype in character (1) in the other five,
the ventrad extension extends well past the upper labial border.
This variation appears commonly in other parts of the range; speci-
mens from Indiana, Edmonson County, Kentucky, and Jackson Coun-
ty, Florida, lack a ventrad extension to the black cap. Character (2)
also lacks exclusiveness: one of the six paratypes has the black nuchal
collar extending on to vertebral scale V, although in the other para-
types and holotype the collar terminates on scale IV. In a series of
37 T. coronata from coastal plain Mississippi, 62 percent (23) have
the nuchal collar ending on scale IV or anteriorly; in 38 percent (14).
it extends onto scale V. Thus, it appears that two of the main char-
acters used by Schwartz are not sufficiently exclusive to be of diag-
nostic value. This leaves only characters of (3) to be evaluated.
Ventrals and subcaudals of 5 female T. coronata mitrifer types
were compared with a series of 14 female coastal plain T. coronata
coronata from Charleston and Berkeley counties, South Carolina,
specimens which were examined by Schwartz. Admittedly five speci
means is an inadequate sample, but standard tests (the Student T)
were performed anyway, with the following results: there is a
difference in means of ventrals and subcaudals between coastal plain
females (138.3, 42.5) and T. coronata mitrifer females (142.8, 39.5).
The difference in ventrals is significant at the .05 level; that in sub-
caudals is not significant at the same level.
Schwartz states that T. coronata mitrifer has a darker dorsum
than does T. coronuta coronata. I observed that in the few living
T. coronata I have seen those specimens from the Upper Piedmont
were considerably darker than those from the Coastal Plain. In many
preserved specimens this distinction is still remarkably evident, and


1986










BULLETIN FLORIDA STATE MUSEUM


specimens from the Blue Ridge and Upper Piedmont constitute a
distinct group from those of the Coastal Plain. It seemed reason-
able that two distinct populations of Tantilla coronata might exist,
those of the Coastal Plain and those Schwartz called T. coronata
mitrifer, but including as well demes occupying the Upper Piedmont.
The region of intergradation might logically follow the Fall Line.
With this hypothesis in mind, I segregated Tantilla from North and
South Carolina, eastern Tennessee, Georgia, Alabama, Mississippi,
and Louisiana into three groups: Blue Ridge and Upper Piedmont,
Lower Piedmont and Fall Line, and Coastal Plain (table 5).

TABLE 5. SCALE CHAHACTERISTICS OF Tantilla coronata POPULATIONS

Upper Piedmont
Coastal Plain Fall Line and Montane

mean ventrals
males 131.6 134.2 132.0
females 1.38.1 141.9 142.2
mean subcaudals
males 46.9 46.5 46.3
females 43.5 44.4 42.6
mean proportionate tail
length (%)
males 20.7 20.5 20.1
females 18.6 18.5 18.3
mean collar width in
number of scales
males 3.1 2.9 2.6
females 3.1 3.1 2.8
distal vertebral scale
included in collar
(% of sample)
males 3 1.8 11.1 11.0
4 50.0 55.6 89.0
5 48.2 33.3
females 3 11.8
4 65.8 57.1 70.6
5 31.6 42.9 17.6
6 2.6


If one postulates that T. coronata mitrifer, in a wider sense geo-
graphically than Schwartz defined it, is characterized by more ven-
trals, fewer subcaudals, a narrower nuchal collar that tends to lie


Vol. 10









VARIATION AMONG SOUTHEASTERN SNAKES


more anteriorly on the neck, and a shorter tail, in combination with
a darker dorsum, in comparison with T. coronata from the Coastal
Plain, then the data cited tend to support each of these distinctions.
And in almost every character, specimens from the proposed region
of intergradation are intermediate between mitrifer (sensu latu) and
coronata (sensu strictu. Standard statistical comparison (the Student
T test) was made for each of these characters. Differences in mean
number of ventrals of both sexes, position of the band (data lumped,
as males and females show no significant differences) and collar width
in males are significant at the .05 level. Differences in mean number
of subcaudals and proportionate tail length in both sexes, and collar
width in females are not significant at .05.
On the basis of this analysis, I think a weakly defined population
of Tantilla coronata exists in montane regions and the Upper Pied-
mont of North and South Carolina, Georgia, eastern Tennessee, and
Alabama, to which the subspecific designation Tantilla coronata
mitrifer Schwartz might be applied. I consider the variation in scale
characters, however, to be insufficient to warrant such distinction.
Although a striking contrast in coloration of the dorsum apparently
exists, as illustrated by color descriptions below, living material avail-
able to me has been too meager to substantiate the value of this char-
acter. At this time, I think it is best to relegate the subspecies
T. coronata mitrifer to the synonymy of Tantilla coronata Baird and
Girard, and recognize no subspecies of this wide-ranging species.
It is appropriate at this point to define my concept of the species
Tantilla coronata, as developed from examination of 215 specimens
from all areas of its range.
Tantilla coronata Baird and Girard, 1853
HOLOTYPE: USNM 1876, collected in Kemper County, Mississip-
pi, by D. C. Lloyd.
DIAGNOSIS: A species of Tantilla characterized by the presence
of two basal hooks on the hemipenis, one posterolateral in position
and adjacent to the sulcus spermaticus in the basal third of the organ,
and the other, slightly smaller and anteromedial in position, in the
mesal third of the hemipenis. Ventrals range from 123 to 147; sub-
caudals between 34 and 53. Males have fewer ventrals and more
subcaudals than females, and longer tails. Tail to total length ratio
varies from 15.9 to 23.0 percent. A prominent light parietal band is
present in most (96.6 percent) specimens, separating the black cap
of the head from the black nuchal collar. The nuchal collar is usu-
ally (76 percent) 3 or fewer scales in width at midline, ranging from


1966









BULLETIN FLORIDA STATE MUSEUM


2 to 6 (mean, 3.0). On each side are usually one preocular, two
postoculars, one anterior and one posterior temporal, seven supra-
labials, and six infralabials. Supralabials III and IV enter the orbit;
infralabials I to IV contact the anterior chin shields, which may or
may not be separated from the mental plate by infralabials I. Pos-
terior chin shields two-thirds length of anterior. Anal plate divided.
Maxillary teeth (one side) 14 to 18, usually (55 percent) 16. Dorsum
ranges from tan in coastal plain populations to dark brown in mon-
tane localities; venter white.
GEOGRAPHICAL RANGE: T. coronata is found from Florida west
of the Suwannee River westward to the Mississippi River and north
to southern Indiana and Virginia.
A specimen of T. coronata from Coffee County, Alabama (UF
12387) was obtained within a day of its preservation, and the follow-
ing color notes taken. The dorsum was uniform light brown (army
brown, 6-A-10), becoming tan (rose blush, 5-C-9) laterally. The ven-
ter ranged from cream (9-D-2) beneath the tail to pinkish white
(tilleul-buff, 10-A-2) anteriorly. The nuchal collar was black, pre-
ceded by a gray-brown (manon, 6-A-9) parietal band. The black
dorsal surface of the head faded to gray (32-A-1) on internasals and
prefrontals. This specimen is representative of Coastal Plain popu-
lations.
Another specimen (UF 12409) collected in Cobb County, Georgia
presents a vivid contrast to the Coastal Plain population. In life
the dorsum was dark brown (sepia, 8-A-10) distally, becoming slightly
lighter (moose, 8-C-10) at midbody, and light brown (bark, 8-C-1l)
on the anterior third of the body. The sides of the body were dark
tan (bracken, 8-C-12). The pinkish gray (rosedust, 6-B-2) tail venter
became gradually lighter to pinkish white (folkstone, 13-A-3) on the
anterior third. The parietal band was cream (vanilla, 10-C-3), and
the nuchal collar and dorsal surface of the head were black. Identi-
cal coloration was present in a series of three T. coronata (UF 1240D2
from 2 miles south of Millry, Washington County, Alabama, well with-
in the Coastal Plain. In scalation and pattern, these specimens clearly
belong to Coastal Plain T. coronata. As the Black Warrior Rivet
system flows from the Piedmont to the Gulf of Mexico through this
region, it may well be that some gene flow from Upper Piedmont pop-
ulations, which extend south as far as Tuscaloosa, follows this drain-
age system well into the Coastal Plain.
Tantilla coronata extends up the eastern Mississippi Valley through
western Tennessee and Kentucky to southern Indiana. The Indiana
series does not differ appreciably from Coastal Plain Mississippi speci-


Vol. 10










TABLE 6. SUMMARY OF VARIATION IN CHARACTERS LACKING TAXONOMIC SIGNIFICANCE


% of each population varying from normal
Normal Variations
Character condition (No. cases) A B C D E F G

preoculars 1-1 2-1 (1)
1-2 (1) 1.0 4.4 5.3
0-0 (2)

postoculars 2-2 2-0 (1)
2-1 (2) 1.0 4.4 2.6 16.6 9.1 5.3
1-2 (4)
1-1 (3)

supralabials 7-7 7-6 (7)
7-4 (1) 1.5 8.2 3.8 9.1 27.2
6-6 (4)

supralabials III,IV-III,IV III,IV-II,III (1)
in orbit II,III-II,III (3) 1.0 9.1

infralabials 6-6 7-7 (1)
7-6 (2)
6-7 (5)
6-5 (7) 6.2 8.7 2.6 9.1 36.4
5-6 (2)
6-4 (1)
5-5 (3)










TABLE 6. SUMMARY OF VARIATION IN CHARACTERS LACKING TAXONOaMIC SIGNIFICANCE (continued)

% of each population varying from normal
Normal Variations
Character condition (No. cases) A B C D E F G

infralabials I-IV,I-IV I-IV,I-III (9)
contacting I-III,I-IV (5)
anterior I-V,I-IV (1)
chin shields I-IV,I-V (4) 6.7 8.7 2.6 5.6 18.2 5.3 45.5
I-IV,I-II (1)
I-III,I-II (4)
I,II IV,V;I-IV (1)
I-III,V;I-V (1)

temporals 1&1,1&1 1&1&1,1&1&1 (7)
1&1&1,1&1 (7)
1&1,1&1&1 (7) 6.2 4.4 9.0 11.1 .91
1&1,0 (1)
1&1,1&2 (1)

dorsal scale 15-15-15-15 17-15-15-15 (1)
rows 16-15-15-15 (2)
14-15-15-15 (2)
15-14-15-15 (1) 3.1 2.6 5.0 9.1
14-14-15-15 (1)
15-16-15-15 (1)
15-13-15-15 (1)
15-15-15-13 (1)

anal plate divided single (2) 0.5 1.3


A. T. coronata
B. T. relicta relicta
C. T. relicta neilli


D. T. relicta pamlica
E. Seahorse T. relicta
F. relicta x neilli intergrades


G. T. oolitica









VARIATION AMONG SOUTHEASTERN SNAKES


mens. A single specimen was available from southern Virginia, and
resembles those from coastal North Carolina. I have seen only two
T. coronata from Florida, both from the vicinity of Marianna, Jackson
County, which lies west of the Appalachicola River, and these re-
semble the Alabama and Mississippi specimens. Although no speci-
mens are in existence from the region between the Appalachicola
and Suwannee rivers, south of Chattahoochee County, Georgia, T.
coronata may be there. It does not cross the Suwannee River into
peninsular Florida.

VARIATION IN CHARACTERS OF NO TAXONOMIC SIGNIFICANCE.

Most of the variation found in characters, that lack taxonomic
significance is presented in Table 6. Temporals and infralabials
were found to be the most variable scales, and less than 10 percent
of these varied from the usual condition in each population sampled,
except in the T. relicta sample from Seahorse Key and the T. oolitica
sample. Both samples numbered only 11 specimens each, and while
small sample size probably influenced disproportionately the percent-
age of specimens varying from the norm, the facts that both popula-
tions are insular in nature and probably small in numbers may also be
of moment. In the 11 T. oolitica specimens, 4 of the 8 showing scale
aberrations were aberrant in multiple characters. Most cases of mul-
tiple aberrations observed in all samples were correlations between
variations in labial number and the number of supralabials entering
the orbit or infralabials contacting the anterior chin shields. Table
7 presents the percentages of each sample that vary from the normal
scutellation listed in Table 6.

TABLE 7. PERCENTAGES OF ABERRANT SPECIINENS IN
SOUTHEASTERN Tantilla POPULATIONS

% aberrant % with
in at least multiple
N one character aberrations

T. coronata 195 15.4 43.4
relicta x neilli intergrades 19 15.8 1/3
T. relicta neilli 78 19.3 26.6
T. relicta pamplica 18 27.8 1/5
T. relicta relicta 23 30.2 2/7
Seahorse T. relicta 11 36.4 1/4
T. oolitica 11 73.0 4/8
all peninsular populations 165 25.4 31.0


1966










BULLETIN FLORIDA STATE MUSEUM


The peninsular populations clearly show a trend toward more
aberrant individuals than does T. coronata. I attribute this to the
distribution of peninsular Tantilla in discontinuous demes, with gene
flow severely restricted by habitat barriers. The slightly lower per-
centage of aberrant individuals of T. relicta neilli may reflect greater
continuity of gene flow within the sandhills and mesic hammock habi-
tats, in contrast to the insular nature of the occurrence of scrub. No
significant differences were found between the sexes.
Through an oversight on my part, one character, not listed above,
was not recorded for all specimens examined. This is the contact
of the mental plate with the anterior chin shields, and its corollary
condition, separation from them by infralabials I. In view of the
importance attributed to this character in some descriptions of Tan-
tilla species, 53 specimens available to me at the time the oversight
was discovered were checked to determine whether or not the char-
acter might be taxonomically useful. Table 8 tabulates the results.
Clearly, either condition may be encountered within the same
population, and from this small sample, equal frequency is indicated.
No correlations with sex were found. Descriptions of Tantilla species
that stress this character should be re-evaluated.

TABLE 8. CONTACT OF MENTAL AND ANTERIOR CHIN SHIELDS
IN SOUTHEASTERN Tantilla POPULATIONS

N Contact No contact

T. coronata 22 15 7
T. oolitica 3 1 2
T. relicta relicta 6 3 3
T. relicta neilli 10 5 5
7'. relicta pamlica 2 2
relicta x neilli intcrgrades 2 2
Seahorse T. relicta 8 2 6

Total 53 30 23


Maxillary dentition of several Tantilla species was described by
Smith (1940: 61). In the group to which T. coronata belongs the
two abruptly enlarged, grooved rear teeth are not in line with the
anterior smaller teeth and are separated from them by a diastema.
The 83 maxillae, I have examined in this study, 49 from T. coronata,
3 from T. oolitica, and 31 from T. relicta, all have a similar morph-
ology. The diastema varies from one to two tooth spaces in width,


Vol. 10









VARIATION AMONG SOUTHEASTERN SNAKES


and the enlarged teeth are approximately three times the size of the
preceding teeth. The smaller teeth decrease slightly in size as they
approach the diastema, and are strongly recurved. Table 9 presents
data on the total maxillary counts (enlarged plus smaller teeth)
recorded.

TABLE 9. NUMBER OF MAXILLARY TEETH (LEFT SIDE) IN
SOUTHEASTERN Tantilla POPULATIONS

Number of Teeth
14 15 16 17 18 Mean

T. coronata 6 16 26 1 15.3
T. oolitica 1 1 1 15.0
T. relicta relicta 2 6 5 15.2
T. relicta neilli 3 6 4 15.7
T. relicta pamlica 2 3 14.2
7'. relicta, all subspecies 7 15 9 15.4


Although T. coronata tends to have one more maxillary tooth
(55 percent have 16 or more) than does T. relicta (71 percent have
15 or less), there is no significant difference in mean tooth count.
Two few data were recorded concerning the position of the
retracted hemipenis to evaluate this character properly, but the little
information available suggests that it may be of little taxonomic use
in this species group. The retracted hemipenis was found to extend
to subcaudals 9 to 11 in T. coronata, 10 to 12 in T. relicta, and 8 in
the single T. oolitica in which it was measured. The postero-lateral
basal hook is located at subcaudals 3 or 4 in T. coronata, 4 or 5 in
T. relicta, and 3 in T. oolitica. The antero-medial basal hook, ab-
sent in T. relicta, is located at subcaudal 5 in both T. coronata and
T. oolitica. Terminal spination begins at the level of subcaudals 5
to 7 in T. coronata, 7 or 8 in T. relicta, and 5 in T. oolitica.

GROss ECOLOGY
Tantilla r. relicta is restricted, in areas of syntopy with T. r. neilli,
to scrub habitat on St. Lucie fine sand. In areas of allopatry, T. r.
relicta may inhabit sandhills, as at the type locality, Babson Park, or
coastal dunes and scrub. T. r. neilli is found only in sandhills and
dryer mesic hammock areas. 7T. r. relicta appears completely fossorial,
while T. r. neilli, although semifossorial, may be found occasionally
beneath logs, debris or rocks. Both subspecies inhabit the mounds


1966










BULLETIN FLORIDA STATE MUSEUM


of loose sand thrust up by Geomys pinetis, the pocket gopher. The
distribution of both subspecies is confined to areas of dry soil in Flor-
ida, as is that of T. r. pamlica in the coastal dunes and scrub (figure 2).
Tantilla coronata, however, seems to have broader habitat tolerances,
being found in sandhills, hardwood forests, and pine woods. I have
been informed (J. Dobie, personal communication) that it is fre-
quently collected from rotten stumps in seasonally flooded pine flat-
woods of Mississippi and Louisiana. Collecting notes on 22 museum
specimens include the following situations: rotten stump, in rotten
pine log, in pine straw, under log in sandhills (3), under rocks (9),
and in spider web! Habitat notes on these specimens include pine
woods, pine-hardwood upland, edge of hardwoods, edge of river, and
second growth of thin, oak-forested hillside. Neill (1951: 49) men-
tions collecting T. coronata by "digging into piles of decaying bark
scrap, by uprooting rotten stumps, or by overturning large rocks and
boulders on the wooded hillsides," and finding the eggs in debris
from a rotting longleaf pine in this locality. Neill and Boyles (1957)
discuss the eggs of a female collected "in leaf litter on a rocky ex-
posure along the Black Warrior River" in Tuscaloosa County, Ala-


Aug. 0o A A
A ** A
A*
0 A A'*A A A

0 0A* A A


0 Feb, 0 A
A A


A
0
0


80 120 160 200
A

Fig. 6. Scatter diagrams plotting snout-vent length against month of col-
lection: A, T. coronata; B, T. relicta relicta; C, T. relicta neilli. Solid symbols are
mature individuals; hollow, immature; divided, apparently pubescent. Circles
represent males, triangles, females.


Vol. 10












VARIATION AMONG SOUTHEASTERN SNAKES


0 0


e* A

* A A& A
a A


a
o %ooc


*- *




* A

A A

A
A A

*


S I I i

so 120 160

snout-vent length (mm)


Aug. I


B


A A

*


120
120


Aug.


160


200


* A*

A


0 00


o0 a


200









294 BULLETIN FLORIDA STATE MUSEUM Vol. 10

bama. Hardy (1952: 188) reported five specimens from Roundtop
Mountain, Rutherford County, North Carolina, as "taken in a cleared
field beneath rocks." Schwartz (1953: 156) collected the holotype of
T. coronata mitrifer "beneath a flat rock, about 18 inches in diameter"
on a slope wooded with Pinus echinata and Ilex opaca. One of his
paratypes was "caught on a wooded hillslope beneath a flat rock,"
two others "under one rock in a pine-oak woods," and five were dug
out of the ground from depths of 2 to 12 inches on a steep, rocky
hillside. It seems rather clear from these accounts that while T.
coronata may burrow as most small snakes probably do, it is char-
acteristically secretive and is usually associated with forests contain-
ing pines.
The only habitat data available in the literature on Tantilla oolit-
ica are those of Duellman and Schwartz (1958: 306), who state: "In
southern Florida Tantilla has been collected only on the eastern rim
and on Key Largo. Individuals have been found beneath rocks and
boards in sandy soil in the pine woods, in hammocks, and in edifi-
carian situations. The specimen from Key Largo was dislodged from
a rotten stump." From their color notes they appear not to have
confused T. relicta pamlica from coastal scrubs in southeastern Flor-
ida with T. oolitica. A specimen not available to Duellman and
Schwartz was collected beneath a log in a pasture southwest of Miami,
and the holotype was found beneath a board on a vacant lot in Miami.
Dennis R. Paulson (personal communication) states the holotype and
two others were "collected under logs and trash in an empty lot with
much shrubby growth and a few Pinus elliotii. The area was un-
doubtedly pine woods before clearing, with oolitic limestone over-
lain by some sand. The Martin Co. record is probably more charac-
teristic of central Florida habitat-the specimen was under the dead,
fanned out on the ground, leaves of a Serenoa repens partially
buried under St. Lucie Fine sand, in typical Pinus clausa and Cerati-
ola habitat The Martin County specimen referred to is a Tantilla
relicta pamlica, and the passage quoted well describes the habitat
difference between the two species in southeastern Florida.

SIZE AND SEXUAL MATURITY
Upon plotting snout-vent length and maturity against month of
collection (figure 6), some differences between T. coronata, T. r.
neilli, and T. r. relicta become apparent. T. coronata and T. oolitica
are the largest species, and 'T. relicta the smallest. Within T. relicta,
the nominate subspecies is intermediate, and neilli the largest, with









VARIATION AMONG SOUTHEASTERN SNAKES


T'. relicta pamlica smallest in size, to judge from the small sample of
the latter available. Table 10 presents size statistics of the main
populations of southeastern Tantilla.

TABLE 10. SNOUT-VENT LENGTH (IN MM) OF SOUTHEASTERN
Tantilla POPULAInONs

Smallest Smallest Mature Largest
Juvenile Male Female Male Female

T. coronata 78 134 153 190 217
T. oolitica 92 147 165 171 246
T. relicta relicta 77 120 124 166 187
7'. relicta neilli 80 142 134 190 194
T. relieta pamlica 99 129 146 160 176


T. coronata apparently becomes mature at about 180 mm, T. r
neilli and T. r. relicta at 120 mm. The samples of T. oolitica and
T. r. pamlica are too small to permit such analysis. Males mature
at slightly shorter lengths than do females. I discern four size
groups in T. coronata: juveniles less than 105 mm; immature, young
and young adult individuals from 110 to 150 mm; adults from 150 to
180 mm; and very large specimens over 180 mm. In both the T.
relicta populations considered, only three size groups may exist:
juveniles and young individuals below 120 mm in both; in neilli
adults from 120 to 170 mm and very large specimens over 170 mm;
and in the nominate subspecies adults from 120 to 155 mm and large
individuals over 155 mm. It seems from these approximations that all
three species become mature during their second growth season,
but do not necessarily breed until the following year. Force (1985:
651) found that the Tantilla gracilis population in Oklahoma (where
the active season is shorter than in the southeast) is composed of
three age groups, that males mature at the same age as females but
at a shorter length, and that maturity probably is reached in the
third growth season at about 2V2 years of age.

DIscussion
HEAD PATTERNS.
With respect to the populations of Tantilla relicta, an intriguing
question recurred frequently during this study: What is the relation-
ship of pattern to habitat in this species? The banded populations,
T. r. relicta and T. r. pamlica characteristically inhabit scrub, a plant












BULLETIN FLORIDA STATE MUSEUM


- IA


Fig. 7. a, Coastal Plain T. coronata (UF 12387, Coffee Co., Ala.); b, T.
oolitica holotype (UF 17326, Dade Co., Fla.); c, T. relicta relicta holotype (UF
12421, Polk Co., Fla.); d, T. relicta neilli holotype (UF 12406, Alachua Co., Fla.);
e, T. relicta pamlica holotype (UF 12430-2, Palm Beach Co., Fla.).


Vol. 10


''
r' ,


;


I
I , i
c 'r
ILI~










VARIATION AMONG SOUTHEASTERN SNAKES


association that grows on white St. Lucie Fine sand, and which is a
seral stage in dune succession (Laessle, 1958). The nonbanded T. r.
neilli is found in both sandhills and mesic hammock, but is more
common in the former, and possibly occurs only in those mesic ham-
mocks bordering sandhills areas. In areas of syntopy, relicta and
neilli seem completely segregated ecologically from each other by this
habitat preference, yet in the southern ridge section of Florida where
neilli does not occur, relicta is found in both scrub and sandhills.
I suggest as a purely speculative hypothesis that perhaps selec-
tion acts against uniform head patterns on white sand substrates.
T. relicta is a "submerged basker." It lies, in captivity at least, just
beneath the sand, with the head at the surface. A banded head pat-
tern would tend to break up head outline on the white sand and
render a basking Tantilla less obvious to avian predators, to whom
these diminutive snakes should pose no greater problem to ingest

ventrlsi


males
151








110
sub

males


females b


~f








caudals
females


d
C





'7


Fig. 8. Variation in ventrals and subcaudals: a, T. coronata; b, T. oolitica;
c, T. relicta relicta; d, T. relicta neilli; e, T. relicta pamlica; f, T. relicta from Sea-
horse Key; g, series possibly intergradient between relicta and neilli. Vertical
line is range; horizontal line, mean; hollow bar, two standard deviations of the
mean; solid bar, twice the standard error. Sample size is indicated beneath each
histogram.









BULLETIN FLORIDA STATE MUSEUM


than a large earthworm. The only known instance of avian preda-
tion on Tantilla was one D. Jenni (personal communication) found in
a cattle egret (Bubulcus ibis). I suspect that many of the birds in-
habiting scrub, sandhills, and hammock prey upon Tantilla. Sub-
merged basking is perhaps less dangerous for small snakes in sand-
hills and hammock, both habitats with abundant vegetation on the
ground. The scrub, however, a fire-climax association, typically has
wide areas of open sand bearing little or no vegetation. The ability
to bask with less likelihood of detection from above would thus make
scrub more habitable for banded Tantilla, and reduce the likelihood
of competition by a possibly more effective, blackheaded competitor.
The predominantly nonbandcd condition of disjunct populations of
T. relicta in Charlotte County and on Seahorse Key is conceivably
correlated with the overgrown, mature scrubs found there. Perhaps
as the banded pattern becomes less advantageous a simple Mendelian
dominant pattern reduces the frequency of banded phenotypes in
these populations. This may also account for the presence of non-
banded T. relicta in the mature scrubs east of the St. Johns River.

INTERGRADATION.
Most of the specimens treated above as possible intergrades be-
tween T'. r. relicta and T. r. neilli are not truly intermediate between
neilli and relicta, but show some similarities to both. For instance,
head pattern and ventrals seem closer to neilli, while subcaudals and
tail length closely approximate relicta. Most of these specimens are
from Putnam, Volusia, and Seminole counties, east and south of Lake
George. Others are from scattered areas in Marion, Lake, Levy,
Citrus, and Pinellas counties. I suspect that relicta and neilli do
not intergrade along a continuous front, but that some gene flow
occurs through secondary intergradation, in the sense of Mayr (1963).
These two long-isolated populations have probably come into con-
tact since the closing of the Suwannee Straits and intergradation may
be occurring only in those areas of longest contact, notably in the
area around Lake George. A single specimen is available from Eau
Gallie, Brevard County, which may represent a similar hybrid be-
tween the central populations and T. r. pamlica. In characters of
scutellation it resembles pamlica, but its pattern is nonbandcd. In-
tensive collecting in Volusia County from the St. Johns to the At-
lantic Ocean and south to Cape Canaveral will probably provide a
less speculative interpretation of these Tantilla populations. For the
time being it is reasonable to consider them intergrades.


Vol. 10









VARIATION AMONG SOUTHEASTERN SNAKES


males females



















11
17 .






Fig. 9. Variation in proportionate tail length, expressed as per cent of total
length. Symbols as in Fig. 8.

PLEISTOCENE INFLUENCES UPON SPECIATION.

Probably the most significant influence upon present distribution
of the Floridian herpetofauna was that of fluctuating Pleistocene
physiography. Correlations of ranges of such Florida endemics as
Ophisaurus compressus (McConkey, 1954), Stilosoma extenuatum
(Highton, 1956), Neoseps reynoldsi (Telford, 1959), and Eumeces
egregius (McConkey, 1956; Mount, 1961) have been made with ma-
rine terrace levels, and the present study shows that the same isolat-
ing phenomena probably contributed to divergence within the Florida
Tantilla coronata complex.
Pertinent geological factors in the history of the peninsula have
been presented by Cooke (1939), Laessle (1958), Puri and Vernon


1966









BULLETIN FLORIDA STATE MUSEUM


(1960), and Telford (1959). Terrace levels are thought to represent
marginal marine sediments deposited during cycles of eustatic adjust-
ments in sea level associated with maximal and minimal develop-
ment of ice in the Pleistocene.
Tantilla probably invaded Florida from Mexico, possibly along a
coastal corridor of suitable xeric habitat during the Pliocene or Early
Pleistocene, and populations of this snake became isolated on islands
of the Okefenokee Sea. This may have constituted the initial isola-
tion between precursors of T. r. relicta and T. r. neilli, with ancestral
relicta occupying the island today forming the Lake Wales Ridge,
and ancestral neilli existing on the large island in the present day
Alachua County area. This isolation continued during succeeding
high sea levels. During the early Wisconsin ancestral coronata may
have extended its range to the southeast along a corridor of pine flat-
woods through the St. Johns and Kissimmee river valleys, and an-
cestral relicta to the east and south along Pamlico terrace in dune
habitat.
During the Peorian interglacial, T. oolitica may have been isolated
from the coronata stock on high ground in the Dade County of today,
while intervening coronata populations south of the Suwannee River
were exterminated. The St. Johns and Kissimmee rivers today effec-
tively isolate both T. oolitica and T. r. pamlica from the central popu-
lations. T. r. neilli may be extending its range southward along the
central ridge and the west coast in sandhills habitat, forcing the pos-
sibly less effective competitor, T. r. relicta, into scrub habitat wherever
they have come into contact, accompanied by occasional hybridiza-
tion. Selection against uniform head pattern by predators may have
enabled the banded relicta to remain syntopic with neilli in north
central Florida, where it is restricted to scrub habitat.
Several species (notably Coluber constrictor, Terrapene carolina,
Kinosternon bauri, and Eumeces egregious) on the Florida Keys show
greater affinity to populations in northern Florida and southern Geor-
gia than to those of the intervening peninsula. McConkey (1957) has
suggested that a corridor along the St. Johns and Kissimmee river val-
leys during the Wisconsin glaciation may have provided suitable
habitat through which Eumeces egregious could have reached the
Keys, thus producing the confusing picture of two closely similar pop-
ulations (E. e. egregious and E. e. similis; or T. oolitica and T. coronata)
separated by strongly dissimilar, evolutionarily older populations
(E. e. onocrepis, and T. relicta). I believe the paleozoogeographic
dispersion of T. oolitica was similar.


Vol. 10










1966 V
























Tantilla

oolitica

r.relicti

r.neilli

r.pamlica

intergrades


AVIATION AMONG SOUTHEASTERN SNAKES


*
0


* 0


o'p



Fig. 10. Distribution of Tantilla taxa in peninsular Florida.

SUMMARY

The Tantilla coronata complex in southeastern United States has
been re-evaluated by the study of 881 specimens, 165 of them from
peninsular Florida. Two named subspecies T. coronata wagneri
(Jan) and T. coronata mitrifer Schwartz, are placed in synonymy with
Tantilla coronata Baird and Cirard. The range of T. coronata does
not extend south of the Suwannee River. Two new species, T. oolitica









302 BULLETIN FLORIDA STATE MUSEUM Vol. 10

and T. relicta are described from peninsular Florida. T. oolitica,
most similar to T. coronata in hemipenial structure, is restricted to
Dade and Monroe counties. T. relicta, which ha's but a single basal
hook on the hemipenis, in contrast to two in the other two species,
geographically separates T. coronata from T. oolitica. Three sub-
species of T. relicta are recognized on the basis of differences in pat-
tern, scutellation, body proportions, and habitat. T. r. neilli occurs
in sandhills and mesic hammock of north central Florida. T. r. relicta
inhabits central Florida, being restricted to scrub habitat where syn-
topic with T. r. neilli, but occurring in sandhills as well where allo-
patric in south central Florida. T. r. pamlica is found along a nar-
row strip of coastal dunes and scrub in south eastern Florida. Some
gene flow is indicated, possibly through secondary intergradation,
between T. r. neilli and T. r. relicta in scattered localities, especially
in the Volusia County area. Disjunct populations of T. relicta occur
along the west coast from Charlotte County to Seahorse Key, Levy
County. It is believed that all three species, T. coronata, T. relicta,
and T. oolitica, are derived from a common precursor of the early
Pleistocene, from which they diverged during the successive fluctu-
ations of sea level that characterized the Pleistocene glacial-inter-
glacial stages.

KEY TO THE Tantilla OF SOUTHEASTERN UNITED STATES

1. (a) Hemipenis with two basal hooks ----------- ------2
(b) IHemipenis with one basal hook .-- ...... .... ....................3
2. (a) A light parietal band separating black nuchal collar from black head
dorsum; populations north and west of Suwannee River T. coronata
(b) No light parietal band, black of head and neck continuous (except
Key Largo specimens); southeastern Florida -.. --------T. oolitica
3. (a) No definite light parietal band, black of head and neck almost con-
tinuous; head not pointed but narrowly rounded; ventrals 123-135
and subcandals 51-67 in males, 129-142 and 46-60 in females; sand-
hills and mesic hammocks of northern and north-central peninsular
Florida .. ----- --------------------.-- --...T. relicta neilli
(b) Usually (over 80 percent) a definite light parietal band; head pointed
with tendency toward countersunk lower jaw; ventrals 115-131 and
subcaudals 44-59 in males, 119-134 and 40-55 in females; north-
central and southern Florida ---- ----- ------ .
4. (a) Ventrals 115-118 in males, 119-129 in females; a tendency in most
specimens (90 percent) toward loss of dark pigment on parietals,
both temporals, and snout; coastal dunes and scrubs from Brevard
County to Dade County ..--- ..------------- T. relicta pamlica










VARIATION AMONG SOUTHEASTERN SNAKES


(b) Ventrals 117-131 in males, 120-134 in females; seldom reduction of
dark pigment on head except where light parietal band is situated;
scrub from Marion County south along central ridge and sandhills
of southern Polk County south; isolated populations, usually with no
light parietal band in coastal scrubs of Charlotte, Sarasota, and
Pinellas counties, and on Seahorse Key-...-..-----------T. relicta relicta

LITERATURE CITED

Baird, S. F., and C. Girard
1853. Catalogue of North American reptiles in the museum of the Smithsonian
Institution. Part 1, Serpents. Washington. 172 pp.
Blanchard, F. N.
1938. Snakes of the genus Tantilla in the United States. Field Mus. Nat.
Hist., Zool. Ser., 20: 369-376.
Boulenger, G. A.
1896. Catalogue of the snakes in the British Museum (Natural History). Lon-
don, Taylor and Francis. Vol. 3, 727 p. (1961 reprint, J. Cramer,
Weinheim).
Carr, A. F., Jr.
1940. A contribution to the herpetology of Florida. Univ. Florida Publ.,
Biol. Sci. Ser., III (1): 1-118.
Cooke, C. W.
1939. Scenery of Florida interpreted by a geologist. Bull. Florida Geol. Sur-
vey, 17: 1-118.
Cope, E. D.
1900. The crocodilians, lizards, and snakes of North America. U. S. Natl.
Mus. Report for 1898, Washington, Government Printing Office, p.
153-1270.
Dowling, H. G.
1951. A proposed standard system of counting ventrals in snakes. British J.
Herpetol., 1 (5): 97-99.
Duellman, W. E., and A. Schwartz
1958. Amphibians and reptiles of southern Florida. Bull. Florida State Mus.,
Biol. Sci., 3 (5): 181-324.
Dumeril, A. M. C., and G. Bibron
1854. Erpttologie g6ndrale ou histoire naturelle complete des Reptiles. Paris,
Roret. Tom. VII, part 2: 781-1536.
Force, E. R.
1935. A local study of the opisthoglyph snake Tantilla gracilis Baird and
Cirard. Pap. Michigan Acad. Sci., Arts, Lett., 20, 1934: 645-659.
Garman, S.
1883. The reptiles and batrachians of North America. Mus. Compar. Zool.,
Mem., 8, Cambridge, Harvard Univ., 185 p.
Hardy, J., Jr.
1952. The crowned snake, Tantilla coronata coronata, in North Carolina.
Copeia, 1952 (3): 188.










304 BULLETIN FLORIDA STATE MUSEUM Vol. 10

Highton, R.
1956. Systematics and variation of the endemic Florida snake genus Stilosoma.
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Jan, G.
1862. Enumerazione sistematica delle specie d'ofidi del grupo Calamaridae.
Archiv. Zool., Anat., Fisiol., 2 (1): 1-76.
Jan, G., and F. Sordelli
1866. Iconographie g6n6rale des Ophidiens. Paris. Tome premier, livr. 15.
(1961 reprint, J. Cramer, Weinheim).
Lacssle, A. M.
1958. The origin and successional relationship of sandhill vegetation and sand-
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Maerz, A., and M. R. Paul
1950. A dictionary of color. McGraw-Hill, New York. 207 p.
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1963. Animal species and evolution. Belknap Press of Harvard Univ. Press,
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1954. A systematic study of the North American lizards of the genus Oph-
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McConkey, E. II.
1957. The subspecies of Eumeces egregious, a lizard of the southeastern United
States. Bull. Florida State Mus., Biol. Sci., 2 (2): 13-23.
Neill, W. T.
1951. Notes on the natural history of certain North American snakes. Publ.
Res. Div., Ross Allen's Reptile Instit., 1 (5): 47-60.
Neill, W. T., and J. M. Boyles
1957. The eggs of the crowned snake, Tantilla coronata. Herpetologica, 13:
77-78
Puri, H. S., and R. O. Vernon
1960. Notes of the surficial geology of central peninsular Florida. IN: Late
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1953. A new subspecies of crowned snake (Tantilla coronata) from the south-
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Smith, H. M.
1940. Descriptions of new lizards and snakes from Mexico and Guatemala.
Proc. Biol. Soc. Washington, 53: 55-64.
Snedecor, G. W.
1946. Statistical methods. Iowa State College Press, Ames, 4th edit., 485 p.
Telford, S. R., Jr.
1959. A study of the sand skink, Neoseps reynoldsi Stejneger. Copeia, 1959
(2): 110-119.
Wright, A. H., and A. A. Wright
1957. Handbook of snakes of the United States and Canada. Ithaca, Con-
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