US ISSN: 0025-4231
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BULLETIN OF TME
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DEPARTMENT OF HERPETOLOGY
THE NATURAL HISTORY SOCIETY OF MARYLAND, INC.
MDHS . A Founder Member of the Eastern
Seaboard Herpetological League
/
VOLUME 38 NUMBER 1
MARCH 2002
BULLETIN OF THE MARYLAND HERPETOLOG1CAL SOCIETY
Volume 38 Number 1
March 2002
CONTENTS
A New Subspecies of the Lizard Sceloporus mucronatus (Sauria,
Phrynosomatidae)
Robert G. Webb, Julio A. Lemos-Espinal, and Hobart M.
Smith ................................................................................... 1
Observations on the Diet of Tmchemys gaigeae (Testudines: Emydidae)
James N. Stuart and Charles W. Painter ....................... 15
Evolutionary Speciation in the Alligator Lizards of the Genus Barisia
Hobart M. Smith, Theresa M. Burg and
David Chiszar .................................................................. 23
A New Species of Conophis (Reptilia: Serpentes) From Los Tuxtlas, an Area
of High Endemism in Southern Veracruz, Mexico
Gonzalo Ferez-Higareda, Marco A. Lopez-Luna and
Hobart M. Smith . . . 27
BULLETIN OF THE
Volume 38 Number 1
March 2002
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Herbert S. Harris, Jr.
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Herbert S. Harris, Jr. Tim Hoen
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Volume 38 Number 1
March 2002
A New Subspecies of the Lizard Sceloporus
mucronatus
(Sauria, Phrynosomatidae)
Robert G. Webb, Julio A. Lemos-Espinal, and Hobart M. Smith
Abstract
Specimens from the northwesternmost part of the range of Sceloporus
mucronatus in the Mexican state of Mexico are described as a new subspecies
of S. mucronatus. Pattern and scalation features of the four recognized sub¬
species geographically segregate into two groups, each with two subspecies-
one including the new taxon and nominotypical S. mucronatus, the other S. m.
omiltemanus and S. m. aureolus.
Olson (1998) commented on specimens from near Huichapan, Hidalgo,
Mexico, suggesting a subspecific relationship of S. mucronatus and S. poinsettii.
Auth et al. (2000) employed name-combinations reflecting this relationship.
Earlier, Mindell et al. (1989:61) used a specimen identified as S. poinsettii from
Hidalgo, Mexico (BYU 38634, "Esc. Conalep, Pachuca") as a voucher for
allozyme studies.
The northwesternmost populations of S. mucronatus have some pattern
features that obtain in S. poinsettii (expressed in varying degrees, may be geo¬
graphically variable). Range-wide variation in numbers of dorsal scales and
femoral pores of S. mucronatus is encumbered by that known for S. poinsettii
(personal data, RGW). Scalation of the posterior frontal-frontoparietal area is
often irregular in S. poinsettii (not in S. mucronatus). Pattern features shared
by both species include (1) black and white tail bands (consistent and distally
under tail in poinsettii, distinctness variable and usually absent under tail in
mucronatus), (2) pale intertympanic band, (3) pale scale in black collar above
shoulder, (4) juvenile throat pattern with pale, dark-bordered, median streak,
(5) large females with blue belly patches, and (6) vertebral, dark-blotched
dorsal body pattern (usually only males in some poinsettii populations). Black
longitudinal lines on the back (S. mucronatus aureolus) may occur in speci¬
mens of S. poinsettii. Geographically, the two species (poinsettii and mucronatus)
are widely separated; the southernmost known localities in central and east¬
ern Mexico of S. poinsettii are 32 km (Hwy 45) NW Fresnillo, Zacatecas (UTEP
Bulletin of the Maryland Herpetological Society
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Volume 38 Number 1
March 2002
6179) and 2 km NW Concepcion del Oro, Zacatecas (Ralph W. Axtell 6170),
which are, respectively, about 485 air km NW and 525 air km NNW of the
nearest, northernmost localities for S. mucronatus (Huichapan area, Hidalgo).
A series of 32 specimens from four different sites in Edo. de Mexico
differ from S. m. mucronatus in average size of dorsal scales and aspects of
dorsal color pattern. These characters suggest a new taxon (described herein),
having close taxonomic affinity with nominotypical S. m. mucronatus (see
Comparisons and Key). The Mindell et al. (1989) specimen (see above) prob¬
ably is representative of S. m. mucronatus ; however the specimen (BYU 38634,
Escuela Conalep, ca 1.0 km E Pachuca) cannot be located (Jack W. Sites, in litt.
to RGW, 11 January 2001). Museum codes used in the text to designate speci¬
men repository are BYU (Brigham Young University), UBIPRO (Unidad de
Biotechnologia y Prototipos, Esc. Nac. Estud. Prof. Iztacala, UNAM), UCM
(University of Colorado Museum), and UTEP (University of Texas at El Paso).
Sceloporus mucronatus olsoni nov. subsp.
Type material (all Edo. Mexico, Mexico, collected by Julio Lemos-Espinal).
Holotype, adult female, UCM 61083 (JLE 4503), San Juan Acazuchitlan [GPS
20°08,3.3"N, 99°36/15.8"W], 2646 m, 6 May 2000. Thirty-one paratypes:
UBIPRO 4504, 4506, 5000, 5006, 5008, 5012-13, 5017, 5020; UCM 61082, 61084-
92; UTEP 18566-69 (23 paratopotypes, same collection data as holotype);
UBIPRO 8557-59, UTEP 18571-73, San Miguel at Km 99.5 (Hwy 57) [GPS
20°02,28.9"N, 99°34,26.9MW], 2533 m, 28 May 2000; UBIPRO 8560, Hwy 57 at
Km 104.6 [20°04'24.0"N, 99°36'38.2”W], 2621 m, 28 May 2000; UTEP 18570,
Km 113.4 (Hwy 57) [GPS 20°05'36.3"N, 99°41,26.3"W], 2609 m, 21 May 2000.
Description of holotype. The top of the head is mostly black with an indis¬
tinct pale intertympanic band (indicated by three spots), and a black collar 2-
3 scales wide (with three included pale scales) having uninterrupted anterior
and posterior whitish borders, each about two scales wide. The distinct dor¬
sal body pattern consists of a vertebral black area (fading posteriorly) with
scattered whitish spots. The sides of the body are pale brownish and pattern¬
less. The dorsal pattern of the holotype is illustrated in Fig. 1. The underside
of the head has a diffuse blue and white mottled pattern, and a prominent
midventral, longitudinal pale (and dark-bordered) streak. The black collar
encroaches laterally onto the chest region. Enlarged blue-black groin patches
are attenuated anteriorly to just behind the axillary region, and separated
medially (midbody) by 7-8 scales.
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March 2002
The holotype, SVL 90 mm with incomplete tail (partly severed distally),
has 34 dorsal scales, 41 scales around midbody, 12-13 femoral pores (series
separated medially by about 13 scales), an entire anterior frontal, 1-1
frontoparietals separated by an azygous scale, the prefrontals in medial con¬
tact, 2-2 canthals, 1-1 loreals, 1-1 preoculars, and the anteriormost sublabials
(labiomentals) not contacting the mental.
Diagnosis. A subspecies of Sceloporus mucronatus distinguished from S.
m. aureolus and S. m. omiltemanus in adults having a dorsal pattern of black
vertebral blotches, and from S. m. mucronatus in adults having a reddish color
in life and smaller dorsal scales (see Comparisons and Key); S. m. mucronatus
and S. m. olsoni also occupy distinctive habitats.
Description. Adults of both sexes have a pronounced reddish color in
life (fading in preservative), which may vary in intensity and reflect breeding
condition (specimens collected 6, 21 and 28 May), and is absent in lizards less
than about 70 mm SVL (JLE). The dorsal surface of the head (large-scaled
part) is dark brown to black, paler posteriorly, patternless, but may have tiny
whitish flecks (three dots across parietals most consistent). A pale
intertympanic band is usually faded and indistinct (absent or nearly so in
UBIPRO 5013, UCM 61087, UTEP 18566; most distinct in UBIPRO 8560). The
uninterrupted, black collar (about 3-4 scales wide) has white anterior and
posterior borders two scales wide (both anterior and posterior borders may
be narrowly interrupted medially). A pale scale(s) may occur within the black
collar above the shoulder (UTEP 18571). The vertebral area between the col¬
lar and intertympanic band usually is darker than laterally. The basic dorsal
body pattern in adults of both sexes is a dark (black) vertebral area that is
interrupted by pale transverse solid bars or series of spots to form dark verte¬
bral blotches; the sides of the body are mostly uniformly pale brownish, but
may have some pale scales (e.g., UTEP 18572). Small specimens 54 and 40
mm SVL (UTEP 4580-81, regarded as intergradient, see comments under S.
m. mucronatus) have indistinct vertebral dark areas with prominent white spots
oriented transversely and in dorsolateral rows; the smallest has indistinct
whitish postocular marks. Blotched patterns become more distinct with in¬
creasing size. Adult females have a distinct pattern of either dark vertebral
blotches separated by solid bars, or a mostly continuous black vertebral area
(less well-defined blotches) with intervening white marks staggered and in¬
terrupted (two females in Fig. 1). Patterns may be faded /indistinct in both
sexes of large adults (female, UTEP 18570, 70 mm SVL, and two males in Fig.
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Volume 38 Number 1 March 2002
Fig. 1. Sceloporus mucronatus olsoni from Edo. de Mexico (type-mate¬
rial, see text). Left to right: UTEP 18571, male, 90 mm SVL; UBIPRO 8559,
male, 94 mm SVL; UBIPRO 8560, female, 91 mm SVL; UCM 61083 (holotype),
female, 90 mm SVL.
1). Tails are banded black (widest) and whitish (only about one scale wide),
but the degree of contrast is variable; the underside of tails (reddish in life,
JLE) is usually immaculate (indistinct distal dark banding in UBIPRO 5000,
5008; UCM 61086, 61090; UTEP 18569). The juvenile throat pattern (both sexes)
consists of a coarse dark blue, irregular barring with a dark-bordered, pale
longitudinal median streak; this pattern generally persists in large females
(lateral dark barring diffuse, mostly pale-dark mottling but with evidence of
pale median streak), but is more diffuse and bluish in large males and mostly
uniform blue in the largest males (UTEP 18571, SVL 90 mm). In large males,
the black collar is mostly continuous across the throat, blackish marks occur
on the chest, and black groin patches extend anteriorly as medial borders of
the blue belly patches (separated medially 5-8 scales); black markings may
occur midventrally. Large females (lacking enlarged postanal pair of scales)
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Bulletin of the Maryland Herpetological Society
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March 2002
have the same basic ventral body pattern as males (variable in females but
rather distinct dark-bordered blue belly patches), except for lack of black marks
midventrally, and black across throat (diffuse, incomplete).
Males may attain a slightly larger size than females; the largest male is
about 94 mm SVL (UBIPRO 8559), female 91 mm SVL (UBIPRO 8560). There
are usually two canthals, one loreal, and one preocular (some minor varia¬
tion), and some head scale anomalies (e.g., left prefrontal and anterior frontal
fused, UBIPRO 4504; the anterior frontal one-half divided, UBIPRO 5008).
Dorsal scales average 34.3 (31-37, n - 31), scales around midbody 38.8 (34-43,
n = 31), and femoral pores 13.5 (11-15, n = 62, one leg) or 26.9 (23-30, n = 31,
both legs); the pore series are separated medially by about 11 scales (10.8, 9-13,
n - 31). The anterior frontal is entire (61%, n = 31), and there is usually one
frontoparietal (82%, n = 62, both sides of head) separated by an azygous scale
(frontoparietals in medial contact in 5, separated [no azygous scale] in 2 of 62).
Distribution. Sceloporus m. olsoni occurs in the northwestern part of the
range of the species in the Mexican state of Mexico (along Hwy 57); see map.
Fig. 2. Lizards from the type locality were in rock crevices in a low density
oak forest. The general habitat (JLE) is a semiarid, flat area with rocky hills
and many oaks, Acacia, and Prosopis ; the elevational range is 2533-2646 m
(8308-8679 ft, four localities). Sceloporus torquatus is sympatric (syntopic) with
S. m. olsoni; S. torquatus prefers rock fences and large boulders, whereas S. m.
olsoni occurs in cracks of small rocks near the ground (JLE).
Description of Sceloporus mucronatus
Sceloporus mucronatus is of moderate size with males slightly larger than
females; Smith et al. (1952) recorded the maximal size (presumably male) as
114 mm SVL. The supraoculars are divided, prefrontals usually in medial
contact, anterior frontal usually entire (not longitudinally divided),
frontoparietals usually 1-1 separated by an azygous scale, and canthals usu¬
ally 2-2, loreals 1-1, and preoculars 1-1; the anteriormost sublabial (labiomen¬
tal) rarely contacts the mental. The femoral pore series are widely separated
medially (average about 11-12 scales). Black collars are distinct, unbroken,
with the pale borders usually about two scales wide (rarely one) and entire
(both or only the anterior border may be narrowly interrupted medially).
Blue belly patches are rather widely separated (about 6-8 scales at midbody),
with the largest adult males having solid blue throats and black in chest re¬
gion and midventrally.
Bulletin of the Maryland Herpetological Society
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Volume 38 Number 1
March 2002
20-
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Fig. 2. Map of Mexico City (star) area and adjacent states showing lo¬
calities for Sceloporus mucronatus olsoni (solid circles), presumed intergrades,
S. m. olsoni x S. m. mucronatus (half-solid circles — SW Huichipan and NW
Pachuca, Hgo, and Jilotepec, Mex), and our large sample of S. m. mucronatus
(open circle — Km 19, Hwy 894, Mex). Alvarez and Huerta (1973) mapped the
Pachuca and Jilotepec sites (see text) and other known localities for S.
mucronatus.
Numbers of dorsal scales and femoral pores, and aspects of dorsal and
ventral patterns have some taxonomic utility. Dorsal body patterns consist of
either a black vertebral area /blotches, or black-edged scales /longitudinal lines
on the body. A pale intertympanic band (transverse series of pale scales be¬
tween ear openings; postoccipital band of Olson, 1998) is present or absent.
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Bulletin of the Maryland Herpetological Society
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March 2002
and the black collar varies in width (with or without an enclosed pale scale
above shoulder). Tails (usually immaculate ventrally-distally) have distinct
or indistinct black (widest) and whitish (one or two scales wide) banded pat¬
terns. The juvenile throat pattern (usually dimly barred or mottled) with a
pale, dark-bordered, median streak (= parallel, longitudinal dark lines on
middle of throat in Smith, 1942:357) is present or absent. Adult females (no
enlarged pair of postanal scales) may have either a whitish venter with only
a pale blue suffusion, or rather distinct blue, black-bordered belly patches
(but little or no dark pigment on chest or mid ventr ally).
Three subspecies of Sceloporus mucronatus have been recognized, two
(: mucronatus and omiltemanus) by Smith ("1936"[1938], 1939) and a third
(aureolus) later described by Smith (1942). Alvarez and Huerta (1973) recog¬
nized these three subspecies in their review of S. mucronatus . Individuals gen¬
erally occur on rocks or logs at relatively high elevations in oak or pine-oak
forests. Data for these three subspecies, derived from some additional speci¬
mens and published reports, are discussed below.
Sceloporus m. mucronatus. Our material consists of 64 specimens (UBIPRO
5, 7-8, 30, 33-34, 39, 41, 44, 151, 163, 169, 232, 285-86, 305-06, 2237-38, 2240,
2244-45, 2250, 2254, 2256-57; UCM 61093-114, 61116; UTEP 18574-88) all from
one locality, Edo. de Mexico, Km 19 on Hwy 894 (between Ajusco and
Tianguistenco), 3400 m. The cold-weather collection site is a pine forest ( Pinus
montezumae and P. hartwegii) with some grassland (Festuca and Muhlenbergia);
lizards were taken in an open grassy area among basaltic rocks (JLE),
Heads are dark grayish to blackish, unpatterned. The pale intertympanic
band is usually indistinct (distinct spots [UCM 61100] or small dots [UTEP
18576]) but may be absent. Black collars are about four scales wide and usu¬
ally lack pale scales above the shoulder (distinct in UCM 61093, indistinct
both sides in UBIPRO 5, 41). The dorsal pattern consists of a black vertebral
area (several discrete dark spots, UCM 61107, 73 mm SVL; four dark
crossbands separated by whitish marks, UTEP 18582, 62 mm SVL, both fe¬
males) that is mostly continuous but with some scattered white spots-marks
transversely oriented to form black blotches; the black vertebral area is faded,
indistinct in a large male (UCM 61094, 98 mm SVL). Pale brownish sides of
the body are mostly unmarked or with some slightly paler scales. Tails are
moderately banded (rather distinct in UBIPRO 30, 33, 39; UCM 61096). Juve¬
nile throat patterns (pale medial streak) are present (diffuse or absent in adults).
Adult females (no enlarged postanal scales) may have moderately distinct
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Volume 38 Number 1
March 2002
belly patches and mostly uniform dark throats (e.g., UTEP 18583, 91 mm SVL;
UTEP 18584, 72 mm SVL; UBIPRO 2254, 84 mm SVL; UTEP 18586-87, 83 and
76 mm SVL) and may have some black on chest (UBIPRO 2254, UTEP 18583).
Large adult males have solid blue throats, with black complete across the
throat; chest and midventral areas may be uniformly dark gray to almost
black (UBIPRO 41, UTEP 18578, ca 100 and 95 mm SVL). Dorsal scales aver¬
age 30.0 (27-34, n = 63), midbody scales 40.0 (37-43), and femoral pores 12.0
(10-15, n = 116, one leg) or 24.0 (20-29, n - 55, both legs).
Smith's "Diagnosis" of S. m. mucronatus (//1936,,[1938]:583-584, and Fig.
10 [side and dorsal head scalation]) is repeated (only slight changes) in Smith
(1939:218-220, and Fig. 26). Smith (//1936,,[1938]:P1. XLIX, Fig. 2) illustrated
the dorsal pattern of a Veracruz female. Smith (1942) noted dorsal scales aver¬
aging 29.6 (27-32, n = 49, 93.6% 31 or less) and femoral pores 12.8 (10-17, n =
94). Smith and Laufe (1945) recorded femoral pores averaging 12.0 (10-14, n =
35). Smith et al. (1952, vicinity Las Vigas, Veracruz) noted 10-13 femoral pores
per side, dorsal scales 27-29, and black collars covering about 3-4 scales.
Alvarez and Huerta (1973, Table 1, combined data for seven localities)
noted dorsal scale averages of 29.3-32.4 (27-38) and femoral pores 10.9-15.4
(9-16). However, two of their samples may reflect intergradation with S. m.
olsoni (see below); excluding these two samples, five of their samples of S. m.
mucronatus have combined dorsal scale averages of 29.3-31.7 (27-34), but the
same variation in femoral pores. Thus S. m. mucronatus (above data combined
and including the five samples of Alvarez and Huerta) has larger dorsal scales
(averaging about 29-31, 27-34) than S. m. olsoni (34.3, 31-37).
Two samples of Alvarez and Huerta (1973, Table 1), Pachuca, Hidalgo,
and Xilotepec [= Jilotepec], Mexico (combined femoral pores 11.9-12.1, 11-
15), have the highest counts and averages of dorsal scales (respectively, 32.4,
29-38, n = 20 and 32.4, 29-35, n ~ 11), and are perhaps intergradient. The
Pachuca and Jilotepec samples are geographically nearest to the olsoni locali¬
ties; the former site (recorded as 14 or 15 km W Pachuca, Alvarez and Huerta,
1973:180, 182, and presumably along Highway 85 northwest of city) is some
70 air kilometers distant, whereas the latter site (= Xilotepec or Xlotepec in
Alvarez and Huerta, 1973:179, 182) is only some 10-20 air kilometers south-
southeast, of the olsoni sites along Hwy 57. Six additional specimens from
Edo. de Hidalgo are judged to be intergradient between S. m. olsoni and S. m.
mucronatus (REO 5862-64, 5874, 6.7 mi SW Huichapan, Olson, 1998, and UTEP
4580-81, 10.9 road mi [Hwy 45] E Hidalgo-Queretaro state line [ca 6 mi W
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March 2002
Huichapan], 2103 m, Webb, 1982); these Hidalgo specimens (REO, UTEP)
have small dorsal scales (35.0, 34-36, as in olsoni; Olson, 1998:79, Table 1, re¬
corded 32-39), femoral pores averaging 11.2 (9-13), and the dorsal patterns of
two adult females (REO 5862-63) are most like S. m. mucronatus (as illustrated
in PL XLIX, Fig. 2 in Smith, "1936"[1938]). REO 5874, 83 mm SVL, has a dis¬
tinct intertympanic band (also REO 5862), pale scales in black collar above
shoulder, and a faded dorsal pattern (only indication of anteriormost black
vertebral blotch).
Thus, S. m. mucronatus and S. m. olsoni are similar in the vertebral
blotched dorsal pattern and some other pattern features (see Key). The two
taxa seem to show intergradation, have similar femoral pore counts, but dif¬
fer in average size of dorsal scales and aspects of dorsal pattern (see Com¬
parisons and Key).
Sceloporus m. aureolus. Smith (1942:357) in his description of S. m. aureolus
noted that he had ''described this subspecies in detail as mucronatus
omiltemanus Except for the holotype and 26 topotypes (USNM 112232 and
112233-58) all other listed paratypes of S . m. aureolus (Smith, 1942:356) were
previously assigned to S. m. omiltemanus (Smith, "1936//[1938]:596/ including
description, p. 591, Fig. 12 [side and dorsal head scalation] of EHT-HMS 3080,
and the Veracruz specimen depicted in PL L, Fig. 1). His "Diagnosis"
(//1936//[1938]:591 and Fig. 12) of S. m. omiltemanus was repeated (some
changes) in Smith (1939:220-221, and Fig. 28).
Dorsal body patterns have black-edged scales and black longitudinal
lines (adults). Smith ("1936"[1938]:594, as S. m. omiltemanus) noted a pattern
of light lines (middle of each scale row) separated by narrow black lines. The
pale intertympanic band is absent. The juvenile throat pattern (whitish me¬
dial streak) is lacking. Tails have indistinct pale and dark distal bands. Large
adult males have uniformly blue throats with the chest and median abdomi¬
nal region suffused with slate, and blackish near lateral blue belly patches;
females usually have a whitish ventral surface often with a pale blue suffu¬
sion (Smith, "1936" [1938] :594, 596, as S. m. omiltemanus). Smith (1942) recorded
dorsal scales averaging 34.3 (30-38, n = 59, 91.6% over 31), and femoral pores
14 (11-17, n = 124). Alvarez and Huerta (1973, Jicotlan, Oaxaca [but map-
plotted in Guerrero]) recorded dorsal scales as 34.1 (32-36), and femoral pores
13.0 (12-16).
Our additional material consists of seven specimens from Oaxaca (UTEP
7593-95, 5 km SW Tlaxaico, 2030 m [oak hillside/rock walls]; UTEP 7596, 3
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Volume 38 Number 1
March 2002
km SW Cuquila, 7000 ft+ [oak-pine]; UTEP 7597-99, 7 km S [San Andres]
Chicahuaxtla, 2094 m); improved dirt road mileages (in 1968) between Tlaxiaco
and Cuquila and Chicahuaxtla are respectively 17 and 20 kilometers. Webb
and Baker (1969:148) commented on the Oaxaca specimens. Generally dorsal
body patterns have black-edged scales (more evident in larger lizards) that
may form longitudinal black lines (large male, 96 mm SVL, UTEP 7597); the
only other male (UTEP 7596, 84 mm SVL) has black-edged scales but lacks
longitudinal lines. Heads are dark grayish, patternless, with no pale
intertympanic bands. Black collars are about 5 (4-6) scales wide, usually with
no enclosed pale scales above the shoulder (pale scales indicated in two fe¬
males, UTEP 7593, 7598). In life (2x2 transparencies) dorsal scales were or¬
ange-brown; the borders of the black collar were pale orange-yellow in a fe¬
male (UTEP 7594, 75 mm SVL) but white in a male (UTEP 7596, 84 mm SVL,
also greenish scales on forearms). Throat patterns may show some indistinct
dark barring, but no pale, dark-bordered median streak; one female (UTEP
7595, 67 mm SVL) has the midventral pale streak evident anteriorly. The larg¬
est male (UTEP 7597, 96 mm SVL) with black-bordered, blue belly patches is
mostly dark gray to black in the chest region and in the groins with dark
pigment encroaching onto the preanal area (patchlike); the underside of the
thighs and the midventral belly area are dark, almost black. The ventral pat¬
tern of the smaller male (UTEP 7596, 84 mm SVL, enlarged postanal scales
and hemipenis partly everted) has a uniform pale blue throat but lacks ab¬
dominal belly patches (only pale blue wash as in females). The seven UTEP
specimens from Oaxaca have dorsal scales averaging about 32.7 (32-34), scales
around midbody 36.7 (36-38), and femoral pores 13.0 (12-14, one leg) or 26.0
(25-27, both legs). A female (UTEP 7594) has large (inner row) supraoculars
suggesting one undivided row.
Gehlbach and Collette (1957, as S. m. omiltemanus, near Tlaxiaco, dorsal
patterns not mentioned) noted maximal SVL of males as 101.5 and females
96.5 mm (n = 68); they recorded dorsal scales averaging 33.5 (30-38) and femo¬
ral pores 13.6 (11-16). Lynch and Smith (1965, three from Chicahuaxtla) re¬
corded 32, 33, 33 dorsal scales and 15-15, 15-14, 14-14 femoral pores.
Pending further data Sceloporus mucronatus in the Tlaxiaco-Chicahuaxtla
area, is probably intergradient averaging smaller dorsal scales than in S. m.
omiltemanus , and at least some specimens (Chicahuaxtla male, UTEP 7597,
see above) having a longitudinally black-lined dorsal body pattern (as in
aureolus).
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Bulletin of the Maryland Herpetological Society
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March 2002
Sceloporus m. omiltemanus . Smith (1942:357) noted S. m. omiltemanus as
having fewer dorsal scales (averaging 30, 29-32, 92.3% less than 32) than S. m.
aureolus , and that the latter differs from S. m. omiltemanus in having a lined
dorsal body pattern in adult males; he recorded femoral pores averaging 13.8
(11-16, n = 26), which do not differ appreciably from that of 5. m. aureolus
(four combined samples, see above, averaging 13-14, 11-17). The dorsal pat¬
tern of a Guerrero male is illustrated in Smith (//1936//[1938]:P1. L, Fig. 2).
Alvarez and Huerta (1973, Table 1, dorsal body patterns in adult males not
mentioned) recorded dorsal scales (two samples, respectively, Guerrero and
Sola de Vega, Oaxaca) averaging 30.5 (28-32, n = 7) and 31.6 (28-34, n = 5), and
femoral pores 13.0 (12-16, n = 7) and 13.0 (12-15, n = 5). Another specimen
from Guerrero (UTEF 14522, 3.5 mi SW Omilteme) has 31 dorsal scales, 32
midbody scales, and 13-15 femoral pores.
Comparisons and Key
The foregoing data suggest two distinct groups of subspecies of
Sceloporus mucronatus. One is comprised of the taxa olsoni and mucronatus,
both of which have dorsal body patterns of black vertebral areas that may be
separated (distinctly or indistinctly) into blotches by whitish markings and
share some other pattern features (see Key); adult dorsal patterns of S. m.
olsoni seem to be more pallid and distinct (Fig. 1) than in S. m. mucronatus
(pattern more diffuse) and to differ in having a reddish color in life. Numbers
of femoral pores (one leg) are similar in S. m. olsoni (13.5, 11-15) and S. m.
mucronatus (combined average of three samples, plus five of Alvarez and
Huerta, 12.4 [10.9-15.4], 9-17). The data indicate smaller dorsal scales in olsoni
(34.3, 31-37) than in mucronatus (combined average of two samples, plus five
of Alvarez and Huerta, 30.2 [29.3-31.7], 27-34). The two taxa (our collections)
occupy different habitats with olsoni in semiarid, oak-scrub, and mucronatus
in cold pine forest.
The other distinct populational segment of S. mucronatus comprises
the taxa aureolus and omiltemanus that differ in the dorsal body patterns of
adult males, either black-edged scales (no distinct lines, S. m. omiltemanus) or
dark longitudinal lines (S. m. aureolus ), plus some other pattern features (see
Key). Femoral pores (one leg) overall are similar in S. m. omiltemanus (aver¬
age of three samples, 13.3 [13.0-13.8], 11-16) and aureolus (four samples, 13.4
[13.0-14], 11-17). Dorsal scale counts of omiltemanus (average of three samples,
30.7 [30.0-31.6], 28-34) are lower than in aureolus (four samples, 33.6 [32.5-
34.3], 30-38). Traditionally, the two subspecies have been distinguished by
Bulletin of the Maryland Herpetological Society
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relative size of dorsal scales, 31 or less in omiltemanus and 32 or more in aureolus
(Smith and Taylor, 1950:123).
The two groups of subspecies of Sceloponis mucronatiis (olsoni-mucronatus
and aureolus-omiltemanus) are distinctive in dorsal body patterns; intermedi¬
ate morphological variants between adjacent subspecies of each group,
mucronatus and aureolus , are unknown; the two groups of subspecies possibly
represent two species. Some of the other recorded pattern features distin¬
guishing the two groups (in Key, couplet 1), however, are not totally mutu¬
ally exclusive.
Key to Subspecies of Sceloporus mucronatus
la. Dorsal body pattern with black scale edges and/or narrow, black,
longitudinal lines; no pale intertympanic band; black collars often
5 (4-6) scales wide with no included pale scale above the shoulder;
tails indistinctly banded; juvenile throat pattern lacking pale, dark-
bordered, longitudinal median streak; females lacking well-devel¬
oped blue belly patches; femoral pores averaging 13-14 (11-17, one
leg) . 2
lb. Dorsal body pattern with either black mostly continuous vertebral
area (blotches indistinct) or distinct black vertebral blotches; usu¬
ally some evidence of pale intertympanic band; black collars nar¬
row, no more than 4 scales wide, often with pale scale above shoul¬
der; tails may be distinctly banded; juvenile throat pattern with
longitudinal pale median streak; females with distinct blue belly
patches; femoral pores averaging 12-13 (9-17, one leg) . . 3
2a. Back of adult males with black longitudinal lines along scale edges;
dorsal scales averaging about 33-34 (30-38). S. m. aureolus.
2b. Back of adult males having black scale edges but lacking distinct
longitudinal lines; dorsal scales averaging 30-32 (28-34). S. m.
omiltemanus.
3a. Dorsal scales averaging about 29-31 (27-34); adults with indistinct
dorsal blotched pattern and not reddish in life. S. m. mucronatus.
3b. Dorsal scales averaging 34.3 (31-37); adults with distinct blotched
pattern and reddish in life. S. m. olsoni.
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Acknowledgments
This study was supported by a grant to JLE by the Comision Nacional
para el Conocimiento y Uso de la Bioversidad, under project CQNABIO-U003.
We thank R. Earl Olson for the loan of specimens, and William P. MacKay
(UTEP) for aid in readying Figs. 1 and 2 for publication.
Literature Cited
Alvarez, T., and R Huerta.
1973. Notas sobre Sceloporus mucrunatus [sic] (Reptilia: Iguanidae)
en Mexico. An. Esc. nac. Cienc. biol, Mex., 20:177-184.
Auth, D.L., H.M. Smith, B.C. Brown, and D. Lintz.
2000. A description of the Mexican amphibian and reptile collec¬
tion of the Strecker Museum. Bull Chicago Herp. Soc., 35:65-
85.
Gehlbach, F.R., and B.B. Collette.
1957. A contribution to the herpetofauna of the highlands of
Oaxaca and Puebla, Mexico. Herpetologica, 13:227-231.
Lynch, J.D., and H.M. Smith.
1965. New or unusual amphibians and reptiles from Oaxaca,
Mexico. I. Herpetologica, 21:168-177.
Mindell, D.R, J.W. Sites, Jr., and D. Graur.
1989. Speeiational evolution: a phylogenetic test with allozymes
in Sceloporus (Reptilia). Cladistics, 5:49-61.
Olson, R.E.
1998. Sceloporus poinsetti: its taxonomic affinity with mucronatus.
Bull Maryland Herpetol. Soc., 34:76-82.
Smith, H.M.
"1936"[1938]. The lizards of the torquatus group of the genus Sceloporus
Wiegmann, 1828. Univ. Kansas Sci. Bull, 24:539-693 (pub¬
lished 16 February 1938).
1939. The Mexican and Central American lizards of the genus
Sceloporus . Field Mus. Nat. Hist., Zool Ser., 26:1-397, 31 pis.
Bulletin of the Maryland Herpetological Society
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Volume 38 Number 1
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1942. Mexican herpetological miscellany. Proc. U.S. Nat. Mus.,
92:349-395.
_ ., and L.E. Laufe.
1945. Mexican amphibians and reptiles in the Texas Cooperative
Wildlife collections. Trans. Kansas Acad. Sci., 48:325-354.
and E.H. Taylor.
1950. An annotated checklist and key to the reptiles of Mexico
exclusive of the snakes. Bull. U.S. Nat. Mus. (199):v+253.
Smith, P.W., H.M. Smith, and J.E. Werler.
1952. Notes on a collection of amphibians and reptiles from east¬
ern Mexico. Texas }. Sci., 4:251-260.
Webb, R. G.
1982. Distributional records for Mexican reptiles. Herpetol. Rev.,
13:132.
_ ., and R.H. Baker.
1969. Vertebrados terrestres del suroeste de Oaxaca. An. Inst. Biol.
Univ. Nal. Auton. Mex., 40, Ser. Zool. (1):139-152.
RGW: Department of Biological Sciences , University of Texas at El Paso , El Paso ,
Texas 79968-0519, USA. JLE: Laboratorio de Ecologia, Unidad de Biotecnologia y
Prototipos, Escuela Nacional de Estudios Profesionales Iztacala, UNAM,
Tlalnepantla, Mexico, 54090, Mexico: HMS: Department EPO Biology, Univer¬
sity of Colorado, Boulder, Colorado 80309-8334, USA.
Received: 20 October 2001
Accepted: 17 November 2001
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Observations on the Diet of Trachemys gaigeae
(Testudines: Emydidae)
James N. Stuart and Charles W. Painter
The Big Bend slider, Trachemys gaigeae , is a freshwater turtle of the Rio
Grande drainage system of southern New Mexico, western Texas, eastern
and central Chihuahua, and Coahuila (Ernst, 1992). Most research on T. gaigeae
has focused on its systematic and taxonomic relationship to other sliders in
the T. scripta complex (e.g., Ernst, 1992; Seidel et ah, 1999), while relatively
little attention has been directed to natural history (Ernst et al., 1994;
Degenhardt et al., 1996). In this paper, we review previous publications on
the food habits of T. gaigeae and provide new information obtained by several
methods, including 1) trapping results using various trap baits, 2) fecal sample
analysis, 3) dissection of freshly-killed specimens, 4) stomach-flushing, and
5) observations of specimens feeding in the wild and in captivity.
Materials and methods
Our studies of T. gaigeae were primarily conducted during 1994-1998 at
Bosque del Apache National Wildlife Refuge (BDA) and Elephant Butte Res¬
ervoir (EBR) in the Rio Grande Valley, Socorro and Sierra counties. New
Mexico. Adult and subadult turtles were typically captured in nylon-mesh
hoop-traps (130 cm long, 75 cm diameter) placed in lentic waters > 70 cm
deep. Traps were baited with sardines, watermelon, or banana contained in a
perforated can or wire-mesh bag to prevent consumption by captured turtles.
Turtles less than 80 mm maximum straight-line carapace length (CL) were
too small to be captured in the traps. Captives were retained for ca. 24 h for
measurement and collection of fecal samples which were preserved in 10%
formalin for later examination under a dissecting microscope. Most speci¬
mens were uniquely marked by shell notching and released at the capture
site. Several egg clutches were obtained from wild-caught gravid females
and incubated in the laboratory, and hatchlings were captive-reared for up to
12 months under laboratory conditions in aquaria and, during warmer months
of the year, in outdoor metal tanks (ca. 1.5-2. 2 m diam.). Concurrent with
field studies, several adults were maintained for observation in outdoor metal
tanks and a permanent outdoor pond (6.1 m diam.) on the University of New
Mexico campus. Our identification and taxonomy of aquatic macrophytes in
the study area follows Adams (1998).
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Previous reports
Several publications briefly discuss the food habits of T. gaigeae. Carr
(1952) speculated that the species must be largely carnivorous based on the
riverine habitat in Texas where it is found, whereas Legler (1960b) noted that
the stomach contents of several T. gaigeae from the Rio Conchos, Chihuahua,
consisted of only aquatic vegetation. Price and Hillis (1989) suggested that
adult T. gaigeae, unlike T. scripta in Texas, are exclusively vegetarian. Ernst et
al. (1994) noted that captives readily accepted fish. At BDA in New Mexico,
Stuart (1995) noted the abundance of submerged aquatic macrophytes at cap¬
ture sites, including pondweeds (Fotamogeton pectinatus and Zannichellia sp.),
and filamentous green algae (Chlorophyta), which were presumed food
sources. Wilson et al. (1999) and Morjan and Stuart (2001) both reported fila¬
mentous green algae, fragments of aquatic vascular plants (primarily
Fotamogeton sp.), and parts of crayfish ( Orconectes sp.) from the gastrointesti¬
nal tract and feces of two adult females (182 mm CL and 253.5 mm CL, re¬
spectively) from BDA.
In a study indirectly relevant to T. gaigeae food habits, Garcia (1973)
examined the concentration of methyl mercury in body tissues of "Pseudemys
scripta " (= T. gaigeae), Apalone spinifera, and numerous fish species in EBR, a
major sediment trap on the Rio Grande. Mercury concentrations in liver and
kidney tissues of both turtle species were significantly higher than those in
fishes, whereas overall mercury concentrations were similar in turtles and
predatory fish species, indicating possible bioamplification. Garcia (1973)
suggested the carnivorous habits of both turtles, including scavenging of dead
fishes, could account for high mercury concentrations in these species (me¬
thyl mercury can occur at higher concentrations in dead, decaying fish tissue
than in fresh fish). The study found that mercury concentrations in mixed
algae and bryophytes in the reservoir were also elevated.
Attraction to baited traps
Legler (1960a; 1960b) noted that T. gaigeae, which he considered to be
chiefly herbivorous, was attracted to meat or fish baits. In our study, perfo¬
rated cans of sardines in oil or tomato sauce proved to be an effective attrac-
tant for T. gaigeae for trapping periods of up to 4 days. Sliced watermelon and
banana wrapped in small-mesh screen material was used to a lesser extent as
bait; this method also attracted T. gaigeae, albeit for only the first 24 hrs of
trapping.
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Freshwater fishes (e.g., Cyprinus carpio, Ictalurus sp., Micropterus sp.)
taken as bycatch in traps with T. gaigeae were occasionally sacrificed and left
in traps as an additional attractant for turtles. These fish carcasses were often
found partially consumed during subsequent trap checks, presumably by
the turtles occupying the traps.
Fecal sample analysis
Newly-captured adults and subadults taken at BDA during June- Au¬
gust typically defecated large quantities of loosely-consolidated vegetable
material. Casual examination of this material indicated the presence of aquatic
vascular plants and filamentous algae. One male (167 mm CL) had ingested
parts of a grasshopper (Orthoptera) along with filamentous algae.
A more detailed examination of preserved fecal samples from 14 adults
(160-224 mm CL) captured at BDA in 1994 indicated the following items as
the major foods: fragments of unidentified vascular plants (6 samples); the
filamentous alga Oedogonium (4 samples), a filamentous alga similar to
Rhizoclonium (2 samples); and two unidentified filamentous algae (1 sample).
An unidentified pollen was the sole item in 1 sample. Secondary food items
in these 14 samples included the following: diatoms (i.e., Gomphonema,
Cocconeis, and Epithemia ; presumably epiphytic on the filamentous algae), the
protozoan Vorticella , a desmid alga ( Pediastrum sp.), unidentified coccoid green
algae, and an unidentified fungus. One adult collected from the Rio Grande
near Las Palomas, Sierra County appeared to have pieces of muskgrass ( Chara
vulgaris) in its feces; this alga was observed growing near the capture site. A
fecal sample from a female (173.5 mm CL) trapped near Las Palomas on 13
August 1993 consisted of unidentified vascular plant material and diatoms.
Terrestrial vegetation was also detected in some fecal samples. Three
adult T. gaigeae (140-221 mm CL) collected from the Rio Grande in Brewster
County, Texas in April 1997 and reported by Seidel et al. (1997) were tempo¬
rarily held in captivity before being sacrificed. Feces from these turtles con¬
sisted almost exclusively of fragments of giant reed (. Arundo donax). This grass
was growing along the shoreline in the vicinity of trap sites (W.G. Degenhardt,
pers. comm.). During 19-20 May 1998 in a cove at EBR, we trapped several
large adults that defecated fragments of an unidentified grass. Water depth
at trap sites was < 1 m, and recently-sprouted grasses and forbs along the
shoreline had been inundated by the rising reservoir level.
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Dissection of preserved specimens
Few specimens of T. gaigeae were sacrificed and dissected during our
study. Wilson et al. (1999) reported food items from one such specimen, which
primarily contained large quantities of filamentous algae and pieces of vas¬
cular plants in its stomach and intestinal tract. Similarly, an adult female (235.5
mm CL) that drowned in a trap at BDA on 2 June 1998 had a large quantity of
filamentous algae and fragments of vascular plants (probably P. pectinatus) in
its stomach; the intestinal tract was empty. A melanistic adult male (210 mm
CL) collected at BDA on 2 August 1993 had only filamentous algae in its in-
testinal tract
Stomach-flushing
Parmenter and Avery (1990) described a method for stomach-flushing
live turtles to obtain food items. We successfully tested a similar device on an
adult female T. gaigeae (238 mm CL) captured at BDA on 31 August 1994. The
stomach was filled with two forms of filamentous green algae: Mougeotia sp.
and Rhizoclonium sp. (or a similar taxon).
Observations in the wild
Opportunities to observe natural feeding by T. gaigeae in New Mexico
were very limited due to the high turbidity of water bodies where the species
was found. On 31 July 1997, we observed a small adult at EBR as it floated
near the water surface and cropped filamentous algae growing on the par¬
tially submerged branch of a dead tree.
Indirect information on diet was provided by observations of aquatic
plants in proximity to trap sites. At BDA, T. gaigeae was most frequently trapped
in ponds that supported dense growths of Fotamogeton pectinatus and floating
mats of filamentous green algae, usually from early June to early September.
At EBR, we trapped T. gaigeae near dense growths of milfoil ( Myriophyllum
verticillatum or M. spicatum) in the littoral zone of deep-water coves;
Potamogeton crispus was also present in lesser quantities in the reservoir.
Observations of captive specimens
Adult and subadult turtles maintained in captivity were offered a vari¬
ety of food items including locally collected aquatic plants {Myriophyllum sp.,
Potamogeton pectinatus , and P. crispus); red-leaf and Romaine lettuce; chopped
uncooked pieces of fish, squid, and chicken; and commercially-available pel-
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let food for fish and reptiles. Observation and examination of feces indicated
that all these items were consumed by captives. On several occasions adults
were observed to feed almost exclusively on fish and meat even when veg¬
etable material was also available. Three adult specimens from Brewster
County, Texas that were briefly held in captivity showed a preference for
lettuce and mostly ignored Myriophyllum that was also available. Feeding
occurred at almost any time of day, including evening. Adults maintained in
the outdoor pond were observed cropping filamentous algae from the wall
of the pond. On one occasion, an adult was observed feeding on a cockroach
(Blatta sp.) that fell into its tank.
Hatchling and small juvenile T. gaigeae (30-40 mm CL) readily fed on
commercially-available frozen brine shrimp (Artemia sp.) and Tubifex worms,
and finely-chopped pieces of raw fish and canned squid. Hatchlings main¬
tained in aquaria showed little interest in commercial turtle pellets, filamen¬
tous algae or other plant materials. Feces from hatchlings maintained in out¬
door tanks were composed primarily of unidentified insect parts, although
small amounts of vegetable matter were also present. Outdoor tanks occu¬
pied by hatchlings were noticeably free of mosquito (Culicidae) larvae as com¬
pared to adjacent tanks that lacked hatchlings. A hatchling placed in a small
water bowl with 15 mosquito larvae consumed all the larvae within 2 h. An¬
other hatchling temporarily placed in a shallow outdoor tank for photographic
purposes was observed gleaning aquatic larvae of an unidentified insect while
sitting motionless on the substrate (D. Sias, pers. comm.).
Comments
Our data indicate that T. gaigeae is omnivorous, but that adults and su¬
badults (i.e., individuals > ca.140 mm CL) are primarily herbivorous. Small
invertebrates may be ingested by adults incidentally with vegetation, whereas
larger organisms (e.g., crayfish) are possibly consumed directly. Dead fish
and crayfish may also be scavenged opportunistically. Hatchlings and small
juveniles (30-40 mm CL) appear to feed primarily on small invertebrates. We
captured few individuals of intermediate size (40-140 mm CL) and can only
infer that a gradual dietary shift from primarily invertebrates to vegetation
occurs during this stage of growth.
Previous dietary studies of Trachemys spp., especially T. scripta (sensu
stricto) in the United States, indicate that sliders are opportunistic omnivores
that consume a wide variety of invertebrates, vertebrates, and vegetation (see
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review by Parmenter and Avery 1990). Adult T. script a (sensu stricto) gener¬
ally consume a much greater percentage of vegetation than do hatchlings
and juveniles, reflecting ontogenetic shifts in energy and nutrient require¬
ments, foraging habits, and microhabitat use (Parmenter and Avery, 1990).
Our evidence suggests that T. gaigeae follows the pattern observed in other
species and populations of Trachemys.
Neustophagia, a type of ingestion in which small food particles float¬
ing on the water's surface (neuston) are skimmed and "filtered" in the turtle's
mouth, has been reported in T. scripta and other emydids (Parmenter and
Avery, 1990). Although we have no direct evidence that neustophagia is used
by T. gaigeae , the presence of only pollen in one fecal sample examined sug¬
gests that it may occasionally be employed by this species.
Although we did not observe fish consumption under natural condi¬
tions, our observations of captive turtles suggest the species readily scav¬
enges dead fish when available. The deepwater habitats adjacent to the lit¬
toral areas frequented by T. gaigeae at EBR support a diversity of large fish
species (Sublette et al., 1990). During trapping efforts at EBR in May 1998, we
observed extensive post-spawning mortality of shad (Dorosoma sp.). We sus¬
pect seasonal fish mortality may provide an important food source for T.
gaigeae at EBR, especially early in the annual activity season when aquatic
vegetation is sparse.
Acknowledgments
We thank Louise Lewis for identifying filamentous algae in our samples,
and Michael Seidel, William Degenhardt, and Don Sias for their loan of live
specimens or observations. Fieldwork was partially funded by the U.S. Fish
and Wildlife Service (FWS) and New Mexico Department of Game and Fish
Share With Wildlife Program. Studies at Bosque del Apache National Wild¬
life Refuge were permitted by the FWS.
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March 2002
Literature Cited
Adams, T.P.
1998.
An inventory of herbaceous vascular aquatic plants of pe¬
rennial waters of southwestern New Mexico. Unpubl. M.
Sci. thesis. New Mexico State Univ., Las Cruces, xii + 164
PP-
Carr, A.F., Jr.
1952.
Handbook of Turtles. The Turtles of the United States,
Canada, and Baja California. Comstock Publ. Assoc., Cornell
Univ. Press, Ithaca. 542 pp.
Degenhardt, W.G., C.W. Painter, and A.H. Price.
1996. Amphibians and Reptiles of New Mexico. Univ. of New
Ernst, C.H.
Mexico Press, Albuquerque, xix + 431 pp.
1992. Trachemys gaigeae . Catalog. Amer. Amphib. Rept. (538):l-4.
Ernst, C.H., J.E. Lovich, and R.W. Barbour.
1994. Turtles of the United States and Canada. Smithsonian Insti-
tution Press, Washington and London, xxxviii + 578 pp.
Garcia, J.D.
1973.
A study of mercurials in the Elephant Butte Reservoir eco¬
system. Unpubl. Ph.D. dissertation, Univ. of New Mexico,
Albuquerque. 128 pp.
Legler, J.M.
1960a. A simple and inexpensive device for trapping aquatic
turtles. Utah Acad. Sci. Proc. 37:63-66.
1960b. Remarks on the natural history of the Big Bend slider,
Pseudemys scripta gaigeae Hartweg. Herpetologica 16:139-140.
Morjan, C.L. and J.N. Stuart.
2001 . Nesting record of a Big Bend slider turtle (Trachemys gaigeae)
in New Mexico, and overwintering of hatchlings in the nest.
Southwest. Nat. 46:230-234.
Bulletin of the Maryland Herpetological Society
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Parmenter, R.R. and H.W. Avery.
1990. The feeding ecology of the slider turtle. Pp. 257-266, in J.W.
Gibbons (ed.), Life History and Ecology of the Slider Turtle.
Smithson. Instit. Press, Washington, D.C. xiv + 368 pp.
Price, A.H. and D.M. Hillis.
1989. Biochemical genetics and taxonomic status of Trachemys
gaigeae and the Trachemys scripta complex in Texas. Abstr.
First World Congress of Herpetology, Canterbury, United
Kingdom.
Seidel, M.E., W.G. Degenhardt, and J.R. Dixon.
1997. Geographic distribution: Trachemys gaigeae (Big Bend slider).
Herpetol. Rev. 28:157.
Seidel, M.E., J.N. Stuart, and W.G. Degenhardt.
1999. Variation and species status of slider turtles (genus
Trachemys) in the southwestern United States and adjacent
Mexico. Herpetologica 55:470-487.
Stuart, J.N.
1995. Notes on aquatic turtles of the Rio Grande drainage. New
Mexico. Bull. Maryland Herpetol. Soc. 31:147-157.
Sublette, J.E., M.D. Hatch, and M. Sublette.
1990. The Fishes of New Mexico. Univ. New Mexico Press, Albu¬
querque. xiii + 393 pp.
Wilson, W.D., J.A. Hnida, and J.N. Stuart.
1999. Natural history note: Trachemys gaigeae (Big Bend slider).
Endoparasites. Herpetol. Rev. 30:226.
New Mexico Department of Game and Fish , Conservation Services Division,
P. O. Box 25112 , Santa Fef NM 87504 , USA
E-mail (JNS): JStuart@state.nm.us
Received 13 December 2001
Accepted 30 December 2001
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Bulletin of the Maryland Herpetological Society
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March 2002
Evolutionary Speciation in the Alligator Lizards of
the Genus Barisia
Hobart M. Smith, Theresa M. Burg and David Chiszar
Abstract.
Barisia imbricata auctorum is a superspecies consisting of four sibling
species: B. ciliaris, B. imbricata , B. jonesi and B. planifrons.
Guillette and Smith (1982) recorded detailed information on external
variation in Barisia imbricata (Wiegmann), recognizing four subspecies: B. i.
imbricata, B. i. ciliaris (Smith), B. i. jonesi Guillette and Smith, and B. i. planifrons
(Bocourt).
Good (1988) accepted that arrangement, as have most other authors.
His review of the Gerrhonotinae also included B. levicollis (Stejneger) and B.
rudicollis (Wiegmann) in the genus Barisia, as generally accepted since Tihen
(1949).
Good (1988) also stated (p. 81) that " Barisia levicollis is probably no more
different from B. imbricata than the various subspecies of B. imbricata are from
each other; either its specific status or the subspecific status of some of the B.
imbricata subspecies is therefore open to question."
On the contrary, five categorical differences distinguish B. levicollis from
the subspecies of B. imbricata: a single superciliary (vs 2-4, except for one
anomalous specimen in 192 with 0); 46-51 dorsals (vs 39-45); postoculars 1-2
(vs 3-4); no preocular contact with anterior superciliary (vs 100%); and
preocular in contact with both anterior medial and lateral supraoculars (vs
0%). The last two, however, are contingent upon the first, but that still leaves
three independent categorical differences of B. levicollis from B. imbricata. In
addition, the two species are broadly dichopatric (Fig. 1).
Thus we regard B. levicollis as unassailably a separate species from B.
imbricata.
Each of the subspecies of B. imbricata also has at least one categorical
distinction from the others. Most notable is the unique occurrence, in the B.
imbricata complex, of 12-14 dorsal scale rows in B. i. imbricata; all other sub-
Bulletin of the Maryland Herpetological Society
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j
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Bulletin of the Maryland Herpetological Society
Fig. 1. Distribution of the five members of the Barisia imbricata complex. C, B. ciliaris; lf B. imbricata; J, B. jonesi ;
B. levicollis; P, B. planifrons .
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March 2002
species, as well as B. kvicollis, have 16. It is also dichopatric from B. i. jonesi
and B. i. planifrons , partially parapatric with and partially dichopatric from B.
l ciliaris . Nevertheless, there is no indication of intergradation between the
latter and B, i. imbricata , despite the parapatry that seems to exist. Although
Fig. 1 indicates partial dichopatry, on the basis of available specimens,
parapatry may well occur there as well as elsewhere. B. i. imbricata occupies a
central position; all of the others are peripheral (Fig. 1).
The most readily distinguishable of the three peripheral subspecies is
B. i. ciliaris , which is light tan at least in adults. In the other two, the ground
color is invariably dark, often with still darker markings dorsally.
A noncategorical but partially diagnostic difference of B. i. ciliaris from
B. i. jonesi and B. i. planifrons is the presence of two loreals in 79% (N=45) of
the former, one in 100% of the two others. B. i. ciliaris reaches a greater size
(158 mm SVL) than the other two (133 mm in B. i. jonesi , 122 mm in B. i.
planifrons ). Obviously there is no intergradation of any, because they are widely
dichopatric.
The light dorsal color of B. i. ciliaris is shared in Barisia only with B.
levicollis . Indeed, examples of the former were long thought to be the latter
because of the similarity in coloration, and the name ciliaris was applied in
reference to the fact that, contrary to B. levicollis , it has a full set of superciliaries.
The two dichopatric populations of B. imbricata (B. i. jonesi , B. i. planifrons)
have a highly restricted range, far from each other. They categorically differ
from each other in at least two ways: B. i. jonesi has sharply keeled dorsals
and the lower anterior temporal is in contact with two supralabials, as op¬
posed to obtuse keels and a single supralabial in contact with the lower ante¬
rior temporal in B. i. planifrons . There is only a slight overlap in dorsal scale
count, 34-39 in the latter, 39-42 in the former.
In view of the existence of categorical difference separating all four sub¬
species of B. imbricata , the absence of evidence of intergradation where
parapatry appears to occur (and the dichopatry of other populations), we
suggest that all four taxa are evolutionarily qualified as species. They are not
as distinct from each other as B. levicollis is from the B. imbricata complex, but
species are highly variable in degree of difference. The four in the B. imbricata
complex may be regarded as sibling species (e.g. Mayr and Ashlock, 1991),
which are less readily distinguished from each other than the members of the
B. imbricata complex are from the rest of the species of Barisia. We therefore
Bulletin of the Maryland Herpetological Society
page 25
Volume 38 Number 1
March 2002
propose that the taxa conventionally regarded as subspecies of B. imbricata be
known as B. ciliaris , B. imbricata , B. jonesi and B. planifrons.
Literature Cited
Good, D. A.
1988. Phylogenetic relationships among gerrhonotine lizards.
Univ. California Publ. Zook 121: i-x, 1-139.
Guillette, L. J. Jr. and H. M. Smith.
1982. A review of the Mexican lizard Barisia imbricata , and the
description of a new subspecies. Trans. Kansas Acad. Sci.
85: 13-33.
Mayr, E. and R D. Ashlock.
1991 . Principles of systematic zoology Second edition. New York,
McGraw-Hill, xx, 475 pp.
Tihen, J. A.
1949. A review of the lizard genus Barisia. Kansas Univ. Sci. Bull.
33: 217-255.
HMS , DC: University of Colorado Museum , Boulder, Colorado 80309-0334 .
TB : NMFS Santa Cruz Laboratory, 110 Shaffer Road , Santa Cruz ,
California 95060 .
Received: 18 December 2001
Accepted: 30 December 2001
page 26
Bulletin of the Maryland Herpetological Society
Volume 38 Number 1
March 2002
A New Species of Conophis (Reptilia: Serpentes)
From Los Tuxtlas, an Area of High Endemism in
Southern Veracruz, Mexico
Gonzalo Perez-Higareda, Marco A. Lopez-Luna and Hobart M. Smith
Abstract
Conophis morai is described from a unique habitat for the genus in the
elevated rain forest of Los Tuxtlas, southern Veracruz, south of the range of
C. lineatus, here regarded as a species distinct from C. concolor.
The only Conophis known to occur on Atlantic slopes of Mexico outside
of the Yucatan Peninsula, where C. I concolor occurs, is C. /. lineatus (Dumeril,
Bibron and Dumeril), occurring on the coastal plains and lowlands of central
Veracruz as far south as Lerdo de Tejada (Wellman, 1963; Perez-Higareda
and Smith, 1991), in grasslands and deciduous forest areas.
However, one Conophis was taken by Roberto Mora 12 April 1999 in a
very different, elevated habitat in rain forest on the southeastern slope of San
Martin Tuxtla Volcano, some 60 km southeast of Lerdo de Tejada. We are not
aware of any other voucher specimen taken in Veracruz in a similar habitat,
or that far from the known range of C. lineatus.
The specimen exhibits numerous differences from typical specimens of
C. lineatus in coloration and pattern. In view of these differences, correlated
with distance from other Conophis records, from a unique environment given
to a high degree of endemism, we regard it as representative of a distinct
species here named
Conophis morai sp. nov.
Holotype. UNAM-LT 3662, an adult male, from Ejido Ruiz Cortines on
the southeastern slope of San Martin Tuxtla Volcano, 1050 m, taken by Roberto
Mora 12 April 1999.
Diagnosis. Different from C. lineatus, the only other striped member of
the genus on the east coast of Mexico, in having the dorsolateral and lateral
dark stripes continuous and split by a white line throughout the length of
body, including neck, occupying two scale rows; sublateral stripe continu¬
ous, uninterrupted except on neck; spot at ends of ventrals sharply defined.
Bulletin of the Maryland Herpetological Society
page 27
Volume 38 Number 1
March 2002
not diffuse; both dorsal and ventral edges of supralabials black; ventral sur¬
face of head mostly black, from mental to the first few ventrals; posterior
edge of each ventral on anterior part of body black; rest of venter gray, with
several irregular black spots on midbody ventrals.
Description ofholotype. Scalation much as in other members of Conophis ,
none of which are distinguishable by morphological characters alone. Head
scales normal: 7-8 supralabials, 3rd and 4th entering orbit on the side with 7,
4th and 5th on the other; 9-9 infralabials; 1-1 preoculars; 1-1 loreals; 2-2
postoculars; 2-2 temporals; 19-19-17 scale rows; no keels; 165 ventrals;
subcaudals 66; 664 mm total length. Condition excellent except left side of
head damaged.
Dorsal ground color greenish gray in life, gray in preservative. Dorso¬
lateral stripes broad throughout, beginning on edges of prefrontal and fron¬
tal, on the neck each forming a pair of continuous black lines separated by a
narrow white line throughout the length of the body, on the 7th and 8th scale
rows anteriorly, 6th and 7th posteriorly (Fig. 1). Lateral line broad through¬
out, beginning on rostral, passing through eye, covering the upper edges of
the supralabials (Fig. 3), and splitting into two continuous black lines sepa¬
rated by a white line five scales behind the head, extending throughout the
length of body on 3rd and 4th rows anteriorly, 2nd and 3rd posteriorly.
A sublateral dark line begins on neck as a series of spots on the 1st scale
row, but posterior to the 10th ventral forming a single, continuous black line,
similar to the others, on 1st dorsal scale row throughout the length of the
body, but absent on tail.
A sharply defined black spot on each end of each ventral; ventral sur¬
face of head dark from mental to first few ventrals (Fig. 3). On each side of
each ventral a distinct, sharply defined, oval dark spot, not diffuse or angular
in shape (Fig. 1). The posterior edge on each of the first 20 ventrals is black
(Fig. 4). The rest of the venter is white, except at midbody (Fig. 4), with sev¬
eral irregular black spots on the ventrals, varying in size from almost an en¬
tire ventral to small ones no larger than the regular spot on each end of each
ventral.
Supralabials with black dorsal and ventral edges bordering a continu¬
ous white line along the middle of the scales. Ventral surface of head largely
black, from mental to first few ventrals (Fig. 3).
page 28
Bulletin of the Maryland Herpetological Society
Volume 38 Number 1
March 2002
Fig. 1. Midbody pattern of the
holotype of C. morai.
Fig. 3. Head pattern of the ho¬
lotype of C. morai. Above, ventral
view; below, lateral view.
Fig. 2. Midbody pattern of C.
lineatus, Cotaxtla, Veracruz, from
UNAM-LT 3812.
Fig. 4. Ventral pattern on body.
Left, neck region at ventrals 15-23;
right, midabdomen at ventrals 49-57.
Other irregular dark blotches appear
between ventrals 58-65, one on ven¬
tral 86, and others on ventrals 93-96,
the latter small and diffuse. The rest
of the ventrals are unmarked except
for the dark spot on each end of each
ventral; they continue onto the basal
15 subcaudals, but otherwise the ven¬
ter and subcaudal surfaces are white.
Bulletin of the Maryland Herpetological Society
page 29
Volume 38 Number 1
March 2002
Comparisons. C. maria differs from the nearest taxon of the genus, C.
lineatus , as follows. Character-states for C. 1. lineatus are largely based on
Wellman (1963); GPH also examined a few specimens in Mexican collections.
(1) . Ground color greenish gray, gray in preservative (vs "white, tan-
nish white, or often pale blue" in preservative [Wellman, 1963]).
(2) . Sharply defined, continuous dorsolateral and lateral stripes, broad
on head, on body split into two on adjacent scale rows, enclosing a white line
between, extending the full length of the body (vs generally on a single scale
row, anteriorly often a series of spots, in some expanded only posteriorly
onto the adjacent scale row, but accessory lines discontinuous where present
and light area between gray, not white).
(3) . Sublateral stripe sharply defined and continuous throughout length
of body, on neck appearing as a series of spots, one on each scale of first
dorsal scale row; scale row between sublateral and lateral stripes white (vs a
series of dark-centered scales, in some forming short continuous sections,
often scarcely visible; area between sublateral and lateral stripes gray, not
white).
(4) . Supralabials black dorsally and ventrally, with a white line between
(vs black ventrally only).
(5) . Ventral surface of head largely black (vs unmarked or with a few
small, scattered black flecks).
(6) . Black spot on each end of each ventral sharply defined, oval (vs
diffuse, angular).
(7) . Venter white, anterior ventrals black-edged, midbody ones with
irregular black blotches (vs white, no markings except for spots on ends of
ventrals).
The only other Conophis known from Atlantic slopes of Mexico is what
Wellman (1963) designated C. lineatus concolor. As represented in Mexico, it is
readily distinguished by its unicolor dorsum, lacking stripes. It is known no
closer to the Los Tuxtlas area than Campeche.
Etymology. The patronym honors Biologist Roberto Mora, collector and
donor of the holotype.
page 30
Bulletin of the Maryland Herpetological Society
Volume 38 Number 1
March 2002
Comments. In view of (1) the many unique features of the holotype of C
moral, compared with the adjacent C. lineatus (see preceding), and even all of
the other taxa of the genus (most notably the head and ventral markings),
and (2) occurrence in a sharply different habitat from that characteristic of
the genus, in a dichopatric area of considerable endemism, C. moral is reason¬
ably assured as a distinct species. Further voucher examples will be required
for verification.
No specimens of Conophis are known between the ranges of C. lineatus
and C. morai to the north and the nearest other population of the genus, in
Campeche, northern Yucatan and Belize. A hiatus of some 500 km separates
them. A sharp difference exists between the northern populations and the
cited southern populations; without exception the former is lined, whereas
the southern one is lineless. Farther southeast the unicolor populations blend
with lined ones, and there is no geographic hiatus between them. We there¬
fore propose that C. lineatus is a monotypic species, limited to central Veracruz.
The southern group consists of geographic and/or populational variants of
apparently one species, C concolor ; including the nominal taxa C. I dunni and
C. pulcher,fide Wilson and Meyer (1985), Lee (1996), and Campbell (1998).
One other specimen, an adult male presumably of C. morai , was caught
by GFH September 1981, retained briefly, and later escaped before detailed
observations were made. Field notes indicate that its ground color was gray-
olive. It was found in rain forest at 200 m, San Pabla Siqueda, Balzapote,
municipality of Ssn Andres Tuxtla, Veracruz.
Acknowledgment.
All drawings are by Marcos A. Lopez-Luna.
Bulletin of the Maryland Herpetological Society
page 31
Volume 38 Number 1
March 2002
Literature Cited.
Campbell, J. A.
1998. Amphibians and reptiles of northern Guatemala, the
Yucatan , and Belize. Norman, Oklahoma, Univ. Oklahoma
press, xiii, 380 pp.
Lee, J. C.
1996. The amphibians and reptiles of the Yucatan Peninsula.
Ithaca, New York, Cornell Univ. Press, xii, 500 pp.
Perez-Higareda, G. and H. M. Smith.
1991 . Ofidiofauna de Veracruz: analysis taxonomico y geografico.
Publ. Esp. Instituto de Biologia, UNAM 7: 1-122.
Wellman, J.
1963. A revision of snakes of the genus Conophis (family
Colubridae), from Middle America. Univ. Kansas Publ. Mus.
Nat. Hist. 15: 251-295.
Wilson, L. D. and J. R. Meyer.
1985. The snakes of Honduras. Second edition. Milwaukee, Wis¬
consin, Milwaukee Public Mus. x, 150 pp.
GPH : Estacidn de Biologia Tropical " Los Tuxtlas ," UNAM , Apartado Postal 51,
Catemaco , Veracruz , Mexico.
MALL: Centro de Investigaciones Herpetologicas de Veracruz A. C., Apartado
Postal 473 , Cordoba , Veracruz, Mexico .
HMS: Department ofEPO Biology and Museum, University of Colorado, Boulder,
Colorado 80309-0334 U.S.A.
Received: 18 December 2001
Accepted: 30 December 2001
page 32
Bulletin of the Maryland Herpetological Society
Volume 38 Number 1
March 2002
News and Notes
Reptile and
Amphibian Rescue
410-580-0250
We will take as many un
amvhibians as
with your name and number to
give up an animal for adoption;
or to volunteer to help with our efforts.
OUR CURRENT NEEDS:
• Commercial or Passenger Van
• UV Lights • Power & Hand Tools
• Equipment & Food • Paper Towels
www.reptileinfo.com
Bulletin of the Maryland Herpetological Society
page 33
Volume 38 Number 1
March 2002
News and Notes
TKu-
^EIPTDIUi
S M<Q>
Presented by MARS Preservation Fund, Inc.
We believe in education,
the promotion of captive
breeding and the protection
of critical habitats.
Maryland State Fairgrounds 4-H Building (Daily Show and Seminars)
Days Hotel Timonium (Host Hotel & Professional Lecture Series Saturday Evening)
FEATORONG:
• Captive born reptiles & amphibians for display and sale • Equipment, books & supplies
• Daily book signings • Educational exhibits • Door Prizes • Raffles • Artwork for sale
• Display & photographs of Costa Rican rainforest area purchased by MARS
• Pre-show social gathering with speakers • " Reptiles & Rainforests" art collection
• Seminars throughout the show • "Critter Contact ” hands-on animal display
Professional Lecture Series Saturday evening will feature Dr. Roger Klingenberg (herp veterinarian and author),
Jack Cover (National Aquarium in Baltimore), Jack Hubley (host of syndicated t.v. show, “Wild Moments’’ and
Kathy Love (herpetologist) and another Fabulous Fotos presentation by Bill Love (herpetologist and photographer)
Show proceeds are donated to purchase and protect rainforest and critical habitat.
Now in our ninth year, the MID-ATLANTIC REPTILE SHOW has protected
2,685 acres of critical habitat to date.
ADMISSION: Weekend Pass (includes show both days & lectures) $13 Adults, One Day Pass (Sat. or Sun.) $7 Adults.
Children 6 * 12 and Seniors over 65 - $6 (for one day or weekend pass), Children 5 & under are free.
FOR AOOD1TIIONM DNFORMATQON:
Call the MARS Hotline at 410-580-0250, visit our web site at http://www.reptileinfo.com
or e-mail us at mars@reptileinfo.com
Only registered vendors may display or sell on the premises.
Please leave your pet animals at home! They will not be admitted to the show. Thank you.
page 34
Bulletin of the Maryland Herpetological Society
Volume 38 Number 1
March 2002
News and Notes
Errata
Paulino Ponce Campos apologizes due to a mistake in the name of the
third author in the paper on pages 18-21 in Volume 37, Number 1. Following
is the correct citation:
Paulino Ponce Campos, Sara M. Huerta Ortega, Carlos Nogueira Born ex and
Hobart M. Smith.
2001. National history notes on the southern plateau night liz¬
ard, Xantusia sanchezi. Bulletin of the Maryland Herpeto-
logical Society. Volume 37 Number 1. 18-21.
Bulletin of the Maryland Herpetological Society
page 35
Volume 38 Number 1
March 2002
News and Notes
page 36
Bulletin of the Maryland Herpetological Society
Society Publication
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The Society also publishes a Newsletter on a somewhat irregular
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The Maryland Herpetological Society
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Natural History Society of Maryland, Inc.
2643 North Charles Street
Baltimore, Maryland 21218
Bulletin of the Maryland Herpetological Society
page 37
) VLvL/rss? %vnr?»
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DEPARTMENT OF HERPETOLOGY
THE NATURAL HISTORY SOCIETY OF MARYLAND, INC.
MDHS
A Founder Member of the Eastern
Seaboard Herpetological League
VOLUME 38 NUMBER 2
JUNE 2002
BULLETIN OF THE MARYLAND HERPETOLOGICAL SOCIETY
Volume 38 Number 2
June 2002
CONTENTS
Parasites in Snakes of Thailand
Naiyana Chaiyabutr and Lawan Chanhome . 39
New Reptilian Records from Eastern Iowa
Harlan D. Walley :.-,C . «... . 51
Status of Pseudacris feriarum in Prince William Forest Park, Prince William
County, Virginia
Carol A. Pollio and Sandra L. Kilpatrick . 55
A New Snake Related to Sibon sanniola (Serpentes: Dipsadidae) from Los
Tuxtlas, Veracruz, Mexico
Gonzalo Perez-Higareda, Marco A. Lopez-Luna and
Hobart M. Smith . . . . . . . . . . 62
Book Review
Harlan D. Walley . . . . . . 66
BULLETIN OF THE
Volume 38 Number 2
June 2002
The Maryland Herpetological Society
Department of Herpetology, Natural History Society of Maryland, Inc.
President
Executive Editor
Tim Hoen
Herbert S. Harris, Jr.
Steering Committee
Frank B. Groves
Jerry D. Hardy, Jr.
Herbert S. Harris, Jr. Tim Hoen
Library of Congress Catalog Card Number: 76-93458
Membership Rates
Membership in the Maryland Herpetological Society is $25.00 per year
and includes the Bulletin of the Maryland Herpetological Society. For¬
eign is $35.00 per year. Make all checks payable to the Natural History
Society of Maryland, Inc.
Meetings
Meetings are held monthly and will be announced in the "Herp Talk1
newsletter and on the website, www.naturalhistory.org.
Volume 38 Number 2
June 2002
Parasites in Snakes of Thailand
Naiyana Chaiyabutr and Lawan Chanhome
Abstract
A survey of parasites was conducted in 128 snakes belonging to 17 spe¬
cies of the families Pythonidae, Colubridae, Elapidae and Viperidae. Seventy-
five percent of the snakes examined were found to be infected with endopara-
sites and ectoparasites. Endoparasites included five species of nematodes,
one specie of cestode, one specie of trematode, one specie of Acanthocephala,
one tongue worm and one specie of protozoa. The ectoparasites found were
the hard ticks of the genera Ixodes sp Haemaphysalis sp. and Aponomma sp.
The results indicate that the most abundant parasite in the sample studied is
Kalicephalus laticaudae which adversely affect the captive breeding programme
of the snake farm at Queen Saovabha Memorial Institute (QSMI) and it is the
main cause of mortality in the snakes kept in captivity.
Introduction
The snake farm of the Queen Saovabha Memorial Institute (QSMI)
maintains various species of non-venomous and venomous snakes for both
public display and venom extraction. Venom extracted is used for both im¬
munizing horses for antivenom production and venom research. Prior to 1994,
most of the snakes at the institute had been purchased from dealers. The cap¬
tive breeding program was initiated in order to supply healthy snakes for
antivenom production and decrease the number of the wild snakes captured.
Original breeding stocks due to heavy parasite infestations, including wild-
caught snakes, had a short life span. Heavy parasitic infection in snakes has
been shown to adversely effect reproductive ability (Klingenberg, 1993). Thai¬
land has a diverse snake fauna comprising of 163 species (Jintakune and
Chanhome, 1995), but little is known of their endoparasites and ectopara¬
sites. A number of studies have been reported concerning parasites in snakes
of Malaysia (Ambu et al, 1990; Tat et al, 1980), Philippines (Fishtal and Kuntz,
1964) and Japan (Kagei, 1972; Kagei and Kifune, 1977). The present study
was carried out to survey parasites in different species of Thai snakes at the
snake farm of the QSMI which were purchased from dealers.
Key words: Snake farm, Thai snakes, endoparasites, ectoparasites.
Bulletin of the Maryland Herpetological Society
page 39
Volume 38 Number 2
June 2002
Materials and Methods
The parasitic survey was carried out from March 1996 to October 1998.
The 128 snakes beionging to 17 species from the families Pythonidae,
Colubridae, Elapidae, and Viperidae were examined. The procedures are as
follow:
1) IX race fecal smear
were collected as fresh as possible and stored in the refrigerator
in clean, dry containers with air tight lids. Samples were fixed in 10% formal¬
dehyde solution. Small amount of each sample was applied to a slide, cov¬
ered with a cover glass and then examined with the light microscope.
2) Post-mortem examination
Dead snakes were kept in the refrigerator prior to post-mortem exami¬
nation. The parasites obtained from G.I. tract, muscle, soft tissue and skin
were stored in 70% alcohol for identification. For detailed studies, nematodes
were transferred to lacto-phenol medium; trematodes, cestodes and
acanthocephalas were stained with acid carmine, dehydrated in alcohol,
cleared in methyl salicylate and mounted in permount.
3) Physical examination and skin incisions were performed on living
snakes for ectoparasites and subcutaneous parasites, respectively. Ectopara¬
sites were cleared and studied in Hoyer's medium.
Results
Of the 128 snakes examined, 96 snakes (75%) were found to be infested
with different types of parasites (Table 1 & 2). Five species of nematodes, one
specie of cestode, one specie of trematode, one specie of acanthocephala, one
tongue worm and one specie of protozoan were found.
Venomous snakes
Of the 128 snakes examined, a total of 111 (11 species) were venomous
snakes. Examination of these snakes for parasites showed that 84 (75%) were
infested with parasites (Table 1).
Five species of nematodes were identified as follows: Hook worms,
Kalicephalus laticaudae (fig. 2 & 3) were found in 7 species of snakes (Table 2).
page 40
Bulletin of the Maryland Herpetological Society
Volume 38 Number 2
June 2002
Figure 1. Hookworm
(Kalicephalus laticaudae ) ova,
200x.
Table 1. Number of different species of snakes examined for parasitic infec¬
tion.
' I
Figure 2. The anterior end of
Kalicephalus laticaudae , 200x.
Snake species No. of snakes examined
Venomous
Total number of infected (%)
1. Naja kaouthia
38
23(60.5)
2. Naja siamensis
16
16(100)
3. Ophio phagus hannah
9
9(100)
4. Bungarus fasciatus
8
8(100)
5. Bungarus candidus
9
9(100)
6. Bungarus flaviceps
4
3(75)
1. Daboia russelii siamensis
7
4(57)
8. Calloselasma rhodostoma
5
5(100)
9 . Trimeresurus albolabris
9
3(33)
10. Trimeresurus popeiorum
2
0(0)
11. Trimeresurus kanburiensis
Non-Venomous
4
4(100)
1. Python curtus
1
1(100)
2. Elaphe radiata
4
1(25)
3. Elaphe taeniura ridleyi
2
0(0)
4. Lycodon laoensis
2
2(100)
5. Rhabdophis subminiatus
6
6(100)
6. Enhydris enhydris
2
0(100)
Total 17 species
128
96(75)
Bulletin of the Maryland Herpetological Society page 41
Volume 38 Number 2
June 2002
Table 2. Species of parasite infecting snakes of Thailand.
Species and number of snakes infected
Venomous
Non-Venomous
1
2
o
4
5
6
7
8
9
10 11
1
2
3
4
5
6
Endoparasites
Nematode
Kalicephalus
laticaudae
23 16 0
8
9
0
4
5
3
0
0
1
0
0
2
4
0
Capillaria sp. (ova)
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
Oxyurus sp. (ova)
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Tangua tiara
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
Strongyloides sp .
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Cestodea
Spirometra reptans
(larva)
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
Trematodes
Pamdiastomun sp ,
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
Intestinal, protozoa
Isospom sv.
(sporocyst)
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Aeanthocepnala
Acanihocephalus
ranae
2
0
0
0
0 0
0 0 0 0 0 0
0
0
0
1
0
Tongue worm
Leiperia gracilis
0
0
0
0
0 0
0 0 0 0 0 1
0
0
0
0
0
Ectoparasite
Ixodes sp.
2
0
0
0
0 0
0 0 0 0 0 0
0
0
0
0
0
Haemaphysalis sp.
0
0
0
0
0 0
1 0 0 0 0 0
0
0
0
0
0
Aponomma sp.
2
0
0
0
0 0
0 0 0 0 0 0
0
0
0
0
0
Venomous snakes
1. Naha kaouthia 2. Naja siamensis
3. Ophiophagus hannah 4. Bungarus fasciatus 5. Bungarus cadidus
6. Bungarus flaviceps 7. Daboia russelii siamensis
8. Calloselasma rhodoslotna
9. Trimeresurus albolabris
10. Trimeresurus popeiorum 11. Trimeresurus kanburiensis
Non-Venomous snakes
1 . Python curtus 2. Elaphe radiata
3. Elaphe taeniura ridleyi 4. Lycodon laoensis
5. Rhabdophis subminiatus
6. Enhydris enhydris
page 42 Bulletin of the Maryland Herpeiological Society
Volume 38 Number 2
June 2002
The character of this parasite consists of a buccal capsule which is strongly
developed., globular or subglobular or funnel-shaped, usually with dorsal
groove. The esophageal funnel may contain three small teeth. Males show a
bursa at the posterior end and a trilobed, dorsal lobe projecting beyond lat¬
eral lobes. Two spicules are equal. Females: posterior extremity is long, coni¬
cal; vulva in posterior part of body; uterine branches opposed or parallel
(Ambu et al, 1990). Thin wall oval eggs of this nematode were found in fecal
smears (fig. 1) and accounted for 30.5% of the infection rate in the snakes
examined. A total of 50 worms (20 males and 30 females) were found. Eggs of
Oxyurus sp. were examined from the feces of Ophiophagus hannah (fig 6) and
were larger than the eggs of the hook worm. Larvae and eggs were found in
Naja kaouthia feces were identified as Strongy hides sp. (fig. 7 & 8). Opercu-
lated oval-shaped eggs of Capillaria sp. were also identified from the feces of
these snakes (fig. 9).
Figure 3. The posterior end of Figure 4. The anterior end of
male Kalicephalus laticaudae ), male Tangua tiara , lOOx.
200x.
Figure 5. The posterior end of Figure 6. Finworm ova,
male Tangua tiara, lOOx. (Oxyurus sp.) lOOx.
Bulletin of the Maryland Herpetological Society
page 43
Volume 38 Number 2
June 2002
Figure 7. Strongyloides ova, 4G0x.
Figure 8. Larvae of Strongyloides
lOOx.
Figure 9. Capillaria ova with
both-ends operculum, 200x.
Figure 10. Sparganum of
Spirometra rep tans was removed
by skin incision, in Calloselasma
rhodostoma.
Figure 11. The ventral view of Paradiastomum sp., lOOx.
Cestodes were found in both venomous and non-venomous snakes. It
was juvenile stages called "sparganum” which were collected from dense nod¬
ules in muscles, subcutaneous tissues, coelomic cavities, the mesentery of GI
tract, and gall bladders. Sparganum was found mostly under the skin of the
snakes from South Thailand and were removed by skin Incision (fig. 10).
page 44
Bulletin of the Maryland Herpetoiogical Society
Volume 38 Number 2
June 2002
A trematode (adult liver fluke) found in the fecal sample of Calloselasma
rhodostoma was identified as Paradiastomun sp (fig. 11). One specie of protozoa
was recovered in the fecal smear and intestinal contents of the Monocled
Cobra (Naja kaouthia) and Russell's Viper (Daboia russelii siamensis) which were
collected in central Thailand.
Oocysts of an intestinal protozoan were identified as Isospora sp. which
had two sporocysts (fig. 14).
The ectoparasites found were the hard ticks of the genera Ixodes sp (fig.
15), Haemaphysalis sp (fig. 16) and Aponomma sp (fig 17) and were found under
the body scales, on the head, and around the eyes.
Non- venomous snakes
A total 17 non-venomous snakes (6 species) were examined and 12
snakes (70.6%) were found positive for endoparasites (Table 1 & 2),. Four
species of snakes were found to have a high rate of infection (100%) while
only one specie had 25% and Elaphe taeniara ridleyi showed no infection. All
venomous snakes were negative for Tangua tiara and Leiperia gracilis but they
are found commonly in non-venomous snakes. Only male worms of Tangua
tiara (family Gnathostomatidae) were found in the GI tract in the present
study (fig. 4 & 5). This parasite had a head bulb which was coarsely striated
transversely, unarmed, divided externally into two or four swellings con¬
taining balloonets. The cuticle behind the head bulb was forming a more or
less pronounced collar. The posterior end had caudal alae which was well
developed, eight pairs of caudal papillae. Two spicules were equal, tubular,
with smooth tips (Yamaguti, 1935). In Rhabdophis subminiatus we found the
Acanthocephalus ranae in the soft tissue of the abdominal cavity. Several mor¬
phologies serve to separate acanthocephalans from other parasitic worms.
The most distinguishable character was the presence of numerous hooks on
the protusible proboscis. The samples collected from post-mortem examina¬
tion had five rows of hooks on the proboscis (fig. 12). Leiperia gracilis (fig. 13)
was found in the vomitus of Blood Python (Python curtus). Its size was 4-5
cm. with four pairs of hooks on the head. This worm is related to the Arach¬
nid a and generally lives in bronchi, lungs, and rarely in the heart or head.
Discussion
The high mortality rate of the venomous snakes maintained at the snake
farm of Queen Saovabha Memorial Institute is a major problem. Postmortem
Bulletin of the Maryland Herpetological Society
page 45
Volume 38 Number 2
June 2002
Figurel2. The five rows of
hooks on the proboscis of
Acanthocephaia ranae, lOOx.
Figure 13. The four pairs of
hooks on proboscis (arrow) of
Leiperia gracilis, lOOx.
Figure 14. Isospora oocyst, 400x.
Figure 15. The ventral view of
male tick, Ixodes sp., lOOx.
analysis of the dead snakes in the present study showed most of high mortal¬
ity rate is due to parasitic infection in these snakes. Parasitic infection in snakes
from different countries in the Asian region (Ambu et al., 1990; Fisthal and
Kuntz, 1964; Kagei, 1972; Kagei and Kifune, 1977; Tat et al., 1980) has also
been reported. The distribution of endoparasitic fauna has been reported for
a large number in Malaysian snakes species (Ambu et al., 1990). In the present
study, Kalicephalus laticaudae was found in many more species of venomous
snakes from Thailand then compared with the rest of Southeast Asia. Only
one adult worm of the family Gnathostomatidae was found in GI tract of
Lycodon laoensis and was identified as Tancjua tiara. Ambu, et al. (1990) found
only two male Capillaria sp. but in this study, they were found only as eggs in
Trimeresurus albolabris. The eggs of Oxyurus sp. were also found in Trimeresurus
albolabris. Infection in snakes is probably due to ingestion of fecal contami¬
nated food and water. Strongyloides sp. was identified only to genus level be-
page 46
Bulletin of the Maryland Herpelological Society
Volume 38 Number 2
June 2002
Figurel6. The ventral view of
male tick, Haemaphysalis sp
lOOx.
Figure 17. The dorsal view of
female tick, Aponomma sp.,,
200x.
cause only eggs and larva were found in Naja kaouthia.
Only one specie of cestode was recovered in both venomous and non-
venomous snakes especially the larval stage called sparganum. Sparganum
was found mostly in venomous snakes from South Thailand. Adult worms
are usually found in the small intestine of dogs and cats but the larval stage
can be found in the muscle of snakes, frogs and fishes. The adult worm,
Spirometra replans , has been associated with sparganosis in humans who in¬
gest either raw fish or raw snake. The larval stage can migrate to the brain,
spinal cord and the other parts of the body (Chang et al., 1999; De Roodt et
al, 1993; Fung et al, 1989; Kim et al, 1996; Kron et ah, 1991: Kudesia et al.,
1998; Landero et al., 1991: Norman and Kreutner, 1980; Tsai et al., 1993). No
information is available concerning sparganum in Malasian and Philippines'
snakes (Ambu et al, 1990: Fisthal and Kuntz, 1964; Kagei and Kifune, 1977).
One trematode identified as Pamdiastomun sp. (Kagei et al., 1972) has been
found in snakes in Japan. It has an indirect life cycle with one or several inter¬
mediate hosts, such as snails, frogs and fishes. This parasite actually lives in
both the bile duct and gall bladder and if will pass to the GX tract in case of
heavy infestation. One species of Acanthocephala was recovered from ven¬
omous and non-venomous snakes; Acanthocephalus ranae has an indirect life
cycle. Snakes serve as temporary or secondary intermediate hosts.
Leiperia gracilis (tongue worm) was found in the vomitus of Blood Py¬
thon (Python curtus). Snakes are probably infected by feeding on primary hosts
containing juvenile stages, such as fishes, amphibians, small mammals and
rarely birds.
Bulletin of the Maryland Herpetological Society
page 47
Volume 38 Number 2
June 2002
Isospom is genus of coccidia that affects the epithelial cells of the intes¬
tine, the biliary system, and the kidney. It has a direct life cycle. Naja kaouthia
and Daboia russelii siamensis are infected by ingestion of oocyst contaminated
food and water. The clinical signs in animals are anorexia and diarrhea. In
severe cases can cause digestive disorders, vomit, dehydration, and hemor¬
rhagic enteritis. (Barnard and Upton, 1994).
In the venomous snakes, hard ticks of the genera Ixodes sp., Haema -
physalis sp., and Aponomma §p. were found. They are blood sucking parasite
and the cause of anemia and skin damage that can lead to secondary bacte¬
rial infection. They also have the potential to transmit hemoprotozoas and
viruses to snakes.
Most parasites have an indirect life cycle and snakes serve as the inter¬
mediate host. Parasitic infestation in snakes result in variety of effects from
mild disturbance to fatality The balance of parasite-host relationship is far
different between wild snakes and captive snakes. Conditions in captivity
such as poor husbandry, inadequate diet, and overcrowding can lead to stress
that weakens an immune system (Klingenberg, 1993). This preliminary study
reveals the distribution of helminthes fauna in a large number of snakes in
Thailand. Kalicephalus laticaudae infestation is the most prominent found and
hypothesized to have caused the death of many snake species examined.
Literature Cited
Ambu, S. Krishnasamy, M. and Jefery, J.
1990. A study on endoparasites of some snakes in Peninsular Ma¬
laysia. The Snake, 22: 120-125.
Barnard, S. M. and Upton, S. J.
1994. A veterinary Guide to the Parasites of Reptiles. Vol. I Proto¬
zoa. Kreiger Publishing Company: 51-52.
Chang, J. H., Lin, O. S., and Yeh, K. T.
1999. Subcutaneous sparganosis - a case report and a review of
human sparganosis in Taiwan. Kao Hsiung I Hsueh Ko
Hsueh Tsa Chih. 15(9): 567-571 .
De Roodt, A. R., Suarez, G., Ruzic, A., Bellegarde, E., Braun, Mv and Blanco,
C. M.
1993. A case of human sparganosis in Argentina. Medicina (B
Aires), 53(3): 235-238.
page 48
Bulletin of the Maryland Herpetoiogical Society
Volume 38 Number 2
June 2002
Fisthal, J. H. and Kuntz, R. E.
1964. Amonogenetic and seven digestic Irematodes of amphib¬
ians and reptiles from Phillippines. Proc. Hehm. Soc, Wash¬
ington., 31 (2): 230-240.
Fung, C. F. Ng, T. H., and Wong, W. T.
1989. Sparganosis of the spinal cord: Case report. J Neurosurg,
71(2): 290-292.
Jintakurte, P. and Chanhome, L.
1995. The Venomous Snakes of Thailand (in Thai). Thai Red Cross
Society, Bangkok. 175 pp.
Kagei, N.
1972. Helminthes fauna of Reptiles in Japan I. The Snake, 4: 114-
117.
Kagei, N. and Kifune, T.
1977. Helminthes fauna of Reptile in Japan III. the Snake, 9: 108-
114.
Kim, D. G., Pack, S. H., Chang, K. H., Wang, K. C, Jung, H. W, Kim, H. J.,
Chi, J. G., Choi, K. S., and Han, D. H.
1996. Cerebral sparganosis: clinical manifestations, treatment, and
outcome. J Neurosurg, 85(6): 1066-1071.
Klingenberg, R. J.
1993. Understanding Reptile Parasites, A basic manual for
herpetoculturists & veterinarians. The Herpetocultural Li¬
brary Special Edition. 81 pp.
Kron, M. A., Guderian, R., Guevara, A., and Hidalgo, A.
1991. Abdominal sparganosis in Ecuad or: Case report. Am J
Trop Med Hyg, 44(2): 146-150.
Kudesia, S., Indira, D. B.„ Sarala, D., Vani, S., Yasha, T. C, Jayakumar, P. N.,
and Shankar, S. K.
1998. Sparganosis of brain and spinal cord: unusual tapeworm
infestation (report of two cases). Clin Neurol Neurosurg,
100(2): 148-152.
Bulletin of the Maryland Herpetological Society
page 49
Volume 38 Number 2
June 2002
Landero, A., Hernandez, E, Abasolo, M.A., Rechy, D. A., and Nunez, R
1991 . Cerebral sparganosis cause by Spirometm mansonoides: Case
report. J Neurosurg, 75(3): 472-474.
Norman, S. H. and Kreutner, A. Jr.
1980, Sparganosis: clinical and pathologic observations in ten
cases. South Med. J, 73(3): 297-300.
Tai, O. C, Singh, K. I. And Krishnasamy, M.
1980. A preliminary report on the fecal examination of captive
snake in Malaysia. The Snake, 12: 32-36.
Tsai, M. D., Chang, C. N., Ho, Y. S., and Wang, A. D.
1993. Cerebral sparganosis diagnosed and treatment with stereo¬
tactic techniques: Report of two cases. J Neurosurg. 78(1):
129-132.
Yamaguti, S.
1935. Studies on the Helminth Fauna of Japan. Part II. Reptiles
Nematodes. Japanese ]. Zool., 6(20): 394-403.
Yamaguti, 3
1961 . The Nematodes of Reptiles in Systema Heminthum Vol. III.
Inter Science Publishers, Inc. New Yorks., 127-128.
Naiyana Chaiyabutr: Department of Biology, Faculty of Science, Chulalongkorn
Univeristy, Bangkok, Thailand.
Lawan Chanhome: Queen Saovabha Memorial Institute, The Thai Red Cross
Society, 1871 Rama IV Road, Bangkok, 10330, Thailand, e-mail:
Lawan.C@chula .ac. th
Received 8 September 2001
Accepted 30 December 2001
page 50
Bulletin of the Maryland Herpetoiogical Society
Volume 38 Number 2
June 2002
New Reptilian Records from Eastern Iowa
The most comprehensive work on the snakes of Iowa was Guthrie
(1926), and since that time few publications relating to the herpetofauna of
Iowa have been published, except for Klims tr a (1950) on the herpetofauna
from southeastern Davis County, and Christiansen and Mabry (1985) on the
fauna of Iowa's loess hills region. Christiansen and Bailey's (1988a,b) gener¬
alized papers on the amphibians and reptiles found within the state. Since
that time Bailey and Christiansen (1993) have provided an excellent histori¬
cal review of early research on the herpetofauna, while a recent publication
by Christiansen (1998) focuses on the declining amphibians and reptiles within
the state, having been primarily caused by habitat destruction.
Recently a collection of amphibians and reptiles previously housed
in the Putnam Museum of History and Natural Sciences was donated to the
author by the curator. Dr. Christine S. Chandler, and presently is housed at
the Northern Illinois University, Dekalb, Illinois.
In an effort to update our knowledge of the distribution of certain
amphibians and reptiles of Iowa, I feel it would be noteworthy to verify the
current knowledge of geographical distributions of species found within East¬
ern Iowa. The majority of these records are at least 50 years old, but still of
major importance, as they constitute new records for the state, and should be
of historical importance for future studies
Sauria
Cnemidophorus s. sexlineatus (Eastern Six-lined Racerunner).
MUSCATINE CO: Muscatine, Bloomington twp. August 13, 1961. Bill Webb
and Peter Peterson. (RE 064-66) HDW-NIU 1723-1725.
Eumeces fasciatus (Common Five-lined Skink). JONES CO: 4 mi. West of
Canton, Clay twp. August 6, 1961. Bill Webb. (RE 048) HDW-NIU 1716; Bill
Webb and Robert Schroeder. August 6, 1961. (RE 059-62). HDW-NIU 1713-
1720. August 8, 1961 (RE 63) HDW-NIU 1721.
Serpentes
Coluber constrictor foxii (Blue Racer). JACKSON CO: Maquoketa,
Maquoketa twp.August 9, 1964. Bill Webb. (RE 223) HDW-NIU 1800.
Bulletin of the Maryland Herpetological Society
page 51
Volume 38 Number 2
June 2002
Diadcphis punctatus arnyi (Prairie Ring-necked Snake). JACKSON CO:
Canton, Clay twp. August 6, 1961. Robert Schroeder. (RE 040-41, 058). HDW-
NIU 1738-1740; Near Maquoketa, Maquoketa twp. August 1961. Bill Webb.
(RE 200-203) HDW-NIU 1741-1744.
Elaphe vulpina (Western Fox Snake). SCOTT CO: Princeton, Princeton
twp. 1961. Bill Webb. (RE 005) HDW-NIU 1876.
Heterodon nasicus nasicus (Plains Hog-nosed Snake). MUSCATINE CO:
Muscatine, Bloomington twp. Date unknown. Jim Peck. (RE 004) HDW-NIU
1776. This species is listed as Endangered in Iowa.
Lampropeltis triangulum triangulum (Eastern Milksnake). SCOTT CO:
Davenport, Davenport twp. 1966. Bill Webb. (RE 046) HDW-NIU 1748; Sep¬
tember 6, 1960. Harold Heeschen. (RE 108) HDW-NIU 1750; July 1962. C.B.
Rupp. (RE 219) HDW-NIU 1749.
Nerodia rhombifera rhombifera (Northern Diamond-backed Watersnake).
LOUISE CO: 5 mi. S. Muscatine. September 5, 1961, 20 newborn. D.G. Herold.
(RE 203) HDW-NIU 1891-1910; MUSCATINE CO: Muscatine, Bloomington
twp. September 5, 1961. Bill Webb. HDW-NIU; July 25, 1960, D.G. Herold.
(RE 103) HDW-NIU 1889.
Nerodia sipedon (Northern Watersnake): SCOTT CO: Duck Creek Park,
Davenport, Davenport twp. September 1960. Bill Webb. HDW-NIU 1836;
Princeton slough, Princeton, Princeton twp. September 8, 1961. D.G. Herold.
13 newborn. (RE 198) HDW-NIU; MUSCATINE CO: Muscatine, Bloomington
twp. July 25, 1960. D.G. Herold. (RE 089, 092) HDW-NIU 1831, 1837.
Opheodrys vernalis =Liochlorphis vernalis. (Smooth Greensnake). SCOTT
CO: Davenport, Davenport twp. 1961. Bill Webb. (RE 008-9) HDW-NIU 1711-
1713; Jim Peck (RE131) HDW-NIU 1713. Christiansen (1998) notes that popu¬
lations have suffered declines, and presently has been found on "several state
and country preserves," and is "worthy of concern." LeClere (1998) states,
"uncommon, and having suffered from habitat destruction, pesticides, and
progression." This species is presently listed as Threatened in Iowa.
Pituophis catenifer (Pacific Gophersnake). JACKSON CO: Maquoketa,
Maquoketa twp.. Date unknown. Robert Schroder. (RE-055-57). HDW-NIU
1735-1737; August 27, 1959. Robert Schroder (RE 071) HDW-NIU 1911.
page 52
Bulletin of the Maryland Herpetoiogical Society
Volume 38 Number 2
June 2002
Regina grahamii (Graham's Crayfish Snake). LEE CO: Ft. Madison, Wash¬
ington twp. Date unknown. David Loren. {RE 087) HDW-NIU 1777.
Storeria dekayi wrightorum (Midland Brown snake). SCOTT CO: Daven¬
port, Davenport twp. 1961. Bill Webb. (RE 073) HDW-NIU 1809; May 12, 1961.
Pat Reilly. (RE 129) HDW-NIU 1805; October 5, 1961. Lloyd Abbot (RE 208)
HDW-NIU 1804; October 10, 1965. George R. Cross. (RE 246) HDW-NIU 1821.
Princeton, Princeton twp. October 27, 1960. D.G. Herold. (RE 127, 130) HDW-
NIU 1806-7; September 27, 1964. D.G. Herold. (RE 225) HDW-NIU 1823
Thamnophis radix (Plains Gartersnake). SCOTT CO: Princeton, Princeton
twp. October 1960. D.G. Herold. (RE 124) HDW-NIU 1766; Davenport, Dav¬
enport twp. June 1960 Keith Oden. (RE 122-23, 125) HDW-NIU 1768,1765 and
1764; Davenport, Davenport twp. October 10, 1965. George R. Cross. (RE 246A)
HDW-NIU 1912-1918.
Thamnophis sirtalis parietalis (Red-sided Gartersnake). SCOTT CO:
Princeton, Princeton marsh, Princeton twp.. August 2, 1964. Bill Webb. HDW-
NIU; November 23, 1960, D.G. Herold (RE 117, 120) HDW-NIU 1757, 1760;
Crotalus horridus (Timber Rattlesnake) JACKSON CO: Bellevue, Bellevue
twp. September 7, 1967. Art Stuart. These are listed as one week old neonates.
(RE 260) HDW-NIU 1789-1797; JONES CO: 8 mi. S.E. of Monticello. June 25,
1961. Galen Heim. (RE 215) HDW-NIU 1705.
Literature Cited
Bailey, R.M. and J.L. Christiansen.
1993. The early history and recent trends in Iowa herpetology. J.
Iowa Acad. Sci. 100:83-86.
Christiansen, J.L.
1998. Perspectives on Iowa's declining amphibians and reptiles.
J. Iowa Acad. Sci. 105:109-114.
Christiansen, J.L. and R.M. Bailey.
1988a. The snakes of Iowa. Iowa Conserv. Comm., Des Moines,
(revised 1990) Nongame Technical Ser. (1): 15 pp.
— — and — — .
1998b. The lizards and turtles of Iowa. Iowa Dept. Nat. Resources,
Des Moines, (revised 1997), Nongame Tech. Ser. (3):19 pp.
Bulletin of the Maryland Herpetological Society
page 53
Volume 38 Number 2
June 2002
Christiansen, J.L. and C.M. Mabry.
1985. The amphibians and reptiles of Iowa's loess hills. Proe. Iowa
Acad. Sci. 92:159-163.
Guthrie, J.E.
1926. The snakes of Iowa. Iowa State Agricultural Experiment Sta¬
tion Bulletin (239):147-192.
Klimstra, W.D.
1950. Notes on some amphibians and reptiles from Davis County,
Iowa. J. Iowa Acad. Sci. 24:429-431.
LeClere, J.
1 998, Checklist of the herpetofauna of Iowa. Minnesota Herpetol.
Soc. Occas. Pap. (5):l-23.
Harlan D. Walley , Department of Biology, Northern Illinois University , DeKalb,
Illinois 60115
Received 14 December 2001
Accepted 30 December 2001
page 54
Bulletin of the Maryland Herpetoiogical Society
Volume 38 Number 2
June 2002
Status of Pseudacris feriarum in Prince William
Forest Park, Prince William County, Virginia
An amphibian monitoring program was initiated in Prince William
Forest Park (PRWI), a unit of the National Park System, in 1998 (Pollio 2000).
Habitat location and characterization, anuran call surveys, breeding surveys,
and larval sampling led Resource Management staff to identify native spe¬
cies that were uncommon or not found within the park. The following year a
study was initiated to focus on one of these species, Pseudacris feriarum, the
upland chorus frog. As a result, intensive nocturnal and diurnal field surveys
were conducted during the peak breeding period at prime habitat locations
to determine the population size and reproductive status of P. feriarum.
P. feriarum is a small anuran (19B37 mm SVL) that ranges from northern
New Jersey to the Florida panhandle in the east and west to East Texas and
southeast Oklahoma (Conant and Collins 1991). It is known to occur through¬
out Virginia with greater density in the Piedmont and upper Coastal Plain
(Martoff, et al 1980). Ernst, et al (1997) states that this once common species is
disappearing from Northern Virginia most likely due to increased urban de¬
velopment, habitat loss, or erratic weather patterns. This species can be found
near moist woodland habitats and breeds annually in small ephemeral ponds,
roadside ditches, and other suitable wet habitats. Suitable breeding habitat
for P. feriarum is defined by Mitchell (1998) as grassy areas associated with
shallow water in wetlands without fish, including roadside ditches and ver¬
nal pools in open fields. Breeding can occur from February to May in north¬
ern Virginia. Males typically begin chorusing in late February and females
arrive at ponds within two weeks under normal environmental conditions
(Briggs 1994). Most females arrive at sites over a 2-3 day period for commu¬
nal breeding and deposit clusters of 40B60 eggs attached to vegetation. Eggs
hatch within 3-4 days and transformation occurs within two months. Several
studies on a closely related species (Pseudacris triseriata) indicate that water
depth at time of oviposition plays a significant role in determining survival
rate (Webb 1994, Hecnar and Hecnar 1999). Metamorphosis can be impacted
by availability of food and predator density (Travis 1981, Pearman 1995,
Briston and Kissell, 1996).
Resource Management staff began this project by identifying P feriarum
breeding sites within park boundaries based on historic records and suitable
habitat. Of the 47 amphibian breeding sites identified during surveys con¬
ducted in 1999, only four were selected as having suitable habitat; Rt. 619 # 1,
Bulletin of the Maryland Herpetological Society
page 55
Volume 38 Number 2
June 2002
Rt. 619 # 2, Rt. 619 # 3, and Oak Ridge T ributary (Fig. 1). Two additional sites.
Lake 1 and Lake 4, were selected as potential sites based on habitat, and the
Chopawamsic Backcountry area was selected based on an historic record of
occurrence. All other historic records within the park (Table 1) could not be
used due to the lack of specific locality information.
Visual and auditory surveys were performed weekly beginning in mid-
February 2001 and were increased to two times per week in mid-March. Calls
were classified into four groups: no calls (0); individuals (1); individuals with
Figure 1. Map of Prince William Forest Park, Triangle, Virginia, showing
Pseudacris feriarum sample sites.
page 56
Bulletin of the Maryland Herpetological Society
Volume 38 Number 2
June 2002
limited overlap (2); and chorus, defined as numerous individuals calling with
complete overlap (3). After the first call was heard at a given site, daily sur¬
veys were performed to identify amplexing couples, egg masses, or any sign
of reproduction. Table 2 describes the sites and dates where P. feriarum was
observed, as well as the calling frequency. This species was not observed at
Lake 1, Rt. 619 #3 and the Chopawamsic Backcountry area. Only one male
was heard calling at Lake 4 on 11 April 2001. Oak Ridge Tributary had a maxi¬
mum of three individuals calling at any one time and Rt. 619 #2 had limited
overlap males calling for approximately three weeks. Rt. 619 #1 was the only
site where a full chorus was observed, yet it was completely dry by 11 April
2001, less than one week after the last chorus was heard. No females were
observed at any site and no site had evidence of reproduction. Larval
dipnetting was performed biweekly at the three sites that maintained water,
but none were found.
Resource Management staff had been concerned about the lack of pres¬
ence of P. feriarum in PRWI during the anuran call surveys and this concern
has increased as a result of this study. Historic records from PRWI are incom¬
plete and do not provide adequate information to determine historic popula¬
tion density, however the li terature suggests that this species was once com¬
mon in this area (Ernst, et al 1997, Martoff, et al 1980). It is reasonable to
assume that the main reason for PRWI having few populations is lack of suit¬
able habitat, but whether this lack of habitat is natural or due to human en¬
croachment has yet to be determined. Only six sites were chosen for this study
based on the habitat requirements of the species, and the only site chosen for
historic reasons lacked suitable habitat. The prime site for P. feriarum is Rt.
619 #1, which is a heavily traveled road, and this impact is undoubtedly af-
Table 1. Historic Records of Pseudacr is feriarum in PRWI
Date
Site
Description
Reference
3/1961
PRWI
None
FNH (observers initials)
5/1961
PRWI
None
FNH
6/1961
PRWI
None
FNH
7/1961
PRWI
None
FNH
3/1963
PRWI
None
HEM
4/27/1997
PRWI
Vernal pool
Mitchell 1998
Bulletin of the Maryland Herpetological Society
page 57
Table 2. Pseudacris feriarum Breeding Sites Surveyed in 2001
Volume 38 Number 2
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page 58
Bulletin of the Maryland Herpetoiogical Society
Volume 38 Number 2
June 2002
fecting the population size and breeding success. Males calling there are forced
to compete with the sound of traffic and adults migrating to the site may
have to cross the road. There is high interspecies competition for breeding
sites and the calls of other species have been said to drown out the calls of P.
feriarum during years with erratic weather patterns where breeding seasons
overlap (Briggs 1994, Webb 1994). Duellman and Trueb (1986) stated that
mating success in males is low, around 17.2%, and that coupled with envi¬
ronmental and human pressures has impacted P. feriarum populations
throughout their range.
This study indicates that the population of P. feriarum in PRWI is in
serious decline. Further investigation is needed to determine the factors af¬
fecting this population and to identify management strategies to maintain
and improve its numbers. Continued monitoring will document population
trends; but more intensive habitat surveys, coupled with calling surveys con¬
ducted in early February may assist in identifying additional populations
within the park. Drift fences with pitfall traps should be constructed around
known sites and adults subjected to mark-recapture analysis to determine
population size. Without intervention, P. feriarum will soon disappear from
this part of its historic range.
Ackno wled gment
We would like to thank Jennifer A. Lee, National Park Service, for ac¬
quiring and rectifying geographic data related to this survey, and producing
the map depicted in Figure 1. We would also like to acknowledge the Na¬
tional Park Service, National Capital Region, Washington, D.C., for provid¬
ing funding in support of this investigation.
Literature Cited
Briggs, Kay Marano.
1994. The effects of the ABlizzard of 1993@ on the breeding cycle
of amphibians in northern Virginia. Catesbeiana 14(2): 30-
34.
Britson, Carol A. and Robert E. Kissell, Jr.
1996. Effects of food type on development characteristics of an
ephemeral pond-breeding anuran, Pseudacris triseriaia
feriarum. Herpetologica 52(3): 374-382.
Bulletin of the Maryland Herpetological Society
page 59
Volume 38 Number 2
June 2002
Conant, Roger and John T. Collins.
1991. Peterson Field Guide: Reptiles and Amphibians, East¬
ern/Central North America. Houghton Mifflin Company,
Boston: 327-328.
Duellman, William E. and Linda Trueb.
1986. Biology of Amphibians. The Johns Hopkins University
Press, Baltimore: 671.
Ernst, Carl H., Scott C. Belfit, Steven W. Sekscienski and Arndt F. Laemmerzahl.
1997. The amphibians and reptiles of Ft. Belvoir and northern
Virginia. Bull. MD Kerp. Soc. 33(1): 8-9.
Hecnar, Stephen J. and Darlene R. Hecnar.
1999. Pseudacris triseriata. Herpetological Review 30(1): 38.
Mitchell, Joseph C.
1998. Amphibian decline in the mid-Atlantic region: monitoring
and management of a sensitive resource. Unpublished fi¬
nal report of the Department of Defense Legacy 2000 Pro¬
gram.
Martoff, Bernard S., William M. Palmer, Joseph R. Bailey, Julian R. Harrison
III.
1980. Amphibians and Reptiles of the Carolinas and Virginia. The
University of North Carolina Press, Chapel Hill: 127.
Pearman, Peter B.
1995. Effects of pond size and consequent predator density on
two species of tadpoles. Oecologia 102: 1-8.
Pollio, Carol A.
2000. Amphibians and abandoned mines spawn collaboration of
scientific disciplines. National Park Service Natural Re¬
source Year in Review 2000: 11.
Travis, Joseph.
1981. Control of larval growth variation in a population of
Pseudacris triseriata (Anura: Hylidae). Evolution 35(3): 423-
432.
page 60
Bulletin of the Maryland Herpetological Society
Volume 38 Number 2
June 2002
Webb, Dr. Glenn R.
1994. The colonization of chorus frogs and other amphibians in
artificial ponds. Reptile and Amphibian Magazine: July/
August: 57-63.
Carol A. Pollio , US Fish and Wildlife Service, 4401 N. Fairfax Drive, Suite 840,
Arlington, VA 22203. e-mail: caroljpollio@fws.gov
Sandra L. Kilpatrick, Research Contractor (formerly National Fail c Service,
Triangle, VA)
Received 22 March 2002
Accepted 9 May 2002
Bulletin of the Maryland Herpetoiogical Society
page 61
Volume 38 Number 2
June 2002
A New Snake Related to Sibon sanniola (Serpentes:
Dipsadidae) from Los Tuxtlas, Veracruz, Mexico
Gonzalo Perez-Higareda, Marco A. Lopez-Luna and Hobart M. Smith
Abstract.
Sibon linearis is described from Balzapote, Veracruz, Mexico. It most
closely resemlbes S. sanniola , but is -400 km from the nearest known locality
for that species, has a linear instead of blotched pattern, and 20 vs 23-26 dentary
teeth.
A snake found dead on a trail a few years ago near Balzapote, Veracruz,
Mexico, appears to represent a species similar to S. sanniola , endemic to the
Yucatan peninsula, including Belize (Kofron, 1990; Lee, 1996; Campbell, 1998).
The specimen from Los Tuxtlas has a strikingly distinctive dorsal pattern and
a lower number of dentary teeth (20 vs 23-26) than S. sanniola (Kofron, 1990).
Los Tuxtlas is an area of strong endemicity, and is some 400 km from
the nearest known locality, in Campeche, for S. sanniola. Given also the cited
differences from that species, we regard the specimen from Los Tuxtlas as
representative of a hitherto unknown species that we here name
Sibon linearis sp. nov.
Holotype. Universided Nacional Autonoma de Mexico - Los Tuxtlas
(UNAM-LT) 1796, an adult female from Balzapote, municipality of San Andres
Tuxtla, Veracruz, Mexico, 100 m. August 1983, GPH collector. The left side of
the head is crushed, and the posterior infralabials were eaten by ants. Other¬
wise the specimen is in good condition.
Diagnosis. Similar to S. sanniola but with a lineate instead of blotched
pattern, and 20 uv. 23-26 dentary teeth.
Descrption of holotype. Dorsal scale rows 15-15-15, smooth, without
apical pits; ventrals 155; subcaudals 70; anal single; preoculars 3-3; postoculars
2-2; loreals 1-1; temporals 2-2; supralabials 9-9; a postmental separating the
first pair of infralabials; eyes very large, diameter over 3 times distance from
lip. Maxillary teeth 11, dentary 20. Total length 311 mm, SVL 225 mm, tail 86
mm. Body not laterally compressed.
page 62
Bulletin of the Maryland Herpetological Society
Volume 38 Number 2
June 2002
Fig. 1. Dorsal Fig. 2. Midbody pattern of UNAM-LT 1796.
pattern of head
and anterior part
of the body of
UNAM-LT 1796.
Fig. 3. Lateral head scales of
UNAM-LT 1796.
UNAM-LT 1796. The shaded areas
indicate ant-damage.
Bulletin of the Maryland Herpetological Society
page 63
Volume 38 Number 2
June 2002
Body light brown, slightly darker on flanks; a narrow, irregular, con¬
tinuous dark stripe along the vertebral scale row from the rear nape to about
a sixth of the body length, extending here and there onto the adjacent edges
of the paravertebral scale rows. Posteriorly, the stripe continues interrupted
on the vertebral scale row, in a series of short streaks three scales long, sepa¬
rated from each other by an interspace of one scale length; scales of each
streak dark-edged, light-centered. On the fourth scale row is a series of darker
brown streaks 2-3 scales in length, separated from each other by a space 2-3
scales in length; they are diffuse on the tail.
Superior part of head brown, with a median reddish-brown stripe from
rostral to parietals, expanding on the posterior part of the head and becom¬
ing six scales wide on mid-nape; its width diminishes posteriorly to fuse with
the vertebral stripe on the rear part of nape. Dorsal and lateral head scales,
except for the supralabials, heavily pigmented; supralabials yellowish, their
ventral borders dark brown; a small dark spot on side of neck, covering 2-3
scales. The head is pale yellow ventrally, with scattered, dark pigmentation.
Venter light gray in preservative, weakly pigmented, more so on ends of
ventrals and in some areas onto the first dorsal scale row.
Comments. This appears to be a terrestrial species, as are other species
of Sibon. The type was found in a tropical rain forest about two km from the
coast.
The name linearis is a Latin word meaning "of a line."
The accompanying drawings were prepared by MALL.
Literature Cited
Campbell, J. A.
1998. Amphibians and reptiles of northern Guatemala, the
Yucatan and Belize. Norman, Oklahoma, Univ. Oklahoma,
xix, 380 pp.
Kofron, C. P.
1990. Systematics of neotropical gastropod-eating snakes: the
dimidiata group of the genus Sibon, with comments on the
nebulata group. Amphibia-Reptilia 11: 207-223.
page 64
Bulletin of the Maryland Heroetological Society
Volume 38 Number 2
June 2002
Lee, J. C.
1996. The amphibians arid reptiles of the Yucatan peninsula.
Ithaca, New York, Comstock, xii, 500 pp.
GPH: Estacion de Biologia Tropical , Institute de Biologia, UNAM, Apartado
Postal 51, Catemaco, Veracruz, Mexico.
MALL: Centro de Investigaciones herpetoldgicas de Veracruz A. C., Apartado
Postal 473, Cordoba, Veracruz, Mexico.
HMS: Department ofEPO Biology, University of Colorado, Boulder : Colorado
80309-0334 U.S.A.
Received 27 March 2002
Accepted 10 May 2002
Bulletin of the Maryland Herpetoiogical Society
page 65
Volume 38 Number 2
June 2002
News and Notes
Book Review
Amphibians and Reptiles of Pennsylvania and the Northeast, by Arthur
C. Hulse, C.J. McCoy and Ellen J. Censky. 2001. Cornell University Press,
Ithaca, NY 14850. x + 419 pp., 133 color plates, 83 maps. Hardcover. US$39.95.
ISBN 0-80143768-7.
While most field guides are faiiiy standard in subject and content mat¬
ter, giving only brief descriptions, and natural history notes, this book goes
beyond that stage, and provides detailed information on natural history and
ecology for each of the 83 species found within the northeastern states. Where
information is lacking on reproduction and general natural history for a spe¬
cific species in Pennsylvania, information has been provided from other
sources within the species range, giving both professional or amateur herpe¬
tologists a clearer picture of the life history of each species.
It is with regret that C. J. McCoy "Jack," co-author of this book died on
July 7, 1993, and was instrumental in the production of this book, having
previous published the "Amphibians and Reptiles in Pennsylvania: Check¬
list, Bibliography, and Atlas of Distribution" (1982).
The book , as stated in the Preface, was written for a broad audience,
and directed both to professional and amateur herpetologists, students, and
anyone having a love for nature, and passion to learn more about the organ¬
isms they might encounter in the field. T he book opens with a table of
contents, followed by a preface and acknowledgments. The introduction pro¬
vides information on landform patterns, climate, vegetation, along with ex¬
planations regarding family, genus, species, standard and common names,
descriptions, comments on confusing species, habitat and habits, reproduc¬
tion, distribution, discussed under each species account. This is followed by
a brief note on observing and collecting amphibians and reptiles, and a highly
illustrate 25 page key to the amphibians and reptiles of Pennsylvania and the
Northeastern United States. The individual species accounts form the bulk of
the book, with 322 pages devoted to 83 species, and preceded by an appendix
of Pennsylvania species mensural and reproductive date, which includes sta¬
tistical data that provides mean length for sexually mature individuals and
some neonates, range, sample size, statistical comparison between male and
female, and size at maturity, along with reproductive data on clutch size.
Individual species accounts are subdivided into sections labeled description,
page 66
Bulletin of the Maryland Herpetological Society
Volume 38 Number 2
June 2002
News and Notes
which provides information on general morphology, sexual size dimorphism,
tadpoles, confusing species, following by habitat and habits, and reproduc¬
tion.
Following the key are a set of 133 color photographs arranged four per
page. Overall the plates are excellent, but plate 132 of the Timber Rattlesnake
is extremely dark, while plate 100 of the eastern worm snake, and plate 104 of
the northern black racer are of rather poor quality. Plate 94 showing a mela-
nistic northern coal skink is of exceptional quality.
Topographical errors are few but on page 133, New Jersey is spelled in
lower case letters and with is spelled ith (=with) in the tadpole Rana virgatipes
description, page 210. Earnst 1970b (-Ernst 1970b) on page 210, and on p. 414
Yearicks et ah (1981) give a title, but lacks the journal citation. Necturus
maculosus is cited as not occurring in southwestern Wisconsin, although this
species is occasionally taken during winter and early spring months in the
Mississippi River by commercial fishermen, while Hemidactylium scutatum
has the range cited as extending through "central New England and south¬
ern Ontario to Wisconsin," which gives the impression that this species does
not occur in Illinois, although relict populations occur within northern and
central portions of the state.
In spite of only minor errors, this book is exceptionally well done, and
a must for anyone interested in the herpetofauna of eastern United States,
and particularly those interested in Pennsylvania. It should remain an au¬
thoritative treastise for many decades to come.
The glossary, bibliography, and index close out this excellent volume.
The bibliography cites over 700 references, although the literature search was
discontinued in 1998, as only one reference is cited for 1999.
Harlan D. Walley , Department of Biology, Northern Illinois University
DeKalb, Illinois 60115 .
Bulletin of the Maryland HerpetoSogical Society
page 67
Volume 38 Number 2
June 2002
News and Notes
Errata:
Paulino Ponce Campos apologizes due to a mistake in the name of the
thier author in the paper on pages 18-21 in Volume 37, Numberl. Following
is the correct citation:
Paulino Ponce Campos, Sara M. Huerta Ortega, Carlos noguiera Gomez
and Hobart M. Smith. 2001. Natural history notes on the southern plateau
night lizard, Santusia sanchezi. Bullet of the Maryland Herpetological Society.
Volume 37 Number 1. 18-21.
page 68
Bulletin of the Maryland Herpetological Society
Volume 38 Number 2
June 2002
News and Notes
Reptile and
Amphibian Rescue
410-580-0250
We will take as many unwanted pet reptiles and
amphibians as space allows.
Leave a message with your name and number to
give up an animal for adoption;
or to volunteer to help with our efforts.
OUR CURRENT NEEDS:
• Commercial or Passenger Van
• UVB Lights • Power & Hand Tools • Bleach
• Equipment & Food • Paper Towels
www.reptileinfo.com
Bulletin of the fdaryiand Herpetological Society
page 69
Volume 38 Number 2
June 2002
News and Notes
Presented by MARS Preservation Fund, Inc.
Mu
REPTILE
5HO
We believe in education,
the promotion of captive
breeding and the protection
of critical habitats.
Maryland State Fairgrounds 4-II Building (Daily Show and Seminars)
Days Hotel Timonium (Host Hotel & Professional Lecture Series Saturday Evening)
featuring
• Captive bom reptiles & amphibians for display and sale • Equipment, books & supplies
• Daily book signings • Educational exhibits • Door Prizes •Raffles • Artwork for sale
• Display & photographs of Costa Rican rainforest area purchased by MARS
• Pre-show social gathering with speakers • “Reptiles & Rainforests ” art collection
• Seminars throughout the show • " Critter Contact hands-on animal display
Professional Lecture Series Saturday evening will feature notable herpetologists to be
announced and our Fabulous Fotos presentation.
Show proceeds are donated to purchase and protect rainforest and critical habitat.
Now in our ninth year, the MID-ATLANTIC REPTILE SHOW has protected
2,685 acres of critical habitat to date.
ADMISSION: Weekend Pass (includes show both days & lectures) $13 Adults, One Day Pass (Sat. or Sun.) $7 Adults.
Children 6 - 12 and Seniors over 65 - $6 (for one day or weekend pass), Children 5 & under are free.
FOR ADDITIONAL INFORMATION:
Call the MARS Hotline at 410-580-0250, visit our web site at http://www.reptiIeinfo.com
or e-mail us at mars@reptileinfo.com
Only registered vendors may display or sell on the premises.
Please leave your pet animals at home! They will not he admitted to the show. Thank you.
page 70
Bulletin of the Maryland Herpefological Society
Volume 38 Number 2
June 2002
News and Notes
Turtle and Tortoise Publications Available from
Chelonian Research Foundation
The following turtle and tortoise publications are produced and published by Chelonian Research Foundation. The
comprehensive breadth of our publications on turtle research is unparalleled and we hope that the Quality and scope
of our materials entice you to subscribe and/or purchase. This information, including full detailed tables of contents
of all our publications, is also available on our website :i w ww.ehelonian.org — which we urge yen to visit.
6 Chelonian Conservation and Biology - International Jour, nil ojTrrtte aid tortoise
Research. CCB is the only ’n'erri&tional professional scientific peer-reviewed journal of
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Pritchard, and backed oy an ed’torial board of 30 of the world’s leading turtle authorities.
Our worldwide distribution is over 1000 to more than 60 nations. Included in the contents
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Bulletin of the Maryland Herpetoiogical Society
page 71
Volume 38 Number 2
June 2002
News and Notes
page 72
Bulletin of the Maryland Herpetological Society
Society Publication
Back issues of the Bulletin of the Maryland Herpetological Society
where available, may be obtained by writing the Executive Editor, A list
of available issues will be sent upon request. Individual numbers in stock
are $5.00 each, unless otherwise noted.
The Society also publishes a Newsletter on a somewhat irregular
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lished are Maryland Herpetofauna Leaflets and these are available at
$.25/page.
Information for Authors
All correspondence should be addressed to the Executive Editor.
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Major papers are those over five pages (double spaced, elite type)
and must Include an abstract. The authors name should be centered un¬
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pers are those papers with fewer than five pages. Author's name is to be
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rections to proof, and must return proof preferably within seven days.
The Maryland Herpetological Society
Department of Herpetology
Natural History Society of Maryland, Inc .
2643 North Charles Street
Baltimore , Maryland 21218
Bulletin of the idaryiand Herpetological Society
page 73
US ISSN:
0025-4231
fff BULLETIN or THC
TRacylanb
f)ecpetological
©oriety
DEPARTMENT OF HERPETOLOGY
THE NATURAL HISTORY SOCIETY OF MARYLAND, INC.
MDHS . A Founder Member of the Eastern
Seaboard Herpetological League
30 SEPTEMBER 2002
VOLUME 38 NUMBER 3
BULLETIN OF THE MARYLAND HERPETOLOGICAL SOCIETY
Volume 38 Numbers
September 2002
CONTENTS
Stomach Content Analysis of 13 North American Toad Species
Kiersten Cook, Sam Droege, Arthur Remngton Kellogg.. 75
Cephalic Dichotomy in the Mangrove Salt Marsh Snake, Nerodia clarkii
compressicauda (Colubridae: Natricinae)
Kenneth L. Krysko, Coleman M. Sheehy III, and
John N. Decker. . . . . . . . . 86
A Replacement Name for Bell's Spiny Lizard, Sceloporus belli
Hobert M. Smith, David Chiszar and
Julio A. Lemos-Espinal . . . . 88
Book Review
Harlan D. Walley
91
BULLETIN OF THE
mbt)8
Volume 38 Numbers September 2002
The Maryland Herpetological Society
Department of Herpetology, Natural History Society of Maryland, Inc.
President Tim Hoen
Executive Editor Herbert S. Harris, Jr.
Steering Committee
Frank B. Groves Jerry D. Hardy, Jr.
Herbert S. Harris, Jr. Tim Hoen
Library of Congress Catalog Card Number: 76-93458
Membership Rates
Membership in the Maryland Herpetological Society is $25.00 per year
and includes the Bulletin of the Maryland Herpetological Society. For¬
eign is $35.00 per year. Make all checks payable to the Natural History
Society of Maryland, Inc.
Meetings
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newsletter and on the website, www.naturalhistory.org.
Volume 38 Number 3
September 2002
Stomach Content Analysis of 13
North American Toad Species
Kiersten Cook
Sam Droege
Arthur Remngton Kellogg
Abstract
New summaries of the diets of 13 species of North American toads are
presented from stomach contents analyses.
Diet is a fundamental facet of a species' life history; basic information,
essential to all ecological studies and conservation plans. From a knowledge
of diet comes insight into a species' evolution, morphology, physiology, di¬
etary specialization, food niche, habitat requirements, activity periods, vul¬
nerability to food base changes; and resource partitioning among species,
across habitats, and over time.
The literature suggests toads are primarily nocturnal, opportunistic
foragers, with diets rich in ants and beetles (Bragg 1940; Smith and Bragg
1949; Clarke 1974; Duellman and Trueb 1986). An excellent summary of what
was known of toad diet prior to 1974 may be found in Clarke (1974). Subse¬
quent major works on toad diet include studies by Toft (1980 a; b; 1985),
Dimmit and Ruibal (1980), Flowers and Graves (1985), and Barrentine (1991).
In contrast to other North American anurans, toads have the ability to
eat arthropods normally found unpalatable to most species (eg. beetles, ants,
wasps, and sowbugs, Clarke (1974)). Toads' catholic tastes, ubiquity, and
local abundance has caused biologists in the past to view toads as potential
pest control agents, with the multiple introduction of the cane toad ( Bufo
marinus) as prominent case-in-point. In this note, we capitalize on some of
the early economic works of these biologists and present previously unpub¬
lished stomach content data for 13 species of toads (Bufo sp.). The data pre¬
sented in this paper stand apart from most previous studies, in that collec¬
tions were made across much of the species' ranges and throughout most of
their activity periods. These data offer, to our knowledge, the largest pub¬
lished sample sizes for the diets of B. alvarius, B. boreas, B. americanus, B. canorus,
B. fowleri, B, hemiophrys, B. puctatus, and B. valliceps and the only record of
diet for B. debilis.
Bulletin of the Maryland Herpetological Society
page 75
Volume 38 Numbers
September 2002
The specimens used in this paper came primarily from museum and
university collections, with collection dates ranging from approximately 1852-
1922. The US Biological Survey's technician. Remington Kellogg (later to
become the Assistant Secretary (Science) of the Smithsonian), processed most
of the toads, although, F. E. L. Beal and C. W. Leister, also at the Bureau, did
some analyses. While we found no explicit records of the methodology used
to quantify stomach contents, we do not suspect that Kellogg's methods dif¬
fered from those being used throughout that laboratory, in particular for birds
(Martin, 1949). The rationale for both bird and toad diet studies was to assess
these animal's economic significance in the control of noxious insects (Kellogg
1922; US Biological Survey 1934).
Stomach contents were sorted to phylum, subphylum, order, or family
level. Proportions were calculated using the food item's volumetric displace¬
ment of water. Gravel and vegetation were also included in the volume. For
each specimen a stomach contents card was filled out, giving the species'
name, life stage; the date, month and year of collection, collection site, alti¬
tude, and the name of the collector, (however, not all of these fields were
recorded consistently). In most cases the date of collection was also pro¬
vided, along with the name of the museum or university that supplied the
specimen. Apart from two popular government tracts on the usefulness of
toads to humanity (Kellogg 1922; 1928), in which data for B. americanus were
presented, these data have never been published. The original cards are
housed at Patuxent Wildlife Research Center and are available for use by
appointment with the second author.
Kellogg tabulated the data for 13 species in the early 1920's. Those tables
were discovered in storage at the Patuxent Wildlife Research Center, double
checked against the original cards, updated taxonomically and mathemati¬
cally, and are presented here. A future statistical analysis of the original data
cards is planned.
Tabulations of the percentage by volume of the stomach contents are
presented in Table 1. The data are divided into Spring (February - June) and
Summer (July - September) diets to accentuate major seasonal differences. In
Table 2 we summarize the distribution of the records for each species by the
States and Countries in which the collections were made and the number of
collections made per month, to document the extent of collections. Diets
were not separated by age class or gender, although a majority of specimens
were identified as adult males.
page 76
Bulletin of the Maryland Herpetological Society
Volume 38 Number 3
September 2002
Table 1. Diets of 13 North American toad species (Bufo sp.) collected
across N. America from 1852 to 1922, presented as percent of the total stom¬
ach volume for n specimens. Families are indented. See text for months that
represent Spring and Summer for each species (Spr. = Spring, Sum. = Sum¬
mer).
B. alvarius
B. americanus
B. boreas
B. cognatus
Month
Spr,
Sum.
Spr.
Sum.
Spr.
Sum.
unknown
n
15
160
245
58
178
19
27
ANNELIDA (Phylum)
0
0.96
0
0
0
0
1.30
MANDIBULATA (Subphy)
0.93
4.83
4.28
7.64
4.07
0.79
0
ISOPODA (Class)
0
1.38
0.30
4.50
0.2
4.21
0
AMPHIPODA (Class)
0
0
0
2.03
0.39
0
0
INSECTA (Class) (misc.)
0.80
0.38
1.84
2.64
2.83
0.89
0.19
THYSANURA (Order)
0
0
0
0
0
0
0
ORTHOPTERA (Order) (misc.)
1.87
0.59
2.07
0.36
0.59
2.37
0
Caelifera/Ensifera (Suborder)
2.47
0.75
2.72
0.07
5.58
1.26
4.74
Gryllidae
2.47
0.44
0.97
0.50
1.40
4.68
1.78
ISOPTERA (Order)
8.20
0
0
0
0
0
1.89
DERMAPTERA (Order)
0
0
0
0.05
0.07
0
0.07
HEMIPTERA (Order)
0.13
3.31
0.79
2.03
1.84
174
3.63
COLEOPTERA (Order) (misc.)
1.67
5.63
3.72
4.66
5.31
2.68
3.44
Carabidae
21.27
14.96
20.36
14.62
7.44
11.79 19.15
Hydrophilidae
1.33
0.61
0.14
0.33
0.60
3.63
0.04
Staphylinidae
0
0.70
1.46
1.22
2.06
0.84
0.70
Lampiridae / Cantharidae
0
0.37
0.26
1.67
0.30
0
0
Elateridae
0.07
2.86
2.03
1.12
1.67
0.68
1.22
Coccinellidae
0
0.30
0.34
1.02
1.40
0.53
0.04
Tenebrionidae
13.53
2.08
1.58
2.71
2.94
8.37
4.26
Scarabaeidae
14.80
10.79
5.22
2.55
2.25
9.63
13.19
Cerambycidae
4.73
0.16
0.49
0.78
0.78
0
0
Chrysomelidae
0
1.20
0.80
0.97
0.51
3.05
6.30
Curculionoidea (Superfamily) 0
3.84
4.24
2.22
2.24
3.84
1.26
TRICHOPTERA (Order)
0
0.16
1.75
0.34
2.61
0
0
LEPIDOPTERA (Order)
0
4.91
3.14
7.74
4.64
11.26 5.78
DIPTERA (Order)
0
2.23
1.43
4.66
4.75
1.68
5.52
HYMENOPTERA (Order) (misc.) 2.67
1.45
1.53
2.91
5.27
1.37
0.89
Formicidae
8.53
17.90
23.24
10.98
24.17
18
23.26
ARACHNIDA (Subphylum) (misc.)11.87
0
0
0.26
0.91
0
0.30
PHALANGIDA (Order)
0
0.01
1.80
1.34
0.58
1.32
0.70
ACARINA / ARANEIDA (Order) 2.67
5.50
1.54
3.36
2.02
5.37
0.37
MOLLUSCA (Phylum)
0
2.36
0.84
0.02
0.57
0
0
Lizard, Bird, Crustacea
0
0
0
0
0
0
0
Toad skins
0
0.63
0.95
0
0
0
0
Animal matter (misc.)
0
1.02
0.94
0
0
0
0
Vegetable food
0
7.41
8.73
14.69
10.19
0
0
Rocks and gravel
0
0.30
0.33
0
0
0
0
Bulletin of the Maryland Herpetological Society page 77
Volume 38 Numbers
September 2002
Table 1. Continued.
B. canorus
B. speciosus
B. debilis
B.fowleri
Spr.
Sum.
Spr.
Sum.
Spr.
Sum.
Spr.
Sum.
n
7
26
16
22
6
12
115
82
ANNELIDA (Phylum)
0
0
0
0
0
0
0
0
MANDIBULATA (Subphy)
2.57
0.71
0.44
0.91
0
0
1.92
0.99
ISOPODA (Class)
0
0
6.31
0.68
0
0
0.63
1.83
AMPHIPODA (Class)
0
0
0
0
0
0
0
0
INSECTA (Class) (misc.)
1.14
7.28
0.88
1.91
0
0
0.30
0.72
THYSANURA (Order)
0
0
0
0
0
00
0
0
ORTHOPTERA (Order) (misc.)
0
0
0
0.45
0
0
0.36
3.37
Caelifera/Ensifera (Suborder)
0
0
0
2.C0
0
0
1.17
0.52
Gryllidae
0
0
1.25
0.05
0
0
2.40
9.11
ISOPTERA (Order)
0
0
2.88
6.55
28.29
0
010
0
DERMAPTERA (Order)
0
0
0.13
1.68
0
0
0
0.12
HEMIPTERA (Order)
0.57
1.73
0.81
2.09
3.29
0.83
2.43
1.61
COLEOPTERA (Order) (misc.)
0
2.39
0.31
3.28
0
0.17
1.38
2.21
Carabidae
16.57
12.74
32.50
15.79
0
0
22.73
32.29
Hydrophilidae
1.14
1.70
0
0
0
0
0.10
0
Staphylinidae
3.71
0.81
0.06
0.27
0
0
0.42
0.16
Lampiridae / Cantharidae
0
0.20
0
0.14
0
0
0.54
0
Elateridae
3.86
2.01
0.81
2.18
1.43
0
5.21
2.96
Coccinellidae
2.86
2.42
0.06
0
0
0
1.34
1.06
Tenebrionidae
4.00
0.97
0.38
4.78
0
0.17
3.14
1.11
Scarabaeidae
0
0.05
8.88
18.29
6.43
0
16.39
9.05
Cerambycidae
0
0
0
0
0
0
1.06
0.73
Chrysomelidae
0
0.25
0.81
1.68
0
0
0.70
0.83
Curculionoidea (Superfamily) 0.57
0.64
1.38
1.14
0
0
6.46
4.62
TRICHOPTERA (Order)
0
0.15
0
0.05
0
0
0.01
0.61
LEPIDOPTERA (Order)
1.00
0.28
5.88
6.28
1.71
0.50
7.03
6.79
DIPTERA (Order)
HYMENOPTERA (Order)
1.00
13.00
0.81
0.91
0.71
0
1.34
0.74
(misc.)
0.14
1.48
0.13
1.41
0
0
3.26
0.80
Foimicidae
ARACHNIDA (Subphylum)
56.00
44.24
31.69
23.43
53.86
98.33
16.45
13.70
(misc.)
0
0
0
1.77
4.29
0
0.03
0.30
PHALANGIDA (Order)
ACARINA/ARANEIDA
2.00
1.12
1.56
0.32
0
0
0
1.04
(Order)
2.86
5.83
2.06
1.96
0
0
2.87
0.85
MOLLUSCA (Phylum)
0
0
0
0
0
0
0
0.21
Lizard, Bird, Crustacea
0
0
0
0
0
0
0
0
Toad skins
0
0
0
0
0
0
0.22
1.66
Animal matter (misc.)
0
0
0
0
0
0
0
0
Vegetable food
0
0
0
0
0
0
0
0
Rocks and gravel
0
0
0
0
0
0
0
0
page 78
Bulletin of the Maryland Herpetological Society
Volume 38 Numbers
September 2002
Table 1. Continued.
B. hemiophrvs
B. punctatus
B. quercicus
B. terrestris
Spr.
Sum.
Spr.
Sum,
Spr.
Sum.
Spr.
Sum.
n
15
23
12
17
75
58
99
8
ANNELIDA (Phylum)
2.33
0
0
0
0
0
0
0
MANDIBULATA (Subphy)
5.87
0
0
0.53
0.33
0
0.25
0
ISOPODA (Class)
0
0
0
0.59
0
0
1.38
0
AMPHIPODA (Class)
0
3.26
0
0
0
0
0
0
INSECTA (Class) (misc.)
0.07
3.52
2.10
0.47
0.20
0.14
2.48
0
THYSANURA (Order)
0
2.61
0
0
0
0
0
0
ORTHOPTERA (Order) (misc.)
0
0.13
0
0.59
0.45
0
0.93
1.99
Caelifera/Ensifera (Suborder)
0.13
3.74
0
0.30
0.05
0.28
1.26
0.66
Gryllidae
0
0
2.62
0
1.03
0.69
2.11
17.05
ISOPTERA (Order)
0
0
0
21.93
0.04
0
0.01
0
DERMAPTERA (Order)
0
0
0
0
0
0
1.23
0
HEMIPTERA (Order)
0.07
2.48
1.63
6.68
3.87
0.40
2.86
9.60
COLEOPTERA (Order) (misc.)
1.93
9.96
0.06
0.77
2.51
1.81
2.79
3.31
Carabidae
23.13
5.26
2.10
9.34
7.43
2.16
12.23
11.59
Hydrophilidae
0.73
2.17
0
0
0.13
0
4.64
6,62
Staphylinidae
5.07
2.61
30.30
1.06
1.87
0.85
1.01
0
Lampiridae / Cantharidae
1.07
0
0
0
0.17
1.69
0.13
0
Elateridae
3.20
0.74
0
0.35
1.59
0.79
0.86
7.62
Coccinellidae
0.47
0
0
0
0.01
0.09
0
0
Tenebrionidae
0
0.09
30.30
2.84
0.51
0.38
5.81
0.99
Scarabaeidae
13.93
0.35
8.68
2.13
3.35
3.36
12.95
20.36
Cerambycidae
0.27
0.13
0
0
0.08
0
1.31
8.11
Chrysomelidae
5.53
1.52
0
2.07
3.07
0.66
1.60
0.17
Curculionoidea (Superfamily) 6.07
1.26
0
0.83
2.67
5.67
6.02
3.311
TRICHOPTERA (Order)
0.33
0
0
0
0.33
0
0.95
0
LEPIDOPTERA (Order)
6.40
1.22
0.23
0.12
0.25
0.29
6.03
4.97
DIPTERA (Order)
2.73
7.48
1.11
0
3.44
0
7.03
0
HYMENOPTERA(Order) (misc.) 0.40
1.39
0.12
0.18
2.69
0.91
1.04
0.33
Formicidae
17.73
47.57
15.56
40.43
59.36
79.13
14.75
3.31
ARACHNIDA (Subphylum)
(misc.)
0.07
0.09
0
2.01
0.88
0.34
1.42
0
PHALANGIDA (Order)
0
0.96
0
0
0.27
0
0.41
0
ACARINA / ARANEIDA
(Order)
1.93
0.70
5.19
1.65
2.09
0.19
4.20
0
MOLLUSCA (Phylum)
0.33
0.78
0
0
0
0.17
2.03
0
Lizard, Bird, Crustacea
0
0
0
0
0
0
0.26
0
Toad skins
0
0
0
5.14
1.33
0
0
0
Animal matter (misc.)
0
0
0
0
0
0
0
0
Vegetable food
0.20
0
0
0
0
0
0
0
Rocks and gravel
0
0
0
0
0
0
0
0
Bulletin of the Maryland Herpetological Society
page 79
Volume 38 Number 3
September 2002
Table 1. Continued
B. valliceps
Spr.
Sum.
n
42
67
ANNELIDA (Phylum)
0
0
MANDIBULATA (Subphy)
4.80
3.70
ISOPODA (Class)
3.47
2.55
AMPEDPODA (Class)
0
0
INSECTA (Class) (misc.)
2.13
0.81
THYSANURA (Order)
0
0
ORTHOPTERA (Order) (misc.)
3.00
4.51
Caelifera/Ensifera (Suborder)
0.30
0.84
Gryllidae
5.46
14.16
ISOPTERA (Order)
1.71
1.57
DERMAPTERA (Order)
1.05
7.16
HEMIPTERA (Order)
3.56
4.09
COLEOPTERA (Order) (misc.)
2.48
2.31
Carabidae
11.09
3.63
Hydrophilidae
1.05
0.15
Staphylinidae
0.33
0.10
Lampiridae / Cantharidae
0.38
0.16
Elateridae
4.05
1.03
Coccinellidae
0.02
0
Tenebrionidae
1.55
1.55
Scarabaeidae
7.12
18
Cerambycidae
1.66
0.34
Chrysomelidae
2.16
0.72
Curculionoidea (Superfamily)
2.93
2.94
TRICHOPTERA (Order)
0
0.61
LEPIDOPTERA (Order)
6,07
3.22
DIPTERA (Order)
1.78
2.49
HYMENOPTERA (Order) (misc.)
0.96
1.51
Formicidae
21.44
15.09
ARACHNIDA (Subphylum) (misc.)
0.14
3.89
PHALANGIDA (Order)
1.59
0.73
ACARINA / ARANEIDA (Order)
3.84
5.60
MOLLUSCA (Phylum)
0.45
0.28
Lizard, Bird, Crustacea
0
0
Toad skins
3.40
0.75
Animal matter (misc.)
0
0
Vegetable food
0
0
Rocks and gravel
0
0
page 80
Bulletin of the Maryland Herpetological Society
Volume 38 Numbers
September 2002
Table 2. States and Countries, number of toads collected in each month;
and modal and median year of collection for 13 species of N. American toads
(Bufo).
B. alvarius
B. americanus B. boreas
B. canorus
B. cognatus
States and Countries
A Z
CAN, DC, IL, AK, CA,
IN, LA, LA, CAN, MT,
MA, MI, MO, ID, NV,
NC, NY, OR, WA
OH, PA, VA,
WI,WV
CA (Yosemite
Nat-1 Park)
AZ, CO,
MEX, MT,
NE, NM,
TX
Feb.
nd
0
0
1
Mar.
nd
4
0
0
5
Apr.
nd
12
10
0
0
May
nd
51
15
4
2
Jun.
nd
93
33
3
11
JuL
nd
170
115
22
15
Aug.
nd
60
45
3
7
Sept.
nd
15
18
1
5
Range of years
1892-1917
1853-1922
1855-1920
1915-1922
1852-1920
Modal year
1917
1919
1919
1915
1893&1920
Median year
1914
1907
1911
1915
1913
Bulletin of the Maryland Herpetological Society
page 81
Volume 38 Number 3
September 2002
Table 2. Continued
B. speciosus
B. debilis
B. fowleri
B. hemiophrys Kpunctatus
States
AZ,MEX,TX
TX
NJ,AL, DC,
GA
ND
AZ, CA, TX
Feb.
0
0
0
0
0
Mar.
1
0
17
0
5
Apr.
1
4
10
0
4
May
7
1
27
0
3
Jun.
7
2
61
14
0
Jul.
19
12
45
18
12
Aug.
2
0
29
5
2
Sept.
1
0
8
0
2
Range of years
1883-1919
1902-1916
1853-1921
1904-1920
1852-1919
Modal year
1916&1917
1916
1917
1915
1882
Median year
1907
1916
1917
1915
1891
page 82
Bulletin of the Maryland Herpetological Society
Volume 38 Numbers
September 2002
Table 2. Continued
States
B. quercicus
Ah, FL, GA
B. terrestris
AL, FL, GA, SC
B. valliceps
GUAT, LA, MEX, IX
Feb.
0
4
6
Mar.
5
2 (Feb.+Mar.=80)
0
Apr.
11
6
4
May
19
6
0
Jun.
40
8
23
Jul.
54
0
5
Aug.
4
0
10 (Jul.+Aug.=19)
Sept.
0
0
0
Range of years
1891-1922
1875-1922
1853-1923
Modal year
1922
1901
1923
Median year
1921
1901
1918
Bulletin of the Maryland Herpetological Society
page 83
Volume 38 Number 3
September 2002
Ackno wled gments
We would like to thank Regina Lanning for her entry of the original
diet tables in a database, Chris Binckley, for his help in literature searching
and entering collection data; and three anonymous reviewers.
Clarke, R. D.
1974.
Literature Cited
Food habits of toads, genus Bufo (Amphibia: Bufonidae).
The American Midland Naturalist. 91: 140-147.
Duellman, W. E. and Trueb, L.
1986. Biology of Amphibians. McGraw-Hill, New York. 670 pp
Kramer, W. C.
1972.
Food of the frog Rana septentrionalis in New York. Copiea.
2: 390-392.
Kellogg, A. R.
1922.
The toad. U. S. Dept, of Agriculture Bureau of Biological
Survey. Bi-664. pp. 7.
1928.
Toads destroy many harmful insects and should be pro¬
tected. Yearbook of Agriculture, pp. 620-622.
Martin, A. C.
1949. Procedures in wildlife food studies. USDI, Wildlife Leaflet
325.
Smith, C. C. and A. N. Bragg.
1949. Observations on the natural history of the anura, VII. Food
Toft, C. A.
and feeding habits of the common species of toads in Okla¬
homa. Ecology. 30: 333-349.
1980a. Feeding ecology of thirteen species of anurans in a seasonal
tropical environment. Oecologia (Berl.). 45: 131-141.
page 84
Bulletin of the Maryland Herpetological Society
Volume 38 Numbers
September 2002
1980b . Seasonal variation in populations of Panamanian litter frogs
and their prey: a comparison of wetter and drier sites.
Oecologia (BerL). 47: 34-38.
1981. Feeding ecology of Panamanian litter anurans: patterns in
diet and foraging mode. Journal of Herpetology 15: 139-144.
1985. Resource partitioning in amphibians and reptiles. Copeia.
1: 1-21
KC, SD: USGS - Patuxent Wildlife Research Center ; Laurel MD, 20708-4038
USA.
ARK: Deceased
Received: 18 August 2002
Accepted: 23 August 2002
Bulletin of the Maryland Herpetological Society
page 85
Volume 38 Number 3
September 2002
Cephalic Dichotomy in the Mangrove Salt
Marsh Snake, Nerodia clarkii compressicauda
(Colubridae: Natricinae)
There have been more than 100 snake taxa reported to have some type
of body element duplication (Cunningham, 1937; Smith and Perez-Higareda,
1988; Pefaur and Sierra, 1995). Smith and Perez-Higareda (1988) proposed
seven separate classifications regarding specific terminology for body ele¬
ment duplication including craniodichotomy, prodichotomy,
proarchodichotomy, opisthodichotomy, urodichotomy amphidichotomy, and
holodichotomy. Here we report the first occurrence of craniodichotomy in
the mangrove salt marsh snake, Nerodia clarkii compressicauda (Fig. 1).
This specimen was captive born on 10 June 2000 from wild-collected
parents originating from Big Torch Key, Monroe County, Florida, USA. After
birth, this snake moved slightly before suddenly dying, and upon further
examination it had severe spinal deformities. This specimen was deposited
in the Florida Museum of Natural History, University of Florida (UF 120890).
Fig. 1. Dicephalic mangrove salt marsh snake, Nerodia clarkii
compressicauda (UF 120890).
page 86
Bulletin of the Maryland Herpetological Society
Volume 38 Numbers
September 2002
Natridne snakes are reported to represent approximately 18% of axial
bifurcation cases in snakes, yet this frequency of observation appears to be
correlated with ease of observation rather than fecundity (Smith and Perez-
Higareda, 1988). Since most duplex individuals die as neonates, species that
are more common in the wild or popular in captivity are more likely to be
observed after birth or hatching than less common species (Smith and Perez-
Higareda, 1988). Nonetheless, duplicity in snakes is a rare phenomenon oc¬
curring in about one out of every 100,000 neonates (Belluomini et a!.. 1978).
Literature Cited
Belluomini, H. E., P. de Biasi, G. Puerto, and V. Borelli.
1978. Bicephalia em Crotalus durissus terrificus (Laurenti) (Serpen tes,
Viperidae, Crotalinae). Mem. hist. Bui an tan, 40/41:117-
121 (for 1976-1977).
Cunningham, B.
1937. Axial bifurcation in serpents. Durham, North Carolina,
Duke University Press, vii, 91 pp.
Pefaur, J. E., and N. Sierra.
1995. New records of dicephalic snakes in museum collections.
Herpetol. Rev. 26(3):127-129.
Smith, H. M., and G. Perez-Higareda.
1988. The literature on somatodichotomy in snakes. Bull. Mary¬
land Herpetol. Soc. 23:139-153.
Kenneth L. Krysko, Coleman M. Sheehy III, and John N, Decker
Florida Museum of Natural History , Division of Herpetology, University of
Florida , Gainesville , Florida 32611 , USA.
Received: 16 January 2002
Accepted: 30 January 2002
Bulletin of the Maryland Herpetological Society
page 87
Volume 38 Number 3
September 2002
A Replacement Name for Bell’s Spiny Lizard,
Sceloporus belli
Hobart M. Smith , David Chiszar and Julio A. Lemos-Espinal
In 1995 we described a new subspecies of Sceloporus undulatus, S. u.
belli , honoring our colleague and long-time specialist in the taxonomy of the
undulatus group of the genus. Dr. Edwin L. Bell. Later, Lemos-Espinal et al.
(1998) elevated the taxon to the rank of species on the basis of sympatry at
least in one locality with S. u. consobrinus. No contrary evidence is yet known.
With considerable chagrin we have learned through our honored col¬
league that his patronym is a junior secondary homonym of Tropidolepis bellii
Gray (1831:44). Tropidolepis Cuvier (1829) is a junior synonym of Sceloporus
Wiegmann (1828), hence Gray's name is now assigned to Sceloporus and is
therefore a senior secondary homonym of Sceloporus belli Smith et al
The description of Tropidolepis bellii Gray is too brief for identification
of the taxon to which the name was applied: " Bell's Tropidolepis. Trop. bellii
Gray. Metallic green scales of the back and upper part of the body, and tail
long, strongly keeled, dagger pointed, the keels forming 14-16 ridges; of the
belly broad, blunt, those of limbs and tail smaller, becoming larger near the
end; length 10 inches. Mus. Bell."
This description probably applies to some member of th eformosus group,
but any of several species could be represented. No specimen number or lo¬
cality was given. No light is shed on the matter in Gray (1845), where one
would expect the name would be entered. It is not, and none of the 10 species
of Tropidolepis dealt with there are credited to the Bell Museum. The 1845
descriptions are briefer than the 1831 description, and differently composed,
so that no similarity in that respect leads to a possible allocation of T. bellii.
The name has to remain at least for the present, and probably for all time, in
limbo, as a nomen dubium.
Regardless of identity, T. bellii Gray remains a senior secondary hom¬
onym of S. belli Smith et al. So far as we are aware, Gray’s name has never
been used since it was proposed; it is cited in none of Boulenger's catalogs of
the lizards of the British Museum (1885, 1887), or in his monograph of
Sceloporus (1897), and of course it was not known to Smith (1939). S. belli Smith
et al. cannot automatically be regarded as a nomen protectum under the 1999
Code of Zoological Nomenclature (Art. 23.9), despite the antiquity and solely
page 88
Bulletin of the Maryland Herpetological Society
Volume 38 Number 3 September 2002
original use of Gray's name, because the former has not been in use at least 1 0
years.
In the interest of nomenclatural stability, the International Commis¬
sion on Zoological Nomenclature could be petitioned for preservation of §.
belli Smith et ah, except that there is so little stability involved. The name is
known and used almost exclusively by taxonomists; there is no extensive
popular usage. Therefore we prefer to let the Principle of Priority reign, re¬
quiring the replacement of S. belli Smith et ah We here propose the substitute
name, Scelopoms edbelli nom. mm, for the unavailable name Sceloporus belli
Smith et ah
Literature Cited
Boulenger, G.A.
1885 Catalogue of the lizards in the British Museum (Natural
History). Second edition. VbL II. London, British Museum
of Natural History, xiii, 497 pp.
1887, Catalogue of the lizards in the British Museum (Natural
History). Second edition. VbL EL London, British Museum
of Natural History xii, 575 pp.
1987. A revision of the lizards of the genus Scelopoms. Proc. ZooL
Soc London 1897; 474-522.
Cuvier, G.L.C. E D.
1829. Le regne animal, distribue d'apre son organisation. NouveMe
edition, revue et augmentee par P. A. Latreille. Vol. 2.
Deterville, Paris, xv, 406 pp.
Gray, }. E.
1831. A synopsis of the species of the kass Kept ilia. Vol. 9, 1-101
in Griffith, E. and E. Pidgeon. The class Reptilia, arranged
by the Baron C. Cuvier... London, Whittaker. 16 vols.
Bulletin of the Maryland Herpetological Society
page 89
Volume 38 Numbers
September 2002
1845. Catalogue of the specimens of lizards in the collection of
the British Museum. London, British Museum of Natural
History, xxviii, 289 pp.
Lemos-Espinal, J. A., H. M. Smith, R. E. Ballinger, G. R. Smith and D. Chiszar.
A contribution to the superspecies concept of the lizard
Sceloporus undulatus: S. u. belli a species. Southwestern Natu¬
ralist 43: 20-24.
Smith, H. M.
1939. The Mexican and Central American lizards of the genus
Sceloporus. Zool Ser. Field Mug. Nat. Hist. 26: 1-397.
. D. Chiszar and J. A. Lemos-Espinal.
1995. A new subspecies of the polytypic lizard species Sceloporus
undulatus (Sauria: Iguarddae) from northern Mexico. Texas
J. Sci. 47: 117-143.
Wiegmarm, A. R A.
1828, Beytrage zur Amphibienkunde. Isis v. Oken 21: 364-383.
HMS: EPO Biology, University of Colorado, Boulder ; Colorado 80309-0334.
DC: Department of Psychology, University of Colorado, Boulder, Colorado ,
80309-0334.
JAL : Laboratorio de Ecologia , UBIPRO , Escuela Nacional de Estudios
Pmfesionales Iztacala, UNAM , Apartado Postal 314 , Avenida de hs Barrios s/n ,
Los Reyes Iztacala , Tlalnepantla, Estado de Mexico , 54090 Mexico .
page 90
Bulletin of the Maryland Herpetological Society
Volume 38 Numbers
September 2002
News and Notes
Book Review
Amphibians of Central and Southern Africa , by Alan Charming, 2001.
Cornell University Press, Sage House, 512 E. State Street, Ithaca, NY. ISBN 0-
8014-3865-9, Cloth $49.95.
The Amphibians of Central and Southern Africa provides the first com¬
prehensive guide to the frogs, toads, and caecilians of the southern third of
Africa, which includes Angola, Botswana, Lesotho, Malawi, Mozambique,
Namibia, Swaziland, South Africa, Zambia, and Zimbabwe, written by the
world's most well-known authority on the Anuran fauna of Central and South¬
ern Africa.
The Introduction provides a synthesized review of the natural history
of the amphibians in countries forming the southern third of Africa. Each
species is described with information on local and current common names,
along with the scientific name, and specific epithet for each species. A brief
description, followed by remarks on distribution and habitat, advertisement
call, breeding tadpoles, in addition to key references on the 205 species of
frogs and toads (along with 2 species of caecilians) covered in detail within
the text. Of these, 173 species are illustrated in color in 24 plates within the
text. The illustrations of the tadpoles are superb and make this book an ex¬
ceptional edition, while the distributional maps are small and of little signifi¬
cance, giving only an approximation of the range for each species covered
within the text. One species of Hemiscus and one species of Amnirana are new
to science, and are being described in details elsewhere. Other aspects cov¬
ered within the Introduction include a brief history of amphibian studies in
Central and Southern Africa stemming from Linnaeus in 1758, a listing of the
major natural history museums in Africa along with addresses which would
prove time saving for those researchers interested in contacting collection
managers, and comments on collecting.
The author stresses the awareness of declining populations of anurans
throughout the world, which has been attributed to human activities in the
majority of the cases. A list of The World Conservation Union (IUCN) Red
Date Book species from a workshop held in Cape Town in 2000, is provided
which cites some 37 species found with southern Africa is provided.
Keys are provided for the nine families of Anurans, along with keys for
species from each family within the species accounts, which cover 356 pages
Bulletin of the Maryland Herpetological Society
page 91
Volume 38 Numbers
September 2002
News and Notes
within the text. This is followed by a short section on fossil frogs, with
Thoraciliacus an illustration from a crater lake in Namaqualand being illus¬
trated, and a 38 page well illustrated section on the identification and termi¬
nology of tadpoles for those known species found in southern Africa. A 28
page bibliography, and systematic and alphabetical index, which is current
up through 1999, round out this impressive volume.
This excellent book should be on the shelf of anyone having an interest
in central and southern African anuran fauna, as it is a comprehensive treat¬
ment of extraordinary importance, and as in former monographs by the au¬
thor they are extremely accurate in details, and filled with expertise.
The moderate cost of this cloth bound volume should warrant anyone
having an interest in anuran biology, and herpetology, and especially those
interested in showing any inclination towards Southern Africa in general
should certainly purchase a copy for their library.
Harlan D. Walley, Department of Biology, Northern Illinois University,
DeKalb, Illinois 60115 , hdw@niu.edu or hdw@inwave.com.
Received: 16 July 2002
page 92
Bulletin of the Maryland Herpetological Society
Volume 38 Number 3
September 2002
News and Notes
Reptile and
Amphibian Rescue
410-580-0250
We will take as many unwanted pet reptiles and
amphibians as space allows.
Leave a message with your name and number to
give up an animal for adoption;
or to volunteer to help with our efforts.
OUR CURRENT NEEDS:
• Commercial or Passenger Van
• UVB Lights • Power & Hand Tools • Bleach
• Equipment & Food • Paper Towels
www.reptileinfo.com
Bulletin of the Maryland Herpetological Society
page 93
Volume 38 Number 3
September 2002
News and Notes
Bulletin of the Maryland Herpetological Society
page 96
Society Publication
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where available, may be obtained by writing the Executive Editor. A list
of available issues will be sent upon request. Individual numbers in stock
are $5.00 each, unless otherwise noted.
The Society also publishes a Newsletter on a somewhat irregular
basis. These are distributed to the membership free of charge. Also pub¬
lished are Maryland Herpetofauna Leaflets and these are available at
$.25/page.
Information for Authors
All correspondence should be addressed to the Executive Editor.
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are to appear in text. Cite all literature used at end in alphabetical order
by author.
Major papers are those over five pages (double spaced, elite type)
and must include an abstract. The authors name should be centered un¬
der the title, and the address is to follow the Literature Cited. Minor pa¬
pers are those papers with fewer than five pages. Author's name is to be
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Style Manual for Biological Journals (1964), American Institute of Biological
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Reprints are available at $.07 a page and should be ordered when
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rections to proof, and must return proof preferably within seven days.
The Maryland Herpetological Society
Department of Herpetology
Natural History Society of Maryland, Inc.
2643 North Charles Street
Baltimore, Maryland 21218
Bulletin of the Maryland Herpetological Society
page 97
Maiyland
Herpetological
.Society
US ISSN: 0025-4231
a^c> ,
BULLETIN Or THE
Tftarylanb
Hccpetologiral
Ooriety
DEPARTMENT OF HERPETOLOGY
MDHS . A Founder Member of the Eastern
Seaboard Herpetological League
31 DECEMBER 2002
VOLUME 38 NUMBER 4
BULLETIN OF THE MARYLAND HERPETOLOGICAL SOCIETY
Volume 38 Number 4
December 2002
CONTENTS
Colonization of Herpetofauna to a Created Wetland
T'Shaka A. Toure and George A. Middendorf . 99
Spatial Distribution in a Neotropical Lizard, Liolaemus Quilmes
(Liolaemidae): Site Fidelity and Overlapping Among Males and Females
Monique Halloy and Cecilia Robles . . . ...118
BULLETIN OF THE
mM)s
Volume 38 Number 4 December 2002
The Maryland Herpetological Society
Department of Herpetology, Natural History Society of Maryland, Inc.
President Tim Hoen
Executive Editor Herbert S. Harris, Jr.
Steering Committee
Frank B. Groves Jerry D. Hardy, Jr.
Herbert S. Harris, Jr. Tim Hoen
Library of Congress Catalog Card Number: 76-93458
Membership Rates
Membership in the Maryland Herpetological Society is $25.00 per year
and includes the Bulletin of the Maryland Herpetological Society. For¬
eign is $35.00 per year. Make all checks payable to the Natural History
Society of Maryland, Inc.
Meetings
Meetings are held monthly and will be announced in the "Herp Talk"
newsletter and on the website, www.naturalhistory.org.
Volume 38 Number 4
December 2002
COLONIZATION OF HERPETOFAUNATO A
CREATED WETLAND
TShaka A. Toure1 and George A. Middendorf
Abstract
The colonization by amphibians and reptiles of a newly created wet¬
land was investigated at a site along Sands Road in Davidsonville, Anne
Arundel County, MD. This 52-hectare artificial wetland was constructed in a
gradient design that resulted in four distinct terraced sites that temporarily
retain rainwater (Fig. 1). This palustrine wetland site, surrounded by an emer¬
gent, young, shrub-scrub, forested area, is characterized by the appearance
of shallow temporarily flooded areas over a clay substrate that remains wet
even during the driest periods of the year with a groundwater depth less
than 1.5 m. The adjacent natural forest bordering the Patuxent River served
as a natural indicator of amphibian and reptile activity and a source for site
colonization. The created wetland site was monitored over two field seasons
(March through September 1995-96) using linear transects, frog calls, drift
fence arrays, pitfall and funnel traps, and dipnets. Sampling, conducted for
54 days revealed a total of twenty-eight species (16 amphibians and 12 rep¬
tiles). The colonization of this created wetland compared favorably in diver¬
sity to adjacent, natural forest. Factors best explaining differences in
herpetofaunal activity, across the different sites within the created wetland,
were density of vegetation surrounding the waterbody and hydroperiod.
Introduction
Amphibians and reptiles remain abundant in the deciduous forests
of the eastern United States despite their worldwide decline due to habitat
destruction, introduced predators and competitors, pesticide pollution, acid
precipitation, and global climate change (Wake and Morowitz 1990; Wake
1991). Previous regional herpetofaunal surveys for Maryland and the Dis¬
trict of Columbia region provide ample historic documentation of amphib¬
ian and reptile distribution within the DC metro region (Kelly et al. 1936;
McCauley 1945, 1949; Mansueti 1949; Stine 1953a, 1953b, Cooper 1960; Harris
1966, 1969, 1975; Conant and Collins 1998). The 63 herpetofaunal species
known for this region include 29 amphibians (14 salamanders and 15 frogs;
Table 3) and 34 reptiles (6 lizards, 10 turtles, and 18 snakes; Table 4). Because
many species found regionally are restricted to very particular and often lo-
Bulletin of the Maryland Herpetological Society
page 99
Volume 38 Number 4
December 2002
calized conditions that are not evenly distributed (Conant and Collins 1998;
Harris 1975), local diversity is often lower than regional diversity. In the nearby
200 ha Jug Bay Wetland Sanctuary (Lothian, Anne Arundel County, MD),
Smithberger and Swarth (1993) documented 39 species of amphibians and
reptiles in a six-year study.
In recent years, wetland creation has been frequently employed as a
mitigation technique to offset natural wetland losses, particularly for losses
from highway construction and other commercial and private development
(Johnston 1994). A vegetative study conducted at the Sands Road site located
in Anne Arundel County, MD (Perry et al. 1997) indicated that this mitigation
effort provided many of the ecosystem functions of a natural forested wet¬
land and suggested that, like a number of other studies (Johnston 1994;
Semiitch and Brodie 1998; Semiitch 2000a, 2000b), created wetlands may be
an effective way to deal with increasing developmental pressures. Yet, few
examinations have been made to determine whether these constructed sites
actually reproduce conditions that create functional habits for wildlife popu¬
lations (Leschisin et al. 1992).
Methods
Sites - The created wetland was constructed as a series of terraces that
allowed water flow from the highest terrace (Site D) across three earthen levees
until finally discharging through the natural forest (NF) and into the Patuxent
River. Each terrace differed slightly with respect to gradient (but averaged
approximately 1.5 m drop per 100 m), in distance from the probable coloniza¬
tion source, and in emergent vegetation (Table 1; Fig. 1). To allow comparison
of species activity and colonization patterns, study sites were located in each
of the four terrace levels and in the adjacent natural forest (NF) adjacent to
the river.
Presence - Species' presence was determined following standardized
protocols (Heyer et al 1994) that involved utilization of transects, drift fences,
funnel and pitfall traps, dipnets and frog call surveys to maximize the prob¬
ability of documenting species colonization and activity in the study area.
Linear transects - Diurnal visual surveys along 100-m linear [diagonal
across site NE-SW direction] transects were conducted at a minimum of 4
times per month within each site. Vernal pools and microhabitats were
sampled along the transect path (Scott 1994).
page 100
Bulletin of the Maryland Herpetological Society
Volume 38 Number 4
December 2002
Figure 1. Sands Road Wetland Sanctuary, Anne Arundel County, Maryland
Drift fences - Drift fence arrays, located on berms bordering each ter¬
race, allowed monitoring of movement. Drift fences (30 m) were constructed
of aluminum flashing (Bury and Corn 1987; Com 1994; Dodd and Scott 1994;
Greenberg et al 1994).
Pitfall traps - Twelve pitfall traps (5-gallon plastic white bucket), six on
each side of the drift fence were set up in each site. There were 60 pitfall traps
constructed for the overall study.
Funnel traps - Two double-ended funnel traps, one on each side of the
drift fence, were established within each site in order to capture snakes (Fitch
1951; Clark 1966; Richter 1995).
Bulletin of the Maryland Herpetological Society
page 101
Volume 38 Number 4
December 2002
Frog call surveys - Call surveys were conducted by recording anuran
choruses for a minimum of two days per month. Surveys were taken prima¬
rily during nocturnal periods, although diurnal calls were recorded (Rand
and Drewry 1994).
Dipnets - The capture of aquatic tadpoles and the observation of sala¬
mander larvae during the day were enhanced by the use of this technique.
The use of dipnets allowed amphibians to be captured in waters with thick
vegetation or debris (Wright and Wright 1949; Crisafulli 1997).
Each individual captured or seen was identified, and whenever pos¬
sible, sexed and measured (snout to vent [SVL] to the nearest mm). Species
identification was confirmed by using Green and Pauley (1987), Conant and
Collins (1998) or through consultation with the curators and staff of the
Smithsonian Institution's Division of Amphibians and Reptiles. For each in¬
dividual, date, time, location within the site, vegetation in the immediate area,
presence or absence of standing water, turbidity, air and water temperatures,
other weather conditions, as well as any other unusual features were recorded
(Heyer et al. 1994; McDiarmid 1994). Voucher specimens for each species were
collected for each site. Following standard practice (Fisani 1977), most am¬
phibian voucher specimens were immersed in a solution of chlorotone, while
tadpoles were transferred directly to formalin. Other specimens were chilled,
frozen, and thawed immediately after expiration. All specimens were tagged
with USNM field tags, preserved in formalin (40% formaldehyde), and then
transferred to ethyl alcohol (70%) (Pisani 1977), and deposited in the
Smithsonian Institution, National Museum of Natural History, Division of
Amphibians and Reptiles collection (USNM).
Table 1. Distance between natural forest (NF) and created wetland (A-D) sites.
Site
Distance from colonization source
Gradient
NF
0
1.5 meters
A
228
1.5 meters
B
321
1.5 meters
C
361
1.5 meters
D
373
1.5 meters
page 102
Bulletin of the Maryland Herpetological Society
Volume 38 Number 4
December 2002
Table 2. Amphibian and reptile species richness, diversity index (Simpson’s
and shannon- Weiener), Hmax and equitability by site.
AMPHIBIANS
Species
Richness
Simpson’s
Shannon-
Weiner
Hmax
Equitability Total
Captures
Natural
Forest
16
0.83
2.95
3.91
075
443
Site A
14
071
2.25
3.7
0.61
580
Site B
13
077
2.3
3.58
0.64
287
Site C
12
0.65
1.74
3.46
0.5
357
SiteD
13
0.83
2.71
3.58
0.75
176
Average
of wetland
sites
13
0.74
2.25
3.58
0.625
350
Standard
deviation
of wetland
sites
0.816497
0.07746
0.397576
0.09798
0.102794
170.4836
Degrees of
freedom-3
Prob.>l
*0.034897 0.329316
0.176517
*0.043474 0.310932
0.623349
REPTILES
Species
Richness
Simpson's
Shannon-
Weiner
Hmax
Equitability Total
Captures
Natural
Forest
4
0.72
1.92
2
0.96
5
Site A
6
079
2.42
2.58
0.94
9
Site B
6
0.54
1.64
In
OQ
0.64
26
SiteC
2
0.22
0.54
1
0.54
8
SiteD
6
0.64
1.94
2.58
075
20
Average
of wetland
sites
5
0.5475
1.635
2.185
0.7175
15.75
Standard
deviation
of wetland
sites
2
0.07746
0.797559
0.79
0.171343
8.732125
Degrees of
freed om--3
Prob.>l
0.651448
0.526292
0.744497
0.829919
0.25193
0.306003
Bulletin of the Maryland Herpetological Society
page 103
Volume 38 Number 4
December 2002
Activity Patterns - To determine seasonal activity patterns each site
was sampled a minimum of 4 days per month between March and Septem¬
ber (Scott 1982; Mitchell et al 1993; Scott 1994; Scott and Woodward 1994) for
a total of 54 days (26 in 1995 and 28 in 1996).
Residency - Amphibian breeding site and habitats in the natural forest
and the constructed wetland sites were monitored to establish whether a given
species' presence was temporary or permanent, i.e. whether the species was
represented by colonizing individuals or by individuals representing a fully
reproductive and self-sustaining population (Campbell and Christman 1982).
Analysis - Means, standard deviations, Simpson's and Shannon-
Weiner Diversity Indices, Hmax, an Equitability Index, and basic statistics were
calculated in Microsoft Excel 97 to examine site variation, including diversity
(Krebs 1989; Stiling 1992; Hayek 1994). An apriori significance probability level
of 0.05 was used for all statistical comparisons.
Results
In total, twenty -eight species of amphibians and reptiles were docu¬
mented in the created wetland at the Sands Road Wetland Sanctuary. These
observations include almost half of the species (63) known for the Washing¬
ton, DC Metro region (Conant and Collins 1998).
While many amphibian species were found in all sites (Table 2), reptile
species were neither widespread nor abundant (Table 4). This is somewhat
expected because amphibians are more suited for wetland environments
which this was. Particularly widespread (at all sites) and abundant (10 or
more observations) amphibians included American toad ( Bufo americanus),
green frog (Rana clamitans), and southern leopard frog (R. sphenocephala utri-
cularia). Species found at all sites but with fewer than 10 observations in
some sites — included cricket frog (Acris crepitans ), Fowler's toad ( B.fowleri ),
pickerel frog (R. palustris), spadefoot toad (Scaphiopus holbrookii), marbled sala¬
mander (Ambystoma opacum), and northern slimy salamander ( Plethodon
glutinosus) (Table 3). While no single species of reptile was found in all sites,
painted turtle (Chrysemys picta) and eastern mud turtle (Kinostemon subrubrum)
were encountered in four of the five sites (Table 4).
While all amphibians observed in the created wetland were documented
in the natural forest, only 4 of the 12 expected reptiles were recorded (Tables
3 and 4). The natural forest with its vernal pools and adequate vegetative
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Bulletin of the Maryland Herpetological Society
Volume 38 Number 4
December 2002
Table 3. Number of amphibians captures or observed along the Patuxent River
in the natural forest and four adjacent created wetland sites (1995/1996).
AMPHIBIANS
Natural
Forest
Site
A
Site
B
Site
C
Site
D
Wetland
Average
Wetland
std
P>t
Hyliidae
Acris
crepitans
12/0
10/2
27/8
0/2
9/0
14.5
14.29452
0.872301
Hyla
chrysoscelis
1/7
0/0
0/0
0/2
l/o
0.75
0.957427
*0.004777
Hyla
cinerea
0/3
0/2
0/0
0/2
0/0
1
1.154701
0.181690
Pseudacris
crucifer
3/0
7/0
7/0
0/0
5/0
4.75
3.304038
0.633044
Pseudacris
tnseriata
P
P
P
P
P
—
_
Bufonidae
Bufo
americanus
76/54
56/225 21/23
34/18
19/7
100.75
120.6576
0.824081
Bufo
fozvleri
6/2
0/83
0/10
3/63
0/10
42/25
37/88029 0.432556
Ranidae
Rana
catesbeiana
5/0
1/0
3/2
0/0
0/8
3/5
2/696846 0.712130
Rana
clamitans
61/2
24/9
18/20
18/5
10/6
27.5
9.882645
*0.036966
Rana
palustris
2/3
3/2
5/2
i/i
8/0
5.5
2.645751
0.86216
Rana
sphenocephala 57/24
15/82
33/85
6/101.
8/43
93.25
29.44345
0.705343
Pelobatidae
Scaphious
holbrookii
0/57
0/28
1/2
0/77
2/35
36.25
30.7395
0.548027
Ambystomatidae
Ambystoma
opacum 43/2
17/6
3/7
18/4
3/1
14.75
9.287088
*0.047231
Ambystoma
maculatum
0/1
0/0
0/0
0/0
0/0
0
0
_
Plethodontidae
Plethodon
glutinosus 0/19
0/6
0/7
0/2
0/1
4
2.94392
*0.014615
Salamandridae
Notophthalamus
viridescens 3/0
1/1
2/1
0/0
0/0
1.25
1.5
0.327652
P= species identified as present based on
als were neither seen nor captured.
calling observations, but individu-
Bulletin of the Maryland Herpetological Society page 105
Volume 38 Number 4
December 2002
coverage provided particularly good habitat for amphibians and thus, prob¬
ably served as the "source" site for amphibian colonization of the created
wetland sites. Abundant species in the Natural Forest Site, e.g. American toad,
green frog, southern leopard frog, spadefoot toad, marbled salamander, and
northern slimy salamander, were also seen in all four created wetland sites.
We note that the only spotted salamander (Amby stoma maculatum) captured
during the two-year study was found only in the natural forest.
In Site A, which at 228 m was the closest created wetland site to the
natural forest, we found 14 species of amphibians and 6 species of reptiles
(Tables 3 and 4). With an abundance of amphibian species— only 2 from the
natural forest site were absent — this site exhibited high amphibian richness
and diversity (Table 1). In fact, the capture rate for amphibians in this site
was greater than that for all other sites — including the natural forest. Three
species, American toad, Fowler's toad, and southern leopard frog, were es¬
pecially abundant. The numbers of reptile species were similarly high and
are mirrored in both high richness and diversity values (Table 1). These high
faunal diversities probably resulted from the fact that species could easily
move the short distance from the adjacent natural forest, the source of coloni¬
zation, and that the site retained water throughout the spring season, making
it attractive for both breeding and feeding.
Site B, located 321 m from the natural forest, revealed 13 amphibians
and 6 reptiles (Tables 3 and 4). A small pond located in this site provided a
source of water during dry periods and was a focus of much herpetofauna
activity (Table 3). This pond and its environs served as refugia for a number
of species, e.g. painted turtle laid eggs in adjacent sandy areas, green frog
and bullfrog ( Rana catesbeiana) tadpoles were observed in the waters, and
during the early spring, a southern leopard frog was observed emerging from
hibernacula. Because it retained water longer than others during the summer
season and provided adequate vegetative coverage, activity on this site was
high and overall diversity was similar to that for Site A (Tables 1 and 2).
Site C, was 361 meters from the natural forest, and only 12 amphibian
species and 2 reptile species were observed (Tables 3 and 4), resulting in the
lowest richness and diversity among all sites (Tables 1). None-the-less, Site C
must have contained some features not abundant in other sites as more
spadefoot toads were observed here than for the other sites (Table 3). It is
possible that the presence of breeding congregations of spadefoot toads re¬
sulted after heavy rains and by habitat features associated with the proxim¬
ity of an old sand quarry.
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Bulletin of the Maryland Herpetological Society
Volume 38 Number 4
December 2002
Site D, the furthest of the sites, at 373 m, from the natural forest with 13
amphibian species and 6 reptile species (Tables 3 and 4), displayed moderate
richness and diversity for amphibians and high richness and diversity for
reptiles (Table 1). Although documented in all study sites, pickerel frog, which
uses a variety of aquatic habitats (including bogs, seeps, grassy meadows,
and the margins and banks of marshes, swamps, brooks, and streams), was
more common here than in other sites. The diversity here may be the result of
a small patchy shrub-scrub area that provided shade for reptiles during the
Table 4. Number of reptiles captures or observed along the Patuxent River in
the natural forest and four adjacent created wetland sites (1995/1996).
REPTILES
Natural
Forest
Site
A
Site
B
Site
c
Site
D
Colubridae
Carphophis
amoenus
0/0
2/1
0/0
0/0
1/2
Nerodia sipedon
0/0
0/0
l/o
0/0
0/0
Virginia valeriae
0/0
2/0
0/0
0/0
0/0
Thamnophis
sirtalis
0/0
1/0
1/0
0/0
I/O
Heterudon
platirhinos
0/0
0/0
I/O
0/0
0/0
Scincidae
Eumeces fasciatus 2/0
0/0
0/0
0/0
0/0
Teiidae
Cnemidophorus
sexlineatus
0/0
0/0
0/0
0/0
1/0
Chelydridae
Chelydra
serpentina
I/O
0/0
1/1
0/1
0/1
Chrysemys picta
0/0
0/1
14/6
0/7
4/7
Emydidae
Terrepene Carolina 0/0
1/0
0/0
0/0
0/0
Kinostemidae
Kinosternon
subrubrum
1/0
0/1
1/3
0/0
2/1
Sternotherus
odoratus
1/0
0/0
0/0
0/0
0/0
Bulletin of the Maryland Herpetological Society page 107
Volume 38 Number 4
December 2002
day and was used by anurans for calling at all hours of the day and evening.
As in Site B, nests and egg hatchlings of painted turtles were found. Despite
the presence of shade and water, the lower richness and diversity, compared
to Site B, may be due to the distance from the colonizing source.
While it might generally appear that the number of species captured
and the species richness for each site was correlated with the distance from
the colonization source, specific trends are confounded by variation in veg¬
etation and the duration of standing water. Species richness and diversity
were significantly different among the sites (t-test; df - 3; p = 0.03). Species
richness of assemblages (Hmax) was significantly more diverse in the natural
forest than the created sites (t-test; df - 3; p = 0.04). For amphibians. Cope's
gray tree frog (Hyla chrysoscelis), green frog, marbled salamander (Ambystoma
opacum), and northern slimy salamander (Flethodon glutinosus) assemblages
were significantly greater in the natural forest (t-test; df = 3; p = 0.004, 0.03,
0.04, 0.01). For reptiles, no significant differences were detected in richness or
diversity between the natural forests and created wetland. During the two-
year study average of total captures for reptiles (15.75) was much smaller
than that seen for amphibians (350) (Table 2).
Conclusions
The creation of wetlands as a mitigation tool is increasingly important
as wetlands have been shown to play a major role in reducing the likelihood
of local extinction (Gibbs 1993) and may offset local extinction effects due to
wetland loss, as well as increasing levels of fragmentation (Gibbs 1998). Ex¬
amination and determination of critical features associated with successful
mitigation via wetland creation is, thus, of great usefulness.
We note that one of the major goals in using created wetlands for miti¬
gation purposes is the establishment of successfully reproducing populations.
It is not enough that species migrate to and occupy a created site, they must
also successfully reproduce. Herpetofaunal activity occurs within a variety
of habitats and microhabitats that provide features essential for foraging,
predator escape, thermoregulation, and reproduction. Successful reproduc¬
tion may require features quite different than those associated with foraging,
escape and thermoregulation, particularly for those species dependent upon
vernal pools, ponds, and vegetative coverage. For instance, created wetlands
that retain water in the spring but not into and through the summer and fall,
may provide features adequate for breeding but not, ultimately, for success¬
ful reproduction. Clearly, wetlands able to provide features critical for repro¬
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Bulletin of the Maryland Herpetological Society
Volume 38 Number 4
December 2002
duction are better suited for colonization and successful establishment of
amphibian and reptile populations.
With its greater habitat diversity, vegetative coverage, and biomass, the
natural forest provided source populations of amphibians and reptiles to the
newly created and previously unoccupied areas. The high levels of amphib¬
ian diversity within a year following site creation is testament to the nearby
location of a colonization source, the ability of these organisms to move dis¬
tances, and to the attractiveness of the created habitat, particularly for the
amphibians. The lower diversity of reptiles in both the natural forest and
created wetlands may simply be due to their generally more secretive nature
coupled with the difficulties of detecting them in mature forests (Gibbons
and Coker 1978; Bennett et al 1980).
The created wetland at Sands Road appears to have great potential as a
suitable habitat for amphibians. The presence of amphibians and reptiles in
all sites suggests that all sites provided microhabitats essentia) for successful
foraging and predator escape. However, the results of the present study sug¬
gest that amphibians responded most positively, e.g. bred successfully, in sites
where the hydroperiod was longest and where vegetative cover present. Resi¬
dent species and reproductive success was documented during the second
field season in sites B and D, the sites exhibiting the highest levels of diver¬
sity. The longer hydroperiods in both sites provided adequate habitat and
sufficient lime for successful breeding and reproduction to occur. The obser¬
vation and capture of large bullfrog tadpoles in site B indicates year-round
presence of water as the species can take two years to mejamorph. Although
not documented as successfully reproducing, the emergence of a southern
leopard frog from its winter hibernacula during the second field season is a
positive indication of features associated with habitat necessary for perma¬
nent residency
Physical habitat characteristics important to colonization should be ex¬
pected to change over time, leading to differential colonization and occupa¬
tion patterns. Examples include distance from the source, amount and depth
of standing water, water turbidity, and vegetation. As these change, so too
will the herpetofaunal community (Burke and Gibbons 1995; Thomas and
Barron 1995; Mitchell 1996; Gibbons et al. 1997). The appearance of northern
slimy salamanders in the second year in all five sites is suggestive of habitat
change as it was not seen in any of the sites in the first year. Another sala¬
mander, spotted salamander, may be of particular interest in designing and
Bulletin of the Maryland Herpetological Society
page 109
Volume 38 Number 4
December 2002
determining long-term habitat suitability of mitigated sites. Colonization by
this salamander is constrained by particular habitat requirements such as
mature forest, vernal pools and ponds and is probably not at all affected by
an inability to travel long distances (Semiitch 1998).
A five-year survey at the nearby (8 km S of the Sands Road site), 200 ha
Jug Bay Sanctuary (Smithberger and Swarth. 1993) revealed 39 species of
amphibians and reptiles: 18 amphibians (11 anurans, 7 salamanders) and 21
reptiles (7 turtles, 3 lizards, 11 snakes). These numbers are only slightly greater
than those documented during the two-year Sands Road study: 16 amphib¬
ians (12 anurans, 4 salamanders) and 12 reptiles (5 turtles, 2 lizards, 5 snakes).
The higher richness in the Jug Bay Sanctuary is likely due to greater variation
in habitat types that include freshwater tidal and non-tidal wetlands, upland
hardwood forest and agricultural fields. The majority of amphibians and rep¬
tiles recorded at Jug Bay were found within both upland and non-tidal wet¬
land areas, while all species recorded in Sands Road where in a palus trine
habitat within a young forest.
By monitoring the seasonal activity and utilization of the created wet¬
land, knowledge was gained on the essential factors to be considered when
constructing suitable habitats for successful colonization and reproduction
of amphibians, reptiles, and other wetland dependent species. Amphibians
warn us about our environment and by studying their colonization to cre¬
ated habitats more information can be attained which may allow us to make
intelligent recommendations on how to better preserve our natural environ¬
ment and wildlife.
It will be interesting to observe changes in herpetofaunal diversity as
the sites mature and the results of this study will provide baseline data for
future studies within the Sands Road Wetland Sanctuary.
Acknowdlegments
We would like to give special thanks to Matthew Perry for project sup¬
port, supplies, and encouragement. Steve Pugh's tireless efforts and assis¬
tance in putting in those drift fences. Steve Gotte and George Zug for their
expertise and assistance in the field. Robert Hoffman, R. McDiarmid, and J.
Mitchell for the never-ending support and encouragement. Robert Reynolds
for review of the manuscript. We also thank the Washington Biologists' Field
Club for its financial support.
page 110
Bulletin of the Maryland Herpetological Society
Volume 38 Number 4
December 2002
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2000a. Principles for management of aquatic-breeding amphibians.
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Wetlands Newktter Jammy - February 2000. pp. 5-6, 13.
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1998. Biological delineation of terrestrial buffer zones for pond¬
breeding salamanders. Conservation Biology 12(5):1113-1119.
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manders: individual repeatability and environmental influ¬
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1993. Reptiles and amphibians of the Jug Bay sanctuary. The Mary¬
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Volume 38 Number 4
December 2002
Stiling, P. D.
1992. Introductory Ecology , pp. 318-335. Prentice-Hall Inc.,
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Wake, D. B.
1991. Declining amphibian populations. Science 253:860.
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3 Li. S. Geological Survey /Biological Resource Discipline ,
Western Ecological Research Center San Diego Station - Irvine Office ,
2883 Irvine Blvdf Irvine, CA 92602
2 Department of Biology, Howard University, Washington, DC 20059
Received: 5 September 2002
Accepted: 28 October 2002
Bulletin of the Maryland Herpetological Society
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Volume 38 Number 4
December 2002
SPATIAL DISTRIBUTION IN A NEOTROPICAL
LIZARD, LIOLAEMUS QUILMES (LIOLAEMIDAE):
SITE FIDELITY AND OVERLAPPING
AMONG MALES AND FEMALES
Monique Halloy and Cecilia Robles
Abstract
We investigated spatial distributions, site fidelity, and overlap in a
neotropical lizard, Liolaemus quilmes. We selected a site (60x60m) within its
distribution in northwestern Argentina, Adults were captured, measured and
marked, and released at the site of capture. We monitored marked individu¬
als during three austral springs and summers. Both males and females showed
site fidelity having similar spatial distributions one year to the next. Male
activity areas were significantly larger than those of females and they over¬
lapped with one to two females. Females also overlapped with one to two
males. Finally, male activity areas overlapped considerably among themselves
whereas those of females did not overlap among themselves.
Little is known about the ecology of lizard species belonging to the
South American genus Liolaemus (Liolaemidae, Frost et aL, 2001, recently el¬
evated from its former status as a subfamily of the Tropiduridae, Frost and
Etheridge, 1989). Spatial distribution is one aspect that can give insight into
the intra- and intersexual interactions of a species and its relationship to re¬
productive strategies. By spatial distribution of a lizard, we considered the
entire area in which an individual moved, i.e., its activity area or home range
(Rose, 1982). Several studies on home range sizes in lizard species of North
America are available (e.g.. Tinkle et al., 1962; Ferner, 1974; Stamps, 1977; Fair
and Henke, 1999; Sheldahl and Martins, 2000) but few have been undertaken
on South American species. Ortiz (1981) reported home range sizes for some
species belonging to the Liolaemus nigromaculatus group and Rocha (1999)
gave home range sizes for Liolaemus lutzae. In this study, we investigate the
spatial distributions, site fidelity and overlap in male and female Liolaemus
quilmes during the austral spring and summer of three consecutive years at a
location in northwestern Argentina.
Liolaemus quilmes , one of more than 160 species in this neotropical
genus (Schulte et al., 2000), is a diurnal, mainly insectivorous, and oviparous
species (Ramirez Pinilla, 1992). It belongs to the darwinii complex (Etheridge,
page 118
Bulletin of the Maryland Herpetological Society
Volume 38 Number 4
December 2002
1993) and it occurs in northwestern Argentina (southern part of Salta prov¬
ince, northwestern part of Tucuman province, and northeastern part of
Catamarca province). The species ranges from about 1600 m to just below
3000 m, occupying arid to semi-arid habitats. Males are slightly larger and
more colorful than females (Etheridge, 1993).
Materials and Methods
Our study site was located at Los Cardones (2700m), Tucuman prov¬
ince, Argentina, well within the distribution of this species. The site is charac¬
terized by firm substrate, large rocks, shrubs and cacti. It corresponds to the
semi-arid Prepuna phytogeographic region of Cabrera and Willink (1980).
We measured a grid of 60x60m, subdividing it into 5x5m quadrats using out¬
door paint to mark rocks with the corresponding coordinates.
The study took place during the austral spring and summer of 1999-
2000 (November to March, referred to as the first period), 2000-2001 (Septem¬
ber to April, second period), and 2001-2002 (October to March, third period).
Almost all the lizards were captured and marked during the first two peri¬
ods (first period: 25 males and 21 females; second period: 22 males and 13
females; third period: 1 male and 1 female). Upon capture, lizards were mea¬
sured and weighed (Table 1). They were then marked with a unique combi¬
nation of two colored beads attached at the base of the tail with a surgical
steel monofilament strand (Fischer and Muth, 1989). All elements used as
well as the area of insertion of the strand were disinfected with alcohol. Marked
lizards were released at the site of capture. This procedure did not seem to
TABLE 1 - Means and standard deviations of snout-vent lengths (SVL),
total lengths (TL), and weights (W) for male and female Liolaemus quilmes
from Los Cardones, Tucuman province, Argentina.
Males (n = 48)
Females (n = 35)
SVL (cm)
6.13 ± 0.48
5.72 ±0.37
TL (cm)
14.32 ±2.27
13.27 ±1.80
W(g)
6.76 ±1.62
5.66 ±1.16
Bulletin of the Maryland Herpetological Society
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Volume 38 Number 4
December 2002
impair the lizards, since they were often found again in the same area soon
after release as well as weeks and months later.
The site was visited two consecutive days per month during the first
two periods and it was visited 12 days, end of October to the first week of
November, and 5 days in March during the third period. During each visit,
we searched for marked lizards at 10:00, 13:00 and 16:00 hours. We walked
systematically, following the established coordinates, the starting point be¬
ing selected randomly before each search. We used binoculars to identify liz¬
ards. When we sighted a lizard, we took note of its identity and its coordi¬
nates.
We used the program CALHOME (A Home Range Analysis Pro¬
gram, MS-DOS Version 1.0, 1994) which uses the minimum convex polygon
method to calculate home ranges or activity areas. Following Rose (1982),
regression analyses were made for males and females, separately, obtaining
in each case that 9 sightings was the minimum number of sightings not corre¬
lated with area (F = 2,37, df = 1, 51, p > 0.05, one-tail; F = 0,47, df = 1, 31, p >
0.05, one tail, respectively). Of the total of 48 marked males, we obtained
activity areas for 43 males (based on a minimum of 4 or more sightings), 21 of
TABLE 2 - Means (X) standard deviations (SD), and ranges of areas (in
m2) occupied by male and female Liolaemus quilmes during different years at
Los Cardones, Tucuman province, Argentina.
N: number of lizards foi which we had a minimum of 9 or more
sightings.
Males
Females
Period
N
X±SD
Range
N
X±SD
Range
November 1999
to March 2000
2
77.0 ±46.0
44.5-109.5
0
- —
—
September 2000
to April 2001
6
132.2 ±82.7
22.5 - 242.5
2
29.2 ±9.5
22.5-36.0
October 2001
to March 2002
13
182.5 ±109.4 66.0-379.5
8
21.9 ±12.6
5.5-28.5
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Bulletin of the Maryland Herpetological Society
Volume 38 Number 4
December 2002
which had 9 or more sightings. Of the total of 35 marked females, we ob¬
tained activity areas for 30 females (based on a minimum of 4 or more
sightings), 10 of which had 9 or more sightings. Although we considered
lizards with fewer than 9 sightings to show the location of the various lizards
in the grid (see results), in the data analyses, we used only the individuals for
which we had a minimum of 9 or more sightings. For example, we compared
areas of different periods only if we had a minimum of 3 individuals with 9
or more sightings. We used the Wilcoxon-Mann-Whitney test (Siegel and
Castellan, 1988).
Results
Activity areas varied across periods particularly for males (Table 2).
Nevertheless, areas for males of the second and third periods were not sig¬
nificantly different (z = -0.88, p = 0.38, two-tailed, Wilcoxon-Mann-Whitney
test). On the other hand, areas for males were significantly larger than those
of females during the third period (z = -3.8, p < 0.001, two-tailed, Wilcoxon-
Mann-Whitney test, Siegel and Castellan, 1988). Areas of males were on aver¬
age 4.5 times larger than those of females during the second period and on
average 8.3 times larger during the third period (Table 2).
Males as well as females were found to remain in a similar area across
the three periods of study (Figs. 1 and 2, respectively). Although we did not
always have a minimum of 9 sightings per lizard per period of study, the area
they occupied year after year still reveals permanence in a certain space. This
can be seen for 11 males, five of them during three consecutive years and the
rest during two consecutive years (Fig. 1). In females, six were found in con¬
secutive years occupying similar locations, two of them during three con¬
secutive years (Fig. 2).
Male activity areas overlapped among themselves considerably dur¬
ing the second and third period (Figs. 3 and 4, respectively). For the first
period, we had fewer marked males and consequently there was less over¬
lap. Therefore, a graphic representation is not provided here. Because we did
not have many males with a minimum of nine or more sightings, we did not
calculate overlap percentage since activity areas or home ranges based on
fewer than nine sightings are probably underestimated (Rose, 1982). Female
activity areas, on the other hand, did not overlap among themselves in none
of the three periods of study although they occasionally came in contact (Figs
3 and 4, second and third period, respectively). There was one exception: two
Bulletin of the Maryland Herpetological Society
page 121
Volume 38 Number 4
December 2002
FIGURE 1. - Site fidelity in male Liolaemus quilmes during three con¬
secutive periods (described in Methods). Individual males are identified by
two-letter codes and a number that corresponds to one of the three periods of
the study. The first period is noted by small dashed lines, the second period
by larger dashed lines and the third by full lines. Activity areas are based on
a minimum of 4 or more sightings and were calculated with the minimum
convex polygon method. Axes are in meters.
females overlapped within the activity area of male FN during the third pe¬
riod (Fig. 4).
During the second period, of 17 males, 10 were found to overlap
with one female, 1 with 2 females, and 6 did not overlap with any female. Of
the 10 females for that same period, 4 had activity areas overlapping with
one male, 4 with 2 males, and two did not overlap with any male (Fig. 3).
page 122
Bulletin of the Maryland Herpetological Society
Volume 38 Number 4
December 2002
FIGURE 2. - Site fidelity in female Liolaemus quilmes during three con¬
secutive periods (described in Methods). Individual females are identified
by two-letter codes and a number that corresponds to one of the three peri¬
ods of the study. The first period is noted by small dashed lines, the second
period by larger dashed lines and the third by full lines. Activity areas are
based on a minimum of 4 or more sightings and were calculated with the
minimum convex polygon method. Axes are in meters.
During the third period, of 17males, 8 had activity areas overlap¬
ping that of one female, 5 overlapped with 2 females, 1 overlapped with 3
females, 2 came in contact with a female activity area, and one did not over¬
lap any female activity area. Of the 15 females, 6 had activity areas overlap¬
ping that of one male, 7 that of 2 males, 1 that of 3 males, and one came in
contact with a male activity area (Fig. 4).
Bulletin of the Maryland Herpetological Society
page 123
Volume 38 Number 4
December 2002
FIGURE 3. - Activity areas of male (full line) and female (dashed line)
Liolaemus quilmes, during the second period of observation (spring and sum¬
mer of 2000-2001). Activity areas are based on a minimum of 4 or more
sightings and were calculated with the minimum convex polygon method.
Individuals with 9 or more sightings are identified by two-letter codes. Axes
are in meters.
Activity areas of male Liolaemus quilmes were significantly larger than
those of females. Our results are similar to those obtained for other iguanian
species in that males tend to have larger home ranges than females, e.g., Uta
stansburiana stejnegeri (Tinkle et al., 1962), Liolaemus kuhlmani (Ortiz, 1981),
Tropidurus torquatus (Giaretta, 1996), Sceloporus virgatus (Abell, 1999).
Both males and females remained in the same area throughout their
active season, some of them being sighted during the three periods of the
page 124
Bulletin of the Maryland Herpetological Society
Volume 38 Number 4
December 2002
FIGURE 4. - Activity areas of male (full line) and female (dashed line)
Liolaemus quilmes, during the third period of observation (spring and sum¬
mer of 2001-2002). Activity areas are based on a minimum of 4 or more
sightings and were calculated with the minimum convex polygon method.
Individuals with 9 or more sightings are identified by two-letter codes (two-
letter codes written in bold correspond to females). Axes are in meters.
study. This site fidelity has been observed in other similar sized iguanian
lizards, e.g., Sceloporus undulatus erythrocheilus (Ferner, 1974); Uta palmeri
(Hews, 1993); Sceloporus occidental is (Sheldahl and Martins, 2000).
Male activity areas overlapped considerably during the second and
third period of the study. Sheldahl and Martins (2000) recorded an overlap of
41.5% in Sceloporus occidentalis, and Ferner (1974) of 52% in Sceloporus undulatus
erytrocheilus. However, other authors have found that, for similar sized
iguanians (e.g., Uta stansburiana stejnegeri, Tinkle et al., 1962; Uta palmeri , Hews,
Bulletin of the Maryland Herpetological Society
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Volume 38 Number 4
December 2002
1993; Liolaemus lutzae, Rocha, 1999), home ranges in males overlapped little
or not at all, possibly indicating territoriality (Tinkle et al., 1962; Stamps, 1977;
Rocha, 1999). Other species of Liolaemus belonging to the nigromaculatus group
have been characterized as either territorial, hierarchical or neither (Ortiz,
1981). In L. quilmes, males were seen patrolling their activity areas by stand¬
ing on rocks, visually scanning the surroundings, legs outstretched and head
bobbing. Visual displays (such as head bobs, lateral presentation, inflation of
the body) were directed at approaching males, occasionally resulting in chas¬
ing and fighting (some of these behavioral patterns have been described for
this species in Halloy, 1996, and for other iguanian species in Carpenter and
Ferguson, 1977). This suggests a hierarchical rather than a territorial system
in this species.
In her review of territorial lizard species. Stamps (1983) found that
female home ranges either do not overlap among themselves or do so very
little. In our study, female home ranges or activity areas did not overlap (ex¬
cept in one case, during the third period). When kept in an outdoor, 4x5m
enclosure, females of this species were found to chase and fight any other
female close by (Halloy, 1996). These traits (small non-overlapping activity
areas and agonistic behavior among females when at close range) suggest
territoriality in these females (as has been shown in many other iguanian
species, Stamps, 1983; but see Hews, 1993, who observed considerable over¬
lap and lack of territoriality in female Uta palmeri).
Male activity areas overlapped those of one to two females. Females
as well, overlapped with one to two males. This is similar to what Rose (1982)
and Abell (1999) found in Sceloporus virgatus in which males overlapped with
two or more females and females with about two males suggesting a
polygynandrous mating system. Although we need more observations, es¬
pecially behavioral, L. quilmes may perhaps be seasonally monogamous (same
male-female pairs were often seen basking on the same rock) as has been
reported in Liolaemus copiapensis (Ortiz, 1981) and in Uta stansburiana (Tinkle
et al. 1962; Fox, 1983; Stamps, 1983), or polygynandrous as in Sceloporus virgatus
(Rose, 1982; Abell, 1999).
Acknowledgements
We are grateful to S. Downes, S. Fox, E. Martins, and anonymous
reviewers for valuable comments on an earlier draft. We thank Marcela
Castillo, Fabricia Guglielmone, Luciana Marangoni, Cecilia Guerra, and Paul
page 126
Bulletin of the Maryland Herpetological Society
Volume 38 Number 4
December 2002
Grosse for field assistance. We thank Recursos Naturales y Suelos of the
Tucuman province (permits # 394-98 and 95-2000) for permission to work in
the field. CONICET (PIP # 4966/97) provided financial support to MH.
Literature Cited
Abell, A. J.
1999. Male-female spacing patterns in the lizard, Sceloporus
virgatus. Amphibia Reptilia 20: 185-194.
Cabrera, A. L., and A. Willink.
1980. Biogeografia de America Latina. Secretaria General de la
Organizacion de los Estados Americanos. Programa Re¬
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Carpenter, C. C. and G. W. Ferguson.
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Etheridge, R.
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Iguania: Tropiduridae) in Northern Argentina. Boll. Mus.
Reg. Sci. nat. Torino 11(1): 137-199.
Fair, W. S. and S. E. Henke.
1999. Movements, home ranges, and survival of Texas horned liz¬
ards (Phrynosoma cornutum). J. Herpetol. 33(4): 517-525.
Ferner, J. W.
1974. Home range size and overlap in Sceloporus undulatus
erythrocheilus (Reptilia: Iguanidae). Copeia 1974(2): 332-337.
Fischer, M., and A. Muth.
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20: 45-46.
Fox, S. F.
1983. Fitness, home-range quality, and aggression in Uta
stansburiana. In: Lizard Ecology, (Eds. R. B. Huey, E. R.
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Pianka & T. W. Schoener). Harvard University Press, Cam¬
bridge, Massachusetts, pp. 149-168.
Frost, D. R., and R. Etheridge.
1989. A phylogenetic analysis and taxonomy of iguanian lizards
(Reptilia: Squamata). Miscellaneous Publications Museum
of Natural History, University of Kansas 81: 1-65.
. R. Etheridge, D. Janies and T. A. Titus.
2001. Total evidence, sequence alignment, evolution of
polychrotid lizards, and a reclassification of the Iguania
(Squamata: Iguania). American Museum of Natural History
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Halloy, M.
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Hews, D. K.
1993. Food resources affect female distribution and male mating
opportunities in the iguanian lizard Uta palmeri. Anim.
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Ortiz, J. C.
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Ramirez Pinilla, M. P.
1992. Ciclos reproductivos y de cuerpos grasos en dos poblaciones
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1999 . Home range of the Tropidurid lizard Liolaemus lutzae : sexual
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Rose, B.
1982. Lizard home ranges: Methodology and functions. J.
Herpetol. 16(3): 253-269.
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Institute de Herpetologia , Fundacion Miguel Lillo, Miguel Lillo 251 , 4000 San
Miguel de Tucumdn , Argentina, mhalloy@unt.edu.ar (MH); Facultad de Ciencias
Naturales, Universidad Nacional de Tucumdn, Miguel Lillo 205, 4000 San Miguel
de Tucumdn , Argentina, ceciro@tucbbs.com.ar (CR).
Received: 12 October 2002
Accepted: 21 October 2002
Bulletin of the Maryland Herpetological Society
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December 2002
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Volume 39 Number 1
March 2003
Range Extensions and Variational Notes on
Some Amphibians and Reptiles of Jalisco and
Michoacan, Mexico
Paulino Ponce-Campos, Sara M. Huerta-Ortega , Alan Heinze-Yothers and Hobart
M. Smith
Abstract
Herpetological studies in four areas of central western Mexico (the Barranca
of the Rio Santiago, central Jalisco; Guadalajara city, central Jalisco; Sierra de Quila,
central Jalisco; and Sierra de Coalcoman, northwestern Michoacan) have produced
extensions of ranges of Eleutherodactylus pallidus, Eumeces dugesii, Boa constric¬
tor imperator, Leptodeira bressoni, Ramphotyphlops braminus , Salvadora mexicana
and Storeria storerioides. Variational data on these taxa and Enulius oligostichus are
presented.
Introduction
Long-term, on-going studies have been conducted by PP-C and his associates
on the herpetofauna of Jalisco and part of neighboring states, especially on western
slopes.
Four areas have been of special interest, as follows. Of greatest geographic
importance is the is the Barranca of the Rio Santiago, a corridor for the passage of
coastal, low-altitude species eastward toward the plateau, and of plateau species west¬
ward toward the coast. The Barranca has not been well collected, hence its full bio¬
geographic importance remains to be realized. Preliminary efforts were reported by
Ponce-Campos (submitted), Ponce-Campos and Heurta Ortega (1998 and in prerp.)
and Ponce-Campos et al. (2001). We here report range extensions in this area for
Eleutherodactylus pallidus, Boa constrictor imperator and Salvadora mexicana .
The herpetology of the Guadalajara area has been studied sporadically by nu¬
merous authors, among whom Duges is perhaps the earliest, but never synoptically
for that particular area. Perhaps the nearest is in Duges (1889), comparing the
herpetofauna (and other faunae) of Guadalajara with that of Guanajuato. The scat¬
tered records existing at the time were included among those for Mexico as a whole
as early as Cope (1887) and Duges 1896), and as recently as the checklists by Smith
and Taylor (1945, 1948, 1950). Zweifel (1959) examined some questionable early
records for the area.
Bulletin of the Maryland Herpetological Society
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March 2003
The isolated Sierra de Coalcoman is still poorly known despite its proven en-
demicity (e.g., Barisia jonesi Guillette and Smiths 1982 (given specific rank in Smith
et ah 2002); Coniophanes sarae (Ponce-Campos and Smith, 2001 ; Sceloporus insignis
Webb, 1967; Geophis pyburni Campbell and Murphy, 1977, and others). We here
report a range extension of Storeria storerioides in that area.
The Sierra de Quila of central Jalisco has also been collected little in the past.
We here report Eumeces dugesii from there.
Additionally, we here describe the variation in a recently collected specimen
of Enulius oligostichus .
Specimens cited are in the collection of Bosque Tropical A.C., Guadalajara,
Jalisco (BTM), and in the Museum of the Universidad Autonoma de Guadalajara
(UAGM).
Species Accounts
Eleutherodactylus pallidus (Buellman) (Pale Chirping Frog). Found in an area
of subtropical scrub forest, by Alan Heinze-Yothers and Paulino Ponce-Campos June
8, 2000 at 705 m» 20° 56' N, 103° 37' W, specimen BTM-007. The locality is a range
extension 107 km southeast from 30 mi southeast of Tepic, Nayarit (Davis and Dixon,
1957) and 130 km east, from 18.8 mi NW Ahuacatlan, Nayarit (Lynch, 1970), PP-C
found the species at Puerto Vallarta, on the coast of Jalisco. Two specimens are larger
than others recorded of the species. Lynch (1970) recorded the maximum size as
19,3mm. Gur specimen is 22.2mm in SVL and weighed 0.8g. Another specimen
taken in the same area 27 June 2000 and released in situ measured 20.6mm in SVL,
weight 0.6g. Both specimens were calling males.
Eumeces dugesii Thommot (Duges's Skink). Four examples were measured
and photographed (BTM008) in the field and released in the area of Sierra de Quila,
Jalisco, 2,078 m (20° 18' N & 104° 04' W) by PP-C, Rodolfo and Hector Romero-
Contreras and M. en C. Rodolfo Romero-Luna, They were found in oak and pine
forest between 2,078-2, 228m altitude. Three localities are recorded for the species in
Jalisco: 3 mi WSW Mazamitla, Nevado de Colima and Ajijic in Chapala Lake (Peterson
et al, 1995). Our records are 88 km WNW from Ajijic, heretofore the northernmost
record.
Boa constrictor imperator Duadin (Mexican Boa Constrictor), One specimen
(BTM002) was obtained by a local resident, Pedro Esparza Gonzales, from the mu¬
nicipality of Zapopan near Guadalajara, 1074 m (20° AT N, 103° 19* W). It was
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Bulletin of the Maryland Herpetological Society
Volume 39 Number 1
March 2003
found in an area of disturbed subtropical scrub forest during early August 2001 . This
extends its known range from the lowland coasts over all of Jalisco (Garcia and
Ceballos, 1994).
Enulius oligostichus Smith, Arndt and Sherbrooke (Mexican Longtailed
Snake). A specimen of this species will be reported for the first time in Jalisco by
Ponce-Campos (submitted). It was found in disturbed subtropical scrub forest at an
altitude of 1189m (BTM-003). Other records are from lowlands, even near sea level
(Smith et al, 1967; McDiarmid and Bezy, 1971). It is smaller than others reported, at
116mm. The tail is incomplete, at 25mm.
Due to the rarity of the species, the variation exhibited by this specimen is of
special interest. It differs from previous descriptions in having a higher ventral count
(166) (subcaudals 47, tail incomplete); anterior and posterior temporals fused on the
left side (separate on right); supraocular fused with parietal on the right side (separate
on left).
As in other specimens, the dorsal scales are smooth, with a single apical pit;
five supralabials; six infralabials; nasal fully divided; scale rows 15-15-13, the latter
count at the vent. The size and arrangement of the anterior and posterior chinshields,
the shape and size of the maxillary teeth, and the position of the anterior tip of max¬
illa between first and second supralabials, are as in the original description (Smith et
al., 1967).
In life the head is brown, a little darker than the dorsum of the body (gray-
brown); latter color lighter on sides; venter light gray.
Leptodeira bressoni Taylor (Bresson’s Splendid Cat-eyed Snake). A specimen
(BTM-004) was found near Guadalajara in the Barranca del Rio Santiago at 1080m
(20° 46' N, 103° 19' W), 5 March 2002 by a local resident, Rogelio Contreras Esparza.
Also four specimens were found in the canyon by PP-C, AH-Y and Carlos Morfm
(data recorded in the field and released, 21 April 2000; 7 30-75 0mm 20° 54' N, 103°
37' W). These are first known for the area of our interest. Previous records in the state
are by Duellman (1961) at Ranco El Rodeo, Sierra de Ixtlan near San Marcos and
18.4 mi NW Magdalena. The latter two are the nearest to our area, 46km NNE and
71km NW. The Ajijic record (Chapala shore) by Peterson et al. (1995), is the east¬
ernmost record for the species in the state.
Ramphotyphlops braminus (Braminy Blind Snake). This species is con¬
sidered the most abundant species of snake in Guadalajara City (Ponce and Huerta,
in prep.). Examples have been found throughout the city since 1990, even down-
Bulletin of the Maryland Herpetological Society
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Volume 39 Number 1
March 2003
town in the most unlikely habitat, polluted and almost without vegetation (BTM-
012). Some were found near the city, in the Santiago River Canyon, where there
is less human influence, by Adrian Jacobo-Contreras and other local residents, 28
August 2002, 20° 47 N, 103° 20' W (BTM-011), 11 October 2001, 20° 47 N, 103°
19' W, 1 160 m (BTM-013-14). This year at least 6 examples were found in one week
during the rainy season, in August.
The oldest known discovery in the state is from El Chante (Chapala lake shore),
municipality of Jocotepec, Jalisco, a specimen collected by Lila Dipp (a biology
student in UAG at that time) in 1980 (UAGM R-0188). Some other UAGM speci¬
mens from the Guadalajara area include Colonia Lomas del Valle, 12 March 1992
(UAGM R-0249); nr Glorieta Minerva, 19 September 1991 (UAGM R- 173-4); and
Colonia Las Puentes, 7 October 1991 (UAGM R-0241-2). One record is from the
southern coast of Jalisco at Barra de Navided, Jalisco, by SMH-0 22 August 1990
(UAGM R-170). Thirteen- more specimens from Guadalajara are in UAGM and BTM.
The first record for the species in the state was for Puerto Vallarta (Dundee and
Flores, 1991). The species is now known throughout the central, northern and sourthem
parts of Jalisco.
Salvadora mexicana (Dumeril, Bibron and Dumeril) (Mexican Patchnosed
Snake). One (. BTM -001) was obtained 8 December 2001, by Mario-Ruvalcaba-
Venegas in the municipality of Zapopan, near Guadalajara, in an area of disturbed
subtropical scrub forest at 1159m (20° 47' N, 103° 1 9' W). Fordmer records for the
species in the state are limited to coastal areas (Garcia and Ceballos, 1994).
Storeria storeioides (Cope) (Mexican Brown Snake). Two specimens were
found in the municipality of Chinicuila, Michoacan. One was in an area of cloud
forest at 1680m (18° 4F N, 103° 25* W), taken by a local resident, Jesus Bravo-
Sanchez, 3 December 2000 (BTM-005). The second was found in oak-pine forest at
1481m (18° 42' N, 103° 23' W), 4 December 2000 (BTM-006), and was photographed
and released in situ. These are the westernmost records in Michoacan; the nearest
record to these localities is about 50km NE, at Dos Aguas, Michoacan (Duellman,
1961).
Acknowledgments.
We thank the Departamento de Ecologfa y Desarrollo Agropecuario del
municipio de Zapopan, and Eduardo Saha gun, for financial support; Rodolfo and
Hector Romero-Contreras and M. en C» Rodolfo Romero-Luna, Jesus Bravo-Sanchez,
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Bulletin of the Maryland Herpetological Society
Volume 39 Number 1
March 2003
Nicanor Mendoza-Mendoza and Paulino Ponce- Villa for field assistance, also pro¬
vided by the residents of ex-Hacienda del Lazo.
Literature Cited
Campbell J. A. and J. B. Murphy.
1977. A new species of Geophis (Reptilia, Serpentes, Colubridae) from
the Sierra de Coalcom&n, Michoacdn, Mexico. J. Herp. 11(4): 397-
403.
Cope, E. D.
1887. Catalogue of batrachians and reptiles of Central America and
Mexico. Bull. U. S. Nat. Mus. (32); 1-98.
Davis, W. B. and J. R. Dixon.
1957. Notes on Mexican amphibians, with description of a new
Microbatrachylus. Herpetologica 13: 145-147.
Duellman, W. E.
1961. The amphibians and reptiles of Michoacan, Mexico. Univ. Kan¬
sas Publ. Mus. Nat. Hist. (15): 1-148.
Duges, A. A. D.
1896. Reptiles y batracios de los Estados Unidos Mexicanos. La
Naturaleza (2)2: 479-485.
Dundee, H. A. and O. Flores-Villela.
1991. Ramphotyphlops braminus. Herp. Rev. 22(1): 26.
Garcia, A. and G. Ceballos.
1994. Gufa de campo de los reptiles y anfibios de la costa de Jalisco.
Fundacion Ecologica de Cuixmala, A. C. and Institute de Biologia,
UNAM, Mexico. 184 p.
Guillette, L. J., Jr. and H. M. Smith.
1982. A review of the Mexican lizard Barisia imbricata, and the de¬
scription of a new subspecies. Trans. Kansas Acad. Sci. 85: 13-
33.
Lynch, J. D.
1970. A taxonomic revision of the leptodactylid frog genus Syrrhophus
Cope. Univ. Kansas Publ. Mus. Nat. Hist. 20 (1): 1-45.
Bulletin of the Maryland Herpetological Society
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Volume 39 Number 1
March 2003
McDiarmid, R. W. and R. L. Bezy.
1971. The colubrid snake Enulius oligostichus in western Mexico.
Copeia, 1971: 350-351.
Peterson, J. W., H. M. Smith and D. Chizsar.
1995. Some noteworthy amphibians and reptiles from the region of
Chapala, Jalisco, Mexico. Bull. Chicago Herp. Soc. 30: 90-91.
Ponce-Campos, P.
(submitted). Geographic distribution: Enulius oligostichus. Herp Rev.
Ponce-Campos, P. and S. M. Huerta-Ortega.
(in prep.). Anfibios y reptiles de la zona conurbana de Guadalajara y su
perifena. Analisis preliminar. In: Ecologia en Guadalajara.
Ponce-Campos, P. and S. M. Huerta-Ortega.
1998. Geographic distribution: Dryadophis cliftoni. Herp. Rev. 29:176.
Ponce-Campos, P. and S. M. Huerta-Ortega, C. Nogueira-Gomez and H. M. Smith.
200 1 . Natural history notes on the Southern plateau night lizard, Xantusia
sanchezi. Bull. Maryland Herp. Soc. 37: 18-21.
Ponce-Campos, P. and H. M. Smith.
2001. A review of the Stripeless snake (Coniophanes lateritius) com¬
plex of Mexico. Bull. Maryland Herp. Soc. 37: 10-17.
Smith, H. M., R. G. Arndt and W. C. Sherbrooke.
1 967. A new snake of the genus Enulius from Mexico. Nat. Hist. Misc.
Chicago Acad. Sci. (186): 1-4.
Smith, H. M., T. M. Burg and D. Chiszar.
2002. Evolutionary speciation in the alligator lizards of the genus Barisia.
Bull. Maryland Herp. Soc. 36: 23-26.
Smith, H. M. and E. H. Taylor.
1 945. An annotated checklist and key to the snakes of Mexico. Bull. U.
S. Nat. Mus. (187): i-iv, 1-239.
Smith, H. M. and E. H. Taylor.
1 948. An annotated checklist and key to the amphibia of Mexico. Bull.
U. S. Nat. Mus. (194): i-iv, 1-118.
page 6
Bulletin of the Maryland Herpetological Society
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March 2003
Smith, H. M. and E. H. Taylor.
1950. An annotated checklist and key to the reptiles of Mexico exclu¬
sive of the snakes. Bull. LJ. S. Nat. Mus. (199): i-vi, 1-253.
Webb, R. W.
1 967. Variation and distribution of the iguanid lizard Sceloporus bulleri,
and the description of a related new species. Copeia 1967: 202-
213.
Zweifel, R. G.
1959. The provenance of reptiles and amphibians collected in western
Mexico by J. J. Major. Am. Mus. Novit. (1949): 1-9.
PP-C, SH-O, AH-Y - Bosque Tropical , A. C, Apartado Postal 5-515, Gudalajara,
Jalisco 45042, Mexico, poncecp@hotmail.com
HMS - E.P.O. Biology and Museum, University of Colorado, Boulder, Colorado,
U.S.A. 80309-0334.
Received 1 6 November 2002
Accepted 20 November 2002
Bulletin of the Maryland Herpetological Society
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March 2003
A Note Regarding Defensive Behavior in The Short¬
headed Gartersnake (Thamnophis brachystoma).
Limited information exists regarding antipredator or defensive behavior in
Thamnophis brachystoma. In their monograph on the gartersnakes, Rossman et al.
(1999) makes no mention of antipredator behavior for T. brachystoma. Tennant and
Bartlett (2000) report that this species is excitable, and not prone to bite. Hulse et al.
(2001), in addition to commenting on the short-headed gartersnake never attempt¬
ing to bite, also states that it will often thrash about violently and release feces and
musk. Here I report defensive behavior for T. brachystoma, which has not been
reported . The observation was made on 6 May 2002, ca. 1 100 h, just south of PA 36,
and about 12 meters west of the Clarion river at Cook Forest State Park, Cooksburg,
Pennsylvania.
An adult female T. brachystoma was found beneath a small flat rock. The
snake immediately attempted to flee, making rapid undulations with its body. Upon
being grasped, it thrashed about, musked, and voided the contents of its cloaca. After
ca. 10 seconds of energetic writhing, the individual flattened its body dorsoventrally,
opened its mouth, and began repeatedly striking at my left hand. Most of the strikes
failed to make contact. Although it struck with its mouth open it never closed its
mouth upon hitting my hand, and therefore did not bite. Additionally its teeth never
caught onto my skin, nor caused any abrasions. It continued to display this behavior
for approximately one minute, until it was placed on the ground and allowed to crawl
beneath a rock. Flattening of the body caused the snake to appear larger than it was;
whereas open-mouth strikes gave the snake a more threatening appearance. It would
be particularly interesting to investigate the prevalence of these behaviors in this
population, and also to determine if the behaviors are unique to T. brachystoma in
this locality.
A second T. brachystoma, a mature male, was found in the same area. This
individual did not display any behavior that was atypical for the species.
Acknowledgments
I wish to thank W. Shane Snyder for accompanying me in the field. I would
also like to thank the following individuals for reviewing the manuscript and offering
comments and or suggestions, Arthur C. Hulse, James Ball, David Chiszar, and Hobart
Smith.
page 8
Bulletin of the Maryland Herpetological Society
Volume 39 Number 1
March 2003
Literature cited
Hulse, A. C., C. J. McCoy, and E. J. Censky.
2001. Amphibians and Reptiles of Pennsylvania and the Northeast.
Cornell University Press, Ithaca, NY. 419 pp.
Rossman, D. A., N. B. Ford, and R. A. Seigel.
1999. The Garter Snakes: Evolution and Ecology. University of Okla¬
homa Press, Norman, OK. 332 pp.
Tennant, A. and R. D. Bartlett.
2000. Snakes of North America: Eastern and Central Regions. Gulf Pub¬
lishing, Houston, TX. 588 pp.
BrianS. Gray, 1217 Clifton Drive, Erie, Pennsylvania, 16505-5215.
Received: 20 November 2002
Accepted: 4 December 2002
Bulletin of the Maryland Herpetological Society
page 9
Volume 39 Number 1
March 2003
BOOK REVIEW: Wisconsin’s Natural Communities: How to Recognize
them3 Where to Find Them , by Randy Hoffman, 2002. University of Wisconsin
Press, 1930 Monroe St., Madison, Wisconsin, 53711. ISBN 0-299-17080-2, Cloth
$59.95, ISBN 0-299-17084-5, Paper $24.95.
The author has intended this book primarily for the layman, as scientific names
have been excluded from within the text, although the appendix gives both the com¬
mon and scientific names for those species of mosses, lichens, liverworts, mollusks,
snails, and insects not cited in single sources within the text. It truly would have been
a more informative volume had the author provided scientific names along with the
common names cited throughout the text. Everyone familiar with plants, inverte¬
brates or vertebrate species knows that common names are very unstable, and vary
significantly within regions.
The Introduction provides a brief review of the species diversity found within
the state of Wisconsin, with insects numbering nearly 40,000 species, while the am¬
phibians and reptiles are lesser represented, with some 53 species. A comparison of
acres of the 28 plant communities found within Wisconsin shows that since the mid-
1 880’s, only the Southern Red-Oak mixed forest and Dry Pine forest habitats have
increased in dominance into the late 1900’s. All other communities have decreased
significantly, with Sugar-Maple-Basswood, Mesic Prairie, and Wet-mesic Prairie
habitats having been drastically affected. As one would easily surmise, agricultural
and urban commuities have been the major cause for natural community destruction.
The author divided the book into two sections, with the first section describing
the natural communities found within the state of Wisconsin, and part two which
describes the 50 sites discussed within the text. Each site is well- illustrated with line
drawings and maps which depict the specific locality, along with road direction for
each community treated within the text, followed by a list of noteworthy species of
plants, insects, herptiles, birds and mammals that are noted for the specific area. The
author also provides valuable information on population declines for specific spe¬
cies, along with remarks on habitat destruction, and its implications, and general
information on potential species to be encountered in different cite locations.
Eight pages of high quality color plates provide a select overview of certain
communities and species, although this coverage could easily have been expanded
by eliminating some of the line drawings of specific plants and animals throughout
the text, which are of little value.
Overall this excellent book will prove to be a valuable source of information
for both professional biologist and laymen alike. I have noted a number of locations
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Bulletin of the Maryland Herpetological Society
Volume 39 Number 1
March 2003
which I either want to revisit, along with numerous natural communities which I look
forward to visiting. I also would recommend this book to anyone traveling through
the state with a general interest in nature as it provides a wealth of information and
will certainly make routing a specific trip much more enlightening, and productive.
Every Wisconsinite should have a copy in his or her home for reference, knowledge,
and entertainment.
Harlan D. Walley, Department of Biology, Northern Illinois University, Dekalb,
Illinois 60115. hdw@niu.edu .
Received 31 January 2003
Accepted 7 February 2003
Bulletin of the Maryland Herpetoiogical Society
page 11
Volume 39 Number 1
March 2003
Book Review: Herpetology in Montana, by Bryce A. Maxell, J. Kirwin
Werner, Paul Hendricks and Dennis L. Flath. Northwest Fauna Society for North¬
western Vertebrate Biology (5), 138 pp. Wrs. P.O. Box 22313, Seattle, Washington
98122. $12.00 + $1.50 shipping.
The present volume has been a long awaited compendium on the herpetofauna
of Montana, and was written nearly 200 years after the first European herpetological
observations were cited from this state. In the Introduction, the authors provide a
thorough summary of the historical information on Montana from the time of Lewis
and Clark’s arrival in 1805 to the present. This is followed by a checklist of native
species and subspecies, along with dichotomous keys for eggs, larvae, and adults of
amphibians along with a key for identification of juvenile and adults of reptiles
found within the state. A unique aspect which has not been seen in recent herpeto¬
logical literature is a chronological summary of articles on the herpetofauna of Mon¬
tana and Yellowstone National Park between 1800 and 2002, along with a figure
showing the number of articles cited in the bibliography with information on the 29
native species and number of voucher specimens observed.
The individual species accounts provide comments on habitat, along with
information on the Earliest Literature and Voucher Records, Maximum Elevation,
and Voucher Record Summary, followed by a Bibliographic Index which closely
follows the Catalogue of American Amphibian and Reptiles Species Accounts in
format. Each species has an excellent distributional map having been compiled from
information on 3390 amphibian voucher specimens and 1238 reptile voucher speci¬
mens amassed from different institutions through 2001, along with several more re¬
cent voucher specimens collected during the 2001 and 2002 seasons. The only draw¬
back is the lack of illustrations for any of the species cited within the text, although
this should not distract from the usefulness of this badly needed reference work.
Seven species: Amby stoma tigrinum diaboli, Dicamptodon aterrimus, Spea
intermontana, Bufo hemiophrys, Rana sylvatica, Phrynosoma douglasii and Eumeces
skiltonianus utahensis are questionable species having been found in bordering adja¬
cent states, and possibly will be recorded for Montana with further collecting. The
authors provide information on the status and closest distributional data related to
each of these questionable species, followed by accounts for exotic species or sub¬
species having been reported from Montana. The below listed species, Taricha granu¬
losa, Bufo canorus, Hyla arenicolor, Pseudacris clarkii, Rana catesbeiana, Rana
clamitans, Clemmys marmorata, Trachemys scripta elegans, Terrapene Carolina
triunguis , Terrapene ornata, Phrynosoma coronatum, Phrynosoma platyrhinos, and
Heterodon platirhinos are considered exotic species. The major portion of these
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Bulletin of the Maryland Herpetological Society
Volume 39 Number 1
March 2003
species have been cited from translocation populations, or misidentification, and have
not become established within the state. A bibliography of 543 citations dealing
with the herpetofauna within and immediately adjacent to Montana round out this
excellent publication. I would highly recommend this book to anyone having an
interest in North American herpetofauna, and especially those living within the state
of Montana, as it is without graphical errors, and extremely informative, and will
probably remain the standard text for the state for years to come.
Harlan D. Walley , Department of Biology, Northern Illinois University,
Dekalb, Illinois. 60115. hdw@niu.edu
Received 31 January 2003
Accepted 7 February 2003
Bulletin of the Maryland Herpetological Society
page 13
Volume 39 Number 1
March 2003
News and Notes
■
page 14
Volume 39 Number 1
March 2003
News and Notes
Reptile and
Amphibian Rescue
410-580-0250
We will take as many unwanted pet reptiles and
amphibians as space allows.
Leave a message with your name and number to
give up an animal for adoption;
or to volunteer to help with our efforts.
OUR CURRENT NEEDS:
• Commercial or Passenger Van
• UVB Lights • Power & Hand Tools • Bleach
• Equipment & Food • Paper Towels
www.reptileinfo.com
Bulletin of the Maryland Herpetological Society
page 15
Volume 39 Number 1
March 2003
News and Notes
News from
CORNELL MARITIME PRESS
TIDEWATER PUBLISHERS
P.o. Box 456, Centreville, Maryland 21617
410-758-1075 • 800-638-7641 • 410-758-6849 (fax)
Landmark Field Guide First for Delmarva’s Amphibians and Reptiles
Publication Supports Delaware Nature Society ’s Project on Herpetofauna
February 1 , 2003 — A husband-and-wife team from the Delaware Nature Society has compiled
the first-ever guide to amphibians and reptiles of Delmarva. Jim White and Amy Wendt White
spent nearly fifteen years combing the region’s swamps, woods, and fields.
The result of their work is Amphibians and Reptiles of Delmarva, which includes
seventy-three species with each account containing a detailed description of physical
characteristics, comparisons to similar species, and information on the geographical distribution,
abundance, habitat, reproduction and development, and behavior. From skinks to snakes, from
turtles to frogs, this compact but thorough reference gathers vital information about the Delmarva
Peninsula’s herpetofauna. Herpetology is the branch of zoology dealing with reptiles (turtles,
lizards, and snakes) and amphibians (salamanders and frogs).
In 1986, Jim White, associate director of land and biodiversity management for the
Delaware Nature Society, became a principal investigator in a herpetological survey of the state;
this research evolved into the field guide published recently by Tidewater Publishers in
association with the Delaware Nature Society. Amy White, who is a teacher-naturalist at the
Nature Society, often accompanied her husband on early expeditions. A concerted effort to
organize and write the guide commenced in 1998 as she became more involved with the entire
project.
“We hope the book instills a strong conservation ethic in the area’s residents, an ethic that
leads to better protection of these animals and the habitats in which they live,” the authors said.
The field guide has already garnered tremendous accolades, including praise from Roger
Conant, esteemed Director Emeritus of the Philadelphia Zoo and one of the country’s foremost
herpetologists. “As a book on the herpetology of a small geographic area, this publication is the
most complete and thorough of any I have seen,” said Dr. Conant, author of the Peterson Field
Guide to Reptiles and Amphibians of Eastern and Central North America. “Every species and
subspecies is described in meticulous detail, and the splendid photographs make identification
easy.”
Given its size, the Delmarva Peninsula — encompassing all of Delaware, the northeast
comer of Maryland and its Eastern Shore, and the eastern shore of Virginia — boasts a rich array
of amphibians and reptiles. However, Delmarva’s herpetofauna is often overlooked by the vast
majority of people living in the area. Many have never heard a northern spring peeper call or
seen a salamander, even though a breeding chorus of peepers is common in the spring, and
salamanders are residents of most woodlands in Delmarva.
Misconceptions also abound. For example, people sometimes think they have seen
“cottonmouths” (water moccasins) on Delmarva, even though this venomous species is not
continued
page 16
Bulletin of the Maryland Herpetological Society
Volume 39 Number 1
March 2003
News and Notes
White and White ' Amphibians and Reptiles of Delmarva
Page 2
known to occur north of the mouth of the Chesapeake Bay. Misidentification of species,
especially of snakes, is common, Amy White said. “Fear and misunderstanding lead some
people to mistake harmless snakes as venomous and they kill them needlessly.”
The field guide will be useful to a wide variety of people interested in these fascinating
creatures — from the curious child who brings home a salamander to students, experienced
naturalists, and professionals in need of life history, behavioral, and distributional information.
“This landmark field guide is an invaluable tool in our mission to educate children and
adults about the natural world,” said Michael E. Riska, executive director of the Delaware Nature
Society. “It will generate interest in Delmarva’ s fascinating amphibians and reptiles and
encourage conservation of all of our native animals and plants and their habitats.”
A skilled nature photographer, Jim White routinely captures many species and their
environs on film. More than ninety of his pictures are collected in the guide’s full-color insert,
which serves as a valuable tool for identifying various amphibians and reptiles.
“Dealing with tough environmental conditions often made the photography challenging,”
White said, referring specifically to photographing frogs. White often stood in water up to his
knees, usually at night, sometimes in the rain, and crept as close as he could to the frogs without
scaring them away. “Then you have to wait for the frog to call and try to get a good photograph
with its throat expanded. It takes a lot of effort and luck.”
Jim’s favorite species included in the guide is the bright green Barking Treefrog — which
graces the book’s cover — because he was the first to discover a breeding population of these
frogs on Delmarva. Amy’s favorite is the warty brown Bufo americanus americanus, or more
commonly the American Toad, because it serves as a harbinger of spring and is easier to catch
than most other frogs. But handlers beware: it secretes a powerful toxin that can be quite
unpleasant if it is ingested or comes in contact with the eyes.
Both Jim and Amy White earned degrees from the University of Delaware: Jim in
entomology and applied ecology and Amy in environmental engineering. They are members of
the Delaware Ornithological Society and Delaware Natural History Society. Jim is also a
member of Delaware Partners in Flight and Partners for Amphibian and Reptile Conservation.
Founded in 1 964, the Delaware Nature Society (www.delawarenaturesociety.org) fosters
understanding, appreciation, and enjoyment of the natural world through education; preserves
ecologically significant areas; and advocates stewardship and conservation of natural resources.
The Nature Society maintains two nature centers and manages four nature preserves for
biodiversity, research, and educational programs. The Nature Society is also the Delaware
affiliate of the National Wildlife Federation with input on national issues that concern citizens in
the region.
Amphibians and Reptiles of Delmarva is available for $14.95 from booksellers or
Tidewater Publishers, P.O. Box 456, Centreville, MD 21617 (800-638-7641).
Amphibians and Reptiles of Delmarva
296 pp. 32-page section with full-color photographs.
Figures. Maps. Table. References. Glossary. Index. Checklist.
4Y2 x 7. Paper. ISBN 0-87033-543-X. $14.95.
To receive a review copy of this book, qualified members of the media may contact Gregg Wilhelm, Marketing
Coordinator, at 800-638-7641 . To obtain an electronic file of this release, please send a message to
comell@crosslink.net.
Bulletin of the Maryland Herpetological Society
page 17
Volume 39 Number 1
News and Notes
page 18
Bulletin of the Maryland Herpetological Society
Society Publication
Back issues of the Bulletin of the Maryland Herpetological Society, where
available, may be obtained by writing the Executive Editor. A list of available
issues will be sent upon request. Individual numbers in stock are $5.00 each,
unless otherwise noted.
The Society also publishes a Newsletter on a somewhat irregular basis.
These are distributed to the membership free of charge. Also published are
Maryland Herpetofauna Leaflets and these are available at $. 25/page.
Information for Authors
All correspondence should be addressed to the Executive Editor. Manu¬
scripts being submitted for publication should be typewritten (double spaced)
on good quality 8 1/2 by 11 inch paper with adequate margins. Submit origi¬
nal and first carbon, retaining the second carbon. If entered on a word proces¬
sor, also submit diskette and note word processor and operating system used.
Indicate where illustrations or photographs are to appear in text. Cite all lit¬
erature used at end in alphabetical order by author.
Major papers are those over five pages (double spaced, elite type) and
must include an abstract. The authors name should be centered under the title,
and the address is to follow the Literature Cited. Minor papers are those pa¬
pers with fewer than five pages. Author’s name is to be placed at end of paper
(see recent issue). For additional information see Style Manual for Biological
Journals (1964), American Institute of Biological Sciences, 3900 Wisconsin
Avenue, N.W., Washington, D.C. 20016.
Reprints are available at $.07 a page and should be ordered when manu¬
scripts are submitted or when proofs are returned. Minimum order is 100
reprints. Either edited manuscript or proof will be returned to author for ap¬
proval or correction. The author will be responsible for all corrections to proof,
and must return proof preferably within seven days.
The Maryland Herpetological Society
Department of Herpetology
Natural History Society of Maryland, Inc.
2643 North Charles Street
Baltimore, Maryland 21218
Bulletin of the Maryland Herpetological Society
page 19
Maryland
Herpeiological
.Society
US ISSN: 0025-4231
\o *
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BULLETIN OF THE
Tftarylanb
f)erpeto(ogtcal
Oociety
DEPARTMENT OF HERPETOLOGY
THE NATURAL HISTORY SOCIETY OF MARYLAND, INC.
MDHS . A Founder Member of the Eastern
Seaboard Herpetological League
30 JUNE 2003
VOLUME 39 NUMBER 2
BULLETIN OF THE MARYLAND HERPETOLOGICAL SOCIETY
Volume 39 Number 2 June 2003
CONTENTS
Leptotyphlopidae: Worm Snakes, not Blind Snakes!
Van Wallach . . . . . 21
Presence of the Rio Fuerte Beaded Lizard (Heloderma horridum exasperatum) in
western Chihuahua
Julio A. Lemos-Espinal, David Chiszar and
Hobart M. Smith . . . . . . . . . . . . 46
Reproductive Biology and Population Structure of Eurycea longicauda longicauda
W. T. McDowell and B. A. Shepherd . . . . . . 52
New Distributional Record for the Southern Leopard Frog in Frederick County,
Maryland
Wayne G Hildebrand . . . . . . . 62
Book Review
64
BULLETIN OF THE
mbt)6
Volume 39 Number 2 June 2003
The Maryland Herpetological Society
Department of Herpetology, Natural History Society of Maryland, Inc.
President Tim Hoen
Executive Editor Herbert S. Harris, Jr.
Steering Committee
Frank B. Groves Jerry D. Hardy, Jr.
Herbert S. Harris, Jr. Tim Hoen
Library of Congress Catalog Card Number: 76-93458
Membership Rates
Membership in the Maryland Herpetological Society is $25.00 per year
and includes the Bulletin of the Maryland Herpetological Society. For¬
eign is $35.00 per year. Make all checks payable to the Natural History
Society of Maryland, Inc.
Meetings
Meetings are held monthly and will be announced in the “Herp Talk"
newsletter and on the website, www.naturalhistory.org.
Volume 39 Number 2
June 2003
Leptotyphlopidae: Worm Snakes, not Blind Snakes!
Van Wallach
While it is true that the only scientifically relevant names of species are the
Latinized generic, specific and subspecific epithets, common names are used (some¬
times exclusively) in the secondary and lay literature. Also, numerous papers, mono¬
graphs and even books have been devoted to listing the common names of reptiles
and amphibians. I here make a plea for the correction and standardization of the
common names of the scolecophidian blind and worm snakes of the world
(Typhlopidae and Leptotyphlopidae).
There is a perplexing confusion with the vernacular nomenclature of these
two snake families that dates back many years and I wish to set the record straight for
future publications. Table 1, which lists the common names applied to the three
scolecophidian families in various publications, illustrates the confusion and lack of
standardization in vernacular nomenclature for these snakes. Common names ap¬
plied to the Leptotyphlopidae include “Worm Snakes,” “Blind Snakes,” “Slender
Blind Snakes,” “Thread Snakes,” “Earth Snakes,” “Earthworm Snakes,” “Burrow¬
ing Snakes,” “Blind Worms,” “Bigjawed Blindsnakes,” “Ant-nest Worms,” and
“Ground Puppys.” While this article addresses the typhlopid and leptotyphlopid situ¬
ation, a few comments on the anomalepidids are also included. The Typhlopidae (and
the closely related Anomalepididae) are “Blind Snakes” and the Leptotyphlopidae
are “Worm Snakes” and they should be referred to as such! This is manifestly clear
upon a comparative examination of any representatives of the two families. The dif¬
ferences are clearly significant, which is why taxonomists have placed them in sepa¬
rate families since 1891. Three superficial points (body size, coloration, and eye size)
should suffice to demonstrate that leptotyphlopids are more worm-like and less blind
than typhlopids.
Firstly, leptotyphlopids are smaller in average and maximum size than are
typhlopids: shorter in total length, narrower in body diameter, and thinner in body
proportion. The maximum length in Leptotyphlops is 400 mm (L. macrolepis) with
only eight species (L. anthracinus , L. humilis , L. maximus , L. melanotermus , L.
occidentalism L. septemstriatus , L. tricolor , L. weyrauchi) reaching a maximum length
of 300 mm or longer. Average length in leptotyphlopids is about 150 mm with a
midbody diameter of 3 mm. On the other hand, 15 typhlopids reach more than 600
mm in length (. Ramphotyphlops nigrescens , R. proximus , R. unguirostris , R. waitii ,
Rhinotyphlops acutus , R. mucruso , R. schlegelii , R. somalicus , T. angolensis, T.
congestus, T. decorsei , T. lineolatus , T. punctatus , T. schmidti , T usamharicus) with
Bulletin of the Maryland Herpetological Society
page 21
Volume 39 Number 2
June 2003
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Bulletin of the Maryland Herpetological Society
Table 1, Continued
Source Leptotyphlopidae Typhlopidae Anomalepididae
Behler & King, 1979 Slender Blind Snakes
Stucki-Stim, 1979 Worm Snakes Glass or Blind Snakes
Englemann & Obst, 1 98 1 Slender Blind Snakes Worm or True Blind Snakes
Volume 39 Number 2
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Adler & Halliday, 1986 Thread Snakes Typical Blind Snakes Dawn Blind Snakes
De Lisle et al„ 1986 Blind Snakes
Mattison, 1986 Blind Snakes Worm or Blind Snakes
Patterson, 1986, 1987 Worm or Thread Snakes Blind Snakes
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Behler, 1999 Slender Blind Snakes
Bartlett & Tennant, 1999 Slender Blind Snakes
Chan-ard et aL, 1999 Blind or Worm Snakes
Komacker, 1999 Blind Snakes Worm Snakes Blind or Worm Snakes
Tennant & Bartlett, 1999 Slender Blind Snakes Blind Snakes
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page 28
Bulletin of the Maryland Herpetological Society
Volume 39 Number 2
June 2003
a maximum length of 950 mm in Rhinotyphlops mucruso. The average length of
typhlopids is about 300 mm with a midbody diameter of 10 mm. The thinnest of all
scolecophidians are 11 African Leptotyphlops (with length/width ratios of 100-140),
which is why they are often called “Thread Snakes,” while the stoutest of all are
some African and South American Typhlops (with length/width ratios of less than
20).
Secondly, leptotyphlopids resemble worms in coloration (usually pink or
beige) more than typhlopids do (which are generally brown to black). Many
leptotyphlopids lack pigmentation and appear pink in coloration due to subcutaneous
capillary beds. Most typhlopids are heavily pigmented in shades of gold, brown or
black, often with a pattern of contrasting colors forming stripes, spots, or blotches.
Superficially, Leptotyphlops resembles earthworms in all ways except the presence
of scales.
The following examples illustrate why leptotyphlopids should be called
“Worm Snakes.” Schmidt and Davis (1941), although naming the Leptotyphlopidae
“Blind Snakes,” remarked that they “are strikingly similar to earthworms.” In their
defence it must be mentioned that members of the genus Carphophis are also termed
“Worm Snakes.” This is a prime example of the inadequacy of common names ! Oliver
(1955), Heymann (1975), and Grater (1981) called Leptotyphlops “Blind Snakes”
but mentioned that they were often mistaken for earthworms. Stoops and Wright
(1993) classified Leptotyphlops as “Blind Snakes” but went on to explain that they
are also “called worm snakes because that is exactly how they appear— small, slen¬
der, and resembling worms.” Degenhardt et al. (1996) labelled the Leptotyphlopidae
as “Blind Snakes” but mentioned that they are often called “Worm snakes.” Even
more ambiguous are Conant and Bridges (1939) who referred to the Leptotyphlopidae
as “Blind Snakes” but listed Leptotyphlops dulcis as the “Texas Worm Snake.”
Thirdly, leptotyphlopids have much larger and better developed eyes than
any typhlopids, which are blind to a greater degree than Leptotyphlops based upon
relative eye size and presence of a distinct pupil. All Leptotyphlops have a moderate
to large scolecophidian eye with a distinct pupil; all typhlopids have either a small
eye with a pupil, a solid black eyespot (faint in some species), or no visible eye at all.
A few species of Ramphotyphlops and Typhlops have eyespots that are very faint or
only visible in juveniles but the only truly blind typhlopids are members of the genus
Rhinotyphlops. Whereas the eyes are located beneath a large head shield in all
scolecophidians, several groups of Leptotyphlops (such as the Neotropical albifrons
species group and the African L. macrops) have the eye bulging out from the contour
of the head. The relative eye diameter in comparison with head depth averages from
Bulletin of the Maryland Herpetological Society
page 29
Volume 39 Number 2
June 2003
0.25-0.33 in Leptotyphlops and less than 0.10 in Typhlops.
Leptotyphlopids definitely resemble worms much more than typhlopids do,
and typhlopids are more blind than leptotyphlopids. Thus the most accurate and de¬
scriptive common name for the Leptotyphlopidae is “Worm Snake” and that of the
Typhlopidae is “Blind Snake.” “Thread Snake” is not as accurate a name as “Worm
Snake” because some species of Leptotyphlops are thick-bodied with length/width
ratios of 30 or less.
Klauber (1940) revised the genus Leptotyphlops and he was the first to cor¬
rectly use the appellation “Worm Snakes.” Modem authors continue to refer to North
American Leptotyphlops as “Blind Snakes” (Tennant, 1984; Williamson et al., 1994;
Rossi and Rossi, 1995; Werler and Dixon, 2000; Grismer, 2002). Even more trou¬
bling is that the official vernacular indices of the USA have continued to refer to
Leptotyphlops as “Blind Snakes” (Collins et al., 1978; Collins et al., 1982; Collins,
1990, 1997; Collins and Taggart, 2002) or “Threadsnakes” (Crother, 2000). The us¬
age of “Blind Snakes” for Leptotyphlops probably stems from Conant et al. (1956),
who termed them “Slender Blind Snakes.” This inaccurate terminology has been
perpetuated by the entire Peterson field guide series (Conant, 1958, 1975, Stebbins,
1966, 1985, and Conant and Collins, 1991), the Gulf field guide series (Tennant,
1985; Brown, 1997; Bartlett and Tennant, 1999; Tennant and Bartlett, 1999), the
Audubon Society field guide series (Behler and King, 1979; Behler, 1999), and oth¬
ers (Hahn, 1979).
The most ambiguous case in nomenclature is that by Liner (1994) and Frank
and Ramus (1995) wherein they referred to the Leptotyphlopidae as “Slender Blind
Snakes” and the Typhlopidae as “Blind Worm Snakes,” although Skinner (1973) la¬
belled Leptotyphlops as “Worm-snakes, Thread-snakes, Earth-snakes, and Blind-
worms” in the same breath! Tennant (1998) has even transferred the leptotyphlopid
“Blind Snakes” to the Typhlopidae! Some authors (Conant and Collins, 1998; Tennant
and Bartlett, 1999) lumped the two families together and called both the Typhlopidae
and Leptotyphlopidae “Blind Snakes” but Emsley (1977) labelled both Typhlops and
Leptotyphlops as “Worm Snakes.” Greene (1997) labelled both families as
“Blindsnakes” but also referred to African Leptotyphlops as “Threadsnakes.” Although
Adler and Halliday (1986) and Stidworthy (1989) termed the Leptotyphlopidae as
“Thread Snakes,” they referred to Leptotyphlops dulcis as a “Blind Snake.” Brazaitis
and Watanabe (1992, 1993) used “Slender Thread Snakes” for African Leptotyphlops
and “Blind Snakes” for the Typhlopidae, but then termed Ramphotyphlops braminus
a “Worm Snake.” Chan-ard et al. (1999) referred to Typhlops as “Blind Snakes” but
Ramphotyphlops as “Worm Snakes.” Liner (1994) called Typhlops “Common Blind
page 30
Bulletin of the Maryland Herpetological Society
Volume 39 Number 2
June 2003
Worm Snakes” but listed Ramphotyphlops braminus , whose universally known com¬
mon names are either the “Flowerpot or Brahminy Blind Snake,” as the “Longtail
Blind Snake.” Khan (2002) called the Leptotyphlopidae and Typhlopidae “Worm
Snakes” and then referred to the Leptotyphlopidae as “Thread Snakes” and the
Typhlopidae as “Blind Snakes.”
It is obvious, as every systematist knows, that there is no standardization for
vernacular names, and this is especially evident in the Scolecophidia. I appeal for a
more consistent and accurate terminology for the common names of leptotyphlopids
and typhlopids. In an effort to be as descriptive as possible, all accounts and check¬
lists should refer to the Leptotyphlopidae as “Worm Snakes” and the Typhlopidae as
“Blind Snakes.” The question of how to deal with the nomenclature of the two spe¬
cies of alethinophidian “Worm Snakes” ( Carphophis amoenus and C. vermis ) re¬
mains unanswered. The resemblance of Carphophis to earthworms is limited to their
size and natural habitat. I suggest an alternative common name be applied to this
genus.
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page 46
Bulletin of the Maryland Herpetological Society
Volume 39 Number 2
June 2003
Presence of the Rfo Fuerte Beaded Lizard
(Heloderma horridum exasperatum) in
Western Chihuahua
Julio A. Lemos-Espinal, David Chiszar and Hobart M. Smith
Heloderma horridum exasperatum was described by Bogert and Martin del
campo (1956) on the basis of fifteen specimens and one incomplete skin, all from
three localities in the Rio Fuerte drainage basin of southern Sonora and northern
Sinaloa. Additional records for Sonora and Sinaloa are reported by Hale (1989), Hardy
and McDiarmid (1969) and Ottley (1981a and 1981b). Bogert and Martin del Campo
(1956) mentioned that this subspecies of beaded lizard should range eastward into
the barrancas of western Chihuahua. Recent field work in the regions known as Canon
de Chmipas and Barranca de Batopilas has revealed the presence of beaded lizards in
several localities of western Chihuahua (Figure 1).
Fig. 1. Localities of record (round white dots with a black center) for H. h.
exasperatum . Note the correlation with fotthill contours.
Bulletin of the Maryland Herpetological Society
page 47
Volume 39 Number 2
June 2003
In October 2002 JLE surveyed the herpetology of the vicinity of the town of
Chinipas, Chihuahua. Local residents kindly offered to help us with the survey, and
told us about the presence of beaded lizards, locally called “escorpiones,” in the
mountains that surround the town. All these natives considered the “escorpiones” as
extremely poisonous lizards which should be killed to prevent being bitten by them.
Natives have the belief that the skin of “escorpiones” is useful as a cure for snake¬
bites. When a beaded lizard is killed, the skin is removed and stored to use as medi¬
cine. In addition to efforts to find specimens of H. h. exasperation in the field, Helo-
derma skins were sought in all the villages around the town of Chinipas (Figure 2).
Although we failed to find specimens in the field, we were able to establish the pres¬
ence of H. h. exasperatum through nine skins collected in several localities (Table 1).
Fig. 2. One of the skins from the vicinity of the Chinipas area. The pattern charac¬
teristic diagnostic of H. h. exasperatum is the approximate equality of light and
pigmented areas, as shown here.
page 48
Bulletin of the Maryland Herpetological Society
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June 2003
Table 1. Summary of localities where H. h. exasperatum has been recorded.
Number
Locality
Coordinates
Elevation
Source
Chihuahua:
Meters
UBIPRO 10667
A gua Caliente
27°27'28.0"N
108o3r40.9”W
572
This note
UBIPRO10616
Agua Salada
27°22'54.1"N
108o28'8.6"W
536
This note
UBIPRO 10609
(Chmipas)
Casa Colorada
27°24'13.0"N
108°3r56.2"W
528
This note
UBIPRO 10661
El Limon
27°24,L0"N
108°32'36.0"W
451
This note
UBIPRO 10663-6
La Cienega
27°27'27.5"N
108o34'50.9"W
600
This note
UBIPRO 10662
Machahuivo
27°25T.1"N
108°33'43.6"W
547
This note
UBIPRO 10660
Mesa El Rosario
27°22'48.0"N
108°32,41.1"W
469
This note
UBIPRO 10668
Sinaloa:
Palmarejo
27°23'5L1"N
108°24'32.7"W
615
This note
USNM38116
San Bias
26°4'35.7"N
m°45'45"W
100
Bogert and
Martin del
Campo (1956)
Not Given
13 kms NNE
26°54'3.2"N
400
Hardy and
Sonora:
Baca
108°23’38 .2"W
McDiarmid
(1969)
AMNH64220-4
Alamos
27°1'21.1"N
108°56'2L8"W
450
Bogert and
Martin del
Campo (1956)
Not Given
Between
Toni chi and
NuiLHwy
Mex. 16.
28°24'35.7"N
m°iy2$.rw
Not
given
Hale (1989)
AMNH63698-0
MVZ 50863
63703
Guirocoba
26°54'H.3"N
108°41'33.0,,W
450
Bogert and
Martin del
Campo (1956)
Not Given
Movas
28°9'43.8MN
109°26'21.8,'W
Not Given
Ottley (1981a)
Not Given
16.8 kms
NNW Alamos
27°5'9.7,!N
109°5'3.6"W
Not Given
Ottley (1981b)
Bulletin of the Maryland Herpetological Society page 49
Volume 39 Number 2
June 2003
These records constitute the first evidence of the presence of this subspecies of
beaded lizard in Chihuahua and extends its range ~58 km northeastward of the town
of Alamos, Sonora.
In addition to these records, JLE has been working in the Barranca de Batopilas
for several years. Local people in this region are quite familiar with “escorpiones,”
which are nevertheless considered rare and can be seen only once in a while. Searches
for H. h. exasperatum in this region unfortunately have failed to find specimens in
the field, or to find any other evidence (e.g. skins) of the presence of this subspecies.
However, we are positive that beaded lizards occur there.
Specimens from Chinipas, Chihuahua, are deposited in the herpetological col¬
lection of LJnidad de Biologia, Tecnologia y Prototipos - UN AM (UBIPRO).
Acknowledgments.
Support for this study was provided by the Comision Nacional Para el
Conocimiento y Uso de la Biodiversidad (CONABIO) under projects 003, X004 and
AE003.
Literature Cited
Bogert, C. M. and R. Martin del Campo.
1956. The gila monster and its allies. The relationships, habits, and be¬
havior of the lizards of the family Helodermatidae. Bull. Am. Mus.
Nat. Hist. 109: 1-238.
Hale, S. F.
1989. Alamos field trip 1989: trip notes, part III. Hermosillo to Alamos.
Tucson Herp. Soc. Newsl. 2: 91-93.
Hardy, L. M., and R. W. McDiarmid.
1969. The amphibians and reptiles of Sinaloa, Mexico. Univ. Kansas
Publ. Mus. Nat. Hist. 18(3): 39-262.
Ottley, J. R.
1981a. Geographic distribution: Heloderma horridum exasperatum . Herp.
Rev. 12: 64.
Ottley, J. R.
1981b. Geographic distribution: Heloderma suspectum suspectum. Herp.
Rev. 12: 65.
page 50
Bulletin of the Maryland Herpetological Society
Volume 39 Number 2
June 2003
JLE: Laboratorio de Bcologia UBIPRO-UNAM , Av : de los Barrious s/n , los Reyes
Iztacala , Tlainepantla , Edo. de Mexico 54090 Mexico.
DC: Department of Psychology and Museum, University of Colorado, Boulder,
CO 80309-0345.
HMS : Department ofEPO Biology, University of Colorado, Boulder,
CO 80309-0334.
Received 10 March 2003
Accepted 7 April 2003
Bulletin of the Maryland Herpetological Society
page 51
Volume 39 Number 2
June 2003
Reproductive Biology and Population Structure of
Eurycea longicauda longicauda
W. T. McDowell and B. A. Shepherd
Abstract
The reproductive biology and population structure of the salamander Eurycea
longicauda longicauda (Family Plethodontidae) was studied at three sites in south¬
ern Illinois to determine possible microgeographic variation. Males matured at > 46
mm SVL while females matured at > 49 mm snout-vent length (SVL) with both at
two years of age. Surface activity was from late February through late November.
Mating occurred in October and November with oviposition from November through
early January. Adults were much more abundant in the summer months and than
juveniles.
Introduction
Salamander ovarian and testicular cycles have typically been described by
monthly “vertical” sampling (Tilley, 1977) with gonadal examinations to determine
seasonal changes in reproductive tracts. Interpretations of the reproductive biology
(breeding periods, oviposition periods, etc.) of the longtailed salamander Eurycea
longicauda longicauda (Green) have been made with limited monthly samples/year
and or few animals (Mohr, 1943; Hutchinson, 1956; Franz, 1964; Anderson and Martino,
1966; Guttman, 1989; Petranka, 1998). Ireland (1974) described the spermatogenic
and ovarian cycles and Trauth et al. (1993) described the caudal hedonic glands of E.
1. melanopleura. In southern Illinois, Williams et al. (1984, 1985) described the sper¬
matogenic cycle and seasonal changes in cloacal glands of E. I longicauda and found
reproduction to be late October while Phillips et al. (1999) stated that oviposition was
in underground crevices in late summer and autumn. The ovarian cycle and popula¬
tion structure of E. 1. longicauda have yet to be fully described. Our purposes were to
investigate the reproductive biology (especially the ovarian cycle) and the popula¬
tion structure of E. 1. longicauda in southern Illinois.
Material and Methods
Metamorphosed salamanders were sampled monthly at three sites in southern
Illinois; a woodland stream leading from a rock shelter 11 km N of Glendale, Pope
Key Words: Eurycea. reproductive biology, ovarian cycle, oviposition period, popu¬
lation structure, southern Illinois
page 52
Bulletin of the Maryland Herpetological Society
Volume 39 Number 2
June 2003
Co. (Pope Co. A), another woodland stream without a rockface in Pope Co. 3 km E of
Rosebud (Pope Co. B) and a cave 10 km E of Cobden, Union Co. We collected a
monthly sample of at least N = 20 from all three sites combined. At Pope Co. A 124
salamanders were collected from May 1976 through April 1977 and at Pope Co. B 32
were found in March and April 1982. At Union Co. cave 29 were found during May,
July, and August 1976, and principally August ( N = 22) at an adjoining woodland
stream. A small series of larvae was collected during March 1976 at the cave and
identified as E. /. longicauda. Salamanders were hand collected from beneath rocks
within 1-2 m of the streamsides and on the cave wall surfaces.
Salamanders (N-- 1 85) were euthanized by emersion in 10% chloretone within
48 hr of capture, fixed and preserved in 10% formalin. Vasa deferentia, testes and
oviductal widths of mature adults were measured to the nearest 0.25 mm with a Nikon
dissecting stereomicroscope equipped with an ocular micrometer to determine sea¬
sonal size differences. Gonads of individuals > 35 mm SVL (which is the distance
from the tip of the snout to the posterior margin of the vent) were examined to deter¬
mine sexual maturity and reproductive condition in males by degree of testicular
pigmentation and vasa deferentia shape and degree of pigmentation. Previous studies
have shown that they mature at slightly less than 50 mm SVL (Anderson and Martino,
1966). Female sexual maturity and reproductive condition were determined by ova¬
rian follicle diameter and yolkiness and oviductal shape and width (Anderson and
Martino, 1966). A random subset ( N = 25 or more) of ovarian follicles from each
mature female was measured to the nearest 0.5 mm. The vitellogenic cycle at Pope
Co. A and Union Co. cave was determined from adult female monthly mean follicle
diameter described by a regression equation with significance at P < 0.01. An F test
was used to determine positive correlation between ova development and month of
collection. The population structure at Pope Co. A and Union Co. cave was deter¬
mined by analysis of monthly frequency histograms of each individual snout-vent
length. Voucher specimens are deposited at the Southern Illinois University at
Carbondale Fluid Vertebrate Collection.
Results
Juvenile males (< 45 mm SVL) had unconvoluted, unpigmented or lightly
pigmented vasa deferentia and unpigmented (white) or partially (< 75%) lightly pig¬
mented testes. Mature males (N = 65) with convoluted heavily pigmented vasa defer¬
entia and heavily pigmented testes were > 46 mm SVL, although one individual from
Pope Co. B matured at 44 mm SVL. All mature males from March through May had
thin vasa deferentia (approximately 0.25 mm wide) and testes (approximately 1.25
mm wide). Testes began increasing in width in July, were widest in August and Sep-
Bulletin of the Maryland Herpetological Society
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Volume 39 Number 2
June 2003
tember (2.25 mm) and by early October were 1.25 mm and thin (evacuated). By early
October males had very thick vasa deferentia (approximately 1.0 mm wide) packed
with sperm, which indicated that mating probably occurred in October or November.
Mature males from any given month had testes with unimodal width distributions
indicating an annual breeding cycle. Juvenile females (< 49 mm SVL) had small
unyolked (previtellogenic) follicles and thin straight oviducts. Females (TV =47) from
all three sites with large yellow yolky ovarian follicles or thick, convoluted oviducts
were considered mature. Only females > 49 mm SVL from Pope Co. A and Union
Co. cave from all months sampled (Fig. 1) showed vitellogenesis, hence, female
sexual maturity at these two sites was achieved by this size. All adult females from
ao oo •• o
Q 0 0 O
• ••
L , , , , , f r-T- « 1 1 « ■' ' > r“"» « ■ ' ^
45 SO 56 SO 65
SNOUT-VENT LENGTH (mm)
FIG. 1 . Female follicle diameter from Pope Co. A (solid circles) and Union
Co. cave stream (empty circles) from May 1976 through April 1977 as a function of
snout-vent length. Each circle represents mean follicle size of one female.
page 54
Bulletin of the Maryland Herpetological Society
Volume 39 Number 2
June 2003
February through June had small ova (0.5 mm), thin but convoluted oviducts and
spent ovaries. Vitellogenesis (Fig. 2) began during July and before mating when mean
follicle diameter (1.2 mm) more than doubled in size from June (0.5 mm). Mean
follicle diameter increased to 2.4 mm by September and October with oviductal widths
thickest in October. Monthly follicle diameters from females at Union Co. cave (Fig.
2) agree well with those from Pope Co A indicating no microgeographic variation.
Monthly changes in ovum size were significantly different (F = 29.3, P < 0.01) with
follicle diameters in late spring small and increasing diameters in summer and fall.
Month of collection accounted for almost 90% of the variation in individual female
mean follicle size. Follicle diameter size variance was greatest in September and
FIG. 2. Vitellogenesis of females from Pope Co. A (solid circles) and Union
Co. cave stream (empty circles) from May 1976 through April 1977. Each circle
represents mean follicle size of one female. The line fits the regression equation Y =
-0.05 + .043X and r2 = 0.88.
Bulletin of the Maryland Herpetological Society
page 55
Volume 39 Number 2
June 2003
October as mean diameters increased. Monthly mean ovarian follicle diameters at
Pope Co. A were unimodal in distribution (only one size present in any monthly
sample) indicating an annual female breeding cycle. Because all adult females from
Pope Co. A in late October were gravid, and all those in late February and March
were spent, we suggest that oviposition occurred from November through possibly
early January at this site and base the mean oviposition date as December first. Fif¬
teen larval hatchlings (x = 10.5 mm SVL) from the Union Co. cave were collected on
13 March 1976.
The population structure at Pope Co. A and Union Co. cave (Fig. 3) had distinc¬
tive size groups that are interpreted as age classes. First year juveniles were found
from July through November. Four small juveniles (24-29 mm SVL) from July were
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FIG. 3. Population structure at Pope Co. A and Union Co. cave stream. Open
squares depict non-dissected juveniles, gray squares are females (including both adults
and immatures) and dark squares are males (including both adults and immatures).
page 56
Bulletin of the Maryland Herpetological Society
Volume 39 Number 2
June 2003
presumably recent metamorphs as larval metamorphosis occurred during July 1976 at
Pope Co. A (McDowell, 1989). Slightly larger juveniles (to 32 mm SVL) were found
in the following months. Juveniles 28-32 mm SVL from October 1976 would be 10
months old based on the estimated December first mean ovipositional date and are
about the same size as those from March 1977, but this is probably the result of no or
little growth during winter. Juveniles 26-33 mm SVL from March 1977 were a spring
1976 cohort which had overwintered and would be 11 months old. Juveniles 32-36
mm SVL from May would be 16 months old while October individuals 42-45 mm
SVL would be 20 months old. They would mature in the following winter or spring.
Juveniles had a size range of 24-46 mm SVL. During July and August few second
year juveniles were found which was due to samples skewed for adults and first year
juveniles. Based on age determinations correlated with gonadal examination data,
males and females would mature in the spring of their second year and reproduce for
the first time in the following fall or early winter. Adults (N - 94; sex ratio 1.4 : 1.0)
were much more common in the summer months (especially August) and more com¬
mon than juveniles (1.6 : 1.0). Females grew to a larger size (to 60 mm SVL) than
males exhibiting SSD (sexual size dimorphism). Few salamanders were collected from
November through February but they were common in early spring, summer, and fall.
At Pope Co. B adult females (N- 10) from March and April had follicle diam¬
eters with bimodal distributions. Six mature females with ova 0.5 mm were spent and
had thin oviducts while four other mature females had much larger ova (2.0-2.5 mm)
and ranged from 47-54 mm SVL. These larger ova were atretic, being lightly brown¬
ish, collapsed, very flaccid and were carried over winter. The smallest mature female
(47 mm SVL) had presumably undergone vitellogenesis for the first time. All mature
males (N = 14) had thin vasa deferentia and testes. Eight juveniles and no metamorphs
were collected
Discussion
Several new interesting findings were derived from our study. The surface
population at Pope Co. A was more active throughout the year than those found in
previous studies (Anderson and Martino, 1966; Guttman, 1989; Petranka, 1998). Sala¬
manders from Pope Co. A were found from late February through late November.
Anderson and Martino (1966) found that the earliest spring emergence at temporary
ponds in New Jersey was late April or early May with animals in underground re¬
treats by the end of October. Guttman (1989) found surface activity to end during
August in Ohio. The longer seasonal activity present in our study populations may be
due to the milder winters in southern Illinois. All salamanders at both stream locali¬
ties were collected streamside and not away from water. Attempts were made to col-
Bulletin of the Maryland Herpetological Society
page 57
Volume 39 Number 2
June 2003
lect salamanders in nearby woodland areas but failed, contrary to the study by Ander¬
son and Martino (1966) who found them away from water sites. Sizes at maturity also
differed from those found in previous studies. Males matured at > 46 mm SVL (al¬
though one male from Pope Co. B matured at 44 mm SVL) and females at > 49 mm
SVL (although one female from Pope Co. B matured at 47 mm SVL). Anderson and
Martino (1966) found males to mature at 43-45 mm SVL and females at 47 mm SVL
and both at three years. We found our adults to mature at two years. Metamorph sizes
from July correlate well with those of Anderson and Martino’s (1966), however they
indicated that juveniles were much more abundant than in our study.
Other studies have also indicated a fall or winter reproductive season. Bishop
(1943) in New York described courtship in the laboratory on 18 November while
Cooper (1960) observed courtship in Maryland in the field on 1 8 October. Mohr (1943)
found deposited eggs in Pennsylvania in early developmental stages on 2 January
while Franz (1964) found deposited eggs in Maryland on 23 November. Anderson
and Martino (1966) reported oviposition in January. Oviposition of our salamander
populations in southern Illinois correlate well with the preceding observations but
seemed to occur later in the year (November through early January) than reported by
Phillips et al. (1999) who believed that oviposition occurred in late summer and fall.
The brief description by Guttman (1989) agrees with our determined ovarian cycle.
Ovarian follicle diameters of females at Pope Co. A and Union Co. cave are similar to
those of E. /. melanopleura from Arkansas (Ireland, 1974), but E. /. melanopleura
oviposits from December through March. Future work including using histological
techniques to determine if sperm storage occurs in the female spermathecae, should
determine how long after insemination females deposit their ova and whether there is
variation between time of insemination and subsequent oviposition.
Larval periods at Pope Co. A and B (McDowell, 1989) indicated a spring hatch¬
ing of March and March/April respectively having unimodal distributions of monthly
larval sizes. Differing year larvae were not present. Anderson and Martino (1966)
also found larval hatchlings in March. Newly hatched larvae were not present at any
other times at Pope Co. A, which correlates well with the estimated oviposition pe¬
riod of November through early January. However, larval hatchlings (those with yolk
remnants) have also been found 15 February through 12 June at other sites in south¬
ern Illinois (McDowell, 1988; 1992; unpublished), which may indicate a spring ovi-
positional period. Tilley (1977) found adult female Desmognathus ochropheus with
differing states of vitellogenesis (spent, with small ova, and with large yolky ova all
within a monthly sample) and postulated an extended oviposition season. Eurycea L
longicauda may have a more lengthy and variable oviposition period than previously
believed which warrants investigation of additional populations.
page 58
Bulletin of the Maryland Herpetological Society
Volume 39 Number 2
June 2003
Acknowledgments
For assistance in the field we thank T. Boatright, L. Weaver, and especially H.
Ohtsu. Larval identifications were confirmed by R. Brandon. For critically review¬
ing the manuscript we thank J. Martan, K. Lips, C. Phillips, J. Wheeler, and G.
Labanick. We also thank the landowners for allowing us to legally collect salamanders
on their private property.
Literature Cited
Anderson, J. D. and P. J. Martino.
1966. The life history of Eurycea /. longicauda associated with ponds.
Am. Midi. Natur. 75:257-279.
Bishop, S. C.
1943.
A Handbook of Salamanders. Comstock Publishing, Ithaca, New
York.
Cooper, J. E.
1960.
The mating antic of the long-tailed salamander. Maryland Natur.
30:17-18.
Franz, R.
1964.
The eggs of the longtailed salamander from a Maryland cave.
Herpetologica 20:216.
Guttman, S. I.
1989.
Eurycea longicauda. In R. A. Pfingsten and F. L. Downs (eds.),
Salamanders of Ohio, pp. 204-209. Bull. Ohio Biol. Surv.
Hutchinson, V. H.
1956. Notes on the plethodontid salamanders Eurycea lucifuga
Ireland, P. H.
1974.
(Rafinesque) and Eurycea longicauda longicauda (Green). Occ.
Pap. Nat. Speleol. Soc. 3:1-24.
Reproduction and larval development of the dark-sided salamander
Eurvcea longicauda melanopleura (Green). Herpetolosica 30:338-
343.
McDowell, W. T.
1988. Egg hatching season of Eurycea longicauda longicauda and
Bulletin of the Maryland Herpetological Society
page 59
Volume 39 Number 2
June 2003
Eurycea lucifuga (Caudata: Plethodontidae) in southern Illinois.
Bull. Chicago Herp. Soc. 23: 145.
1989. Larval period variability of Eurycea longicauda longicauda in
southern Illinois. Bull. Chicago Herp. Soc. 24: 75-78.
1 992. An observation on the egg hatching season of Eurycea longicauda.
Bull. Chicago Herp. Soc. 27:150.
Mohr, C. E.
1943. The eggs of the long-tailed salamander, Eurycea longicauda
(Green). Proc. Penn. Acad. Sci. 17:86.
Petranka, J. W.
1 998. Salamanders of the United States and Canada. Smithsonian Insti¬
tute Press. Washington, D. C.
Phillips, C. A., R. A. Brandon, and E. O. Moll.
1999. Field Guide to Amphibians and Reptiles of Illinois. Ill. Nat. Sur.
Man. 8.
Tilley, S. G.
1977. Studies of life histories and reproduction in North American
plethodontid salamanders. In D. H. Taylor and S. I. Guttman, (eds.),
The Reproductive Biology of Amphibians, pp. 1-41. Plenum Press,
New York.
Trauth, S. E., R. D. Smith, A. Cheung, and P. Daniel.
1993. Caudal hedonic glands in the dark-sided salamander Eurycea
longicauda melanopleura (Urodela: Plethodontidae). Proc. Ark.
Acad. Sci. 47:151-153.
Williams, A. A., R. A. Brandon, and J. Martan.
1984. Male genital ducts in the salamanders Eurycea lucifuga and
Eurycea longicauda . Herpetologica 40:322-330.
_ _ J. Martan, and R. A. Brandon.
1985. Male cloacal gland complex of Eurycea lucifuga and Eurycea
longicauda (Amphibia: Plethodontidae). Herpetologica 41:272-
page 60
Bulletin of the Maryland Herpetological Society
Volume 39 Number 2
June 2003
281.
W. T. McDowell 2511 South Illinois Ave. # 103 and B. A. Shepherd, Dept . of
Zoology, Southern Illinois University, Carbondale, Illinois 62901. USA.
Corresponding author E-mail: wtmcdowell@earthlink.net
Received; 28 March 2003
Accepted; 25 April 2003
Bulletin of the Maryland Herpetological Society
page 61
Volume 39 Number 2
June 2003
New Distributional Record for the Southern Leopard
Frog in Frederick County, Maryland
Wayne G Hildebrand
A Southern Leopard Frog, Rana sphenocephala utricularia (Crother, 2001),
was found crossing Putman Road in Frederick County Maryland (N 39.52768°, W
077.44574) (about 16 mi. North Lily Pons) on 20 March 2003. The single specimen
was photographed (Figure 1) and released at the capture site. Breeding choruses were
heard on private land in this area during April 2002. The subsequent capture suggests
a sustained population exists at this location. This occurrence extends the range (Har¬
ris, 1975) of Rana sphenocephala utricularia west on to the Piedmont.
■P'lli
!Slli4lli|l
ilSSIilif
—
«ft
Figure 1. Southern Leopard Frog, Rana sphenocephala utricularia (note the
characteristic light spot on the tympanum)
page 62
Bulletin of the Maryland Herpetological Society
Volume 39 Number 2
June 2003
Literature Cited
Harris, Herbert S., Jr.
1975. Distributional Survey (Amphibia / Reptilia): Maryland and the
District of Columbia. The Bulletin of the Maryland Herpetology
Society. 11(3): 73-170.
Crother, Brian I.
200 1 . Scientific and Standard English Names of Amphibians and Rep¬
tiles of North America North of Mexico, With Comments Re¬
garding Confidence in Our Understandings. Society for the study
of Amphibians and Reptiles Herpetological Circular No. 29.
Wayne G Hildebrand Maryland Calling Amphibian Coordinator
North American Amphibian Monitoring Program
Keymar, Maryland 2 17 57 , wayneh@netstorm.net
Received: 26 March 2003
Accepted: 26 March 2003
Bulletin of the Maryland Herpetological Society
page 63
Volume 39 Number 2
June 2003
News and Notes
Book Review
Life-Histories of the Frogs of Okefinokee Swamp, Georgia* North Ameri¬
can Salientia {Amir a) No. 2» by Albert Hazen Wright, 2002. xxi +509 pp. + 45 pis.
Cornell University Press, Sage House, 5 12 E. State Street, Ithaca, NY. ISBN 0-8014-
4046-7, Cloth $49.95.
The original volume was first published in 1932 by the Macmillan Com¬
pany, New York. It represents a classic among works in biology, and is unsurpassed
in herpetology. Its original edition is extremely rare, and is only found in a few major
libraries. I am honored to own an original autographed copy which I received from
the author during my high school years, and to have the opportunity to reviewing the
updated version. The present reprinted edition has been reprinted after 70 years and
is provided with a splendid new forward and afterward by J. Whitfield Gibbons de¬
scribing Wright’s college years and associations with colleagues who accompanied
Wright on the original Cornell Expeditions.
The author provides a highly enlightening review of previous studies on the
amphibian fauna of the Okefinokee Swamp area, along with a discussion of the spe¬
cies known to inhabit the region, followed by a brief discussion of the life processes
pertaining to his data provides. This is followed by a detailed life-history study for
each of the 22 species of anurans inhabiting the region. Of the 22 species, Scaphiopus
holbrooki , Bufo quercicus, B. terrestris , Acris gryllus , Pseudacris nigrita , P.
occidentalism P ocularis , Hyla andersonii, H, cinereaf H. femoralis , H. gratiosa , H.
ornata, H. squirrella, H. versicolor, Rana clamitans, R. grylio, R, heckscheri » R.
septentrionalis, R. sphenocephala, R. virgatipes and. Gastrophryne carolinensis , only
Rana aesopus has been relegated as a subspecies of Rana capita . The author pro¬
vides information on the range, general appearance, measurements, habitat, first ap¬
pearance, general habits, voice, mating, ovulation, eggs, hatching period, tadpoles,
larval period, transformation of tadpoles, food, autumn disappearance, affinities, and
lastly a bibliography for each of the species covered within the text for each of the 22
species under study. The plates provide black and white illustrations of amplexation,
eggs, mature tadpoles, tadpole mouthparts, and transformed froglets. It is surprising
that only three of the nomenclatural designations provided by Wright in his species
account differ from present day nomenclature.
Any student of amphibology will certainly welcome the reprinting of this
superb classic in the field of Herpetology. This well bound reprinting is reasonably
page 64
Bulletin of the Maryland Herpetological Society
Volume 39 Number 2
June 2003
News and Notes
priced, and should be on the bookshelf of anyone interested in anuran biology, along
with anyone interested in Herpetology or Natural History in general.
Harlan D. Walley, Department of Biology, Northern Illinois University,
Dekalb, Illinois 60115, email: hdw@niu.edu
Received 21 February 2003
Bulletin of the Maryland Herpetological Society
page 65
Volume 39 Number 2
June 2003
News and Notes
page 66
Bulletin of the Maryland Herpetological Society
Volume 39 Number 2
June 2003
News and Notes
Reptile and
Amphibian Rescue
410-580-0250
Wfe will take as many unwanted pet reptiles and
amphibians as space allows.
Leave a message with your name and number to
give up an animal for adoption;
or to volunteer to help with our efforts.
OUR CURRENT NEEDS:
• Commercial or Passenger Van
• UVB Lights • Power & Hand Tools • Bleach
• Equipment & Food • Paper Towels
www.reptileinfo.com
Bulletin of the Maryland Herpetological Society
page 67
■
Volume 39 Number 2
News and Notes
“ . i . . . . rr: — r“ -
June 2003
page 69
Society Publication
Back issues of the Bulletin of the Maryland Herpetological Society, where
available, may be obtained by writing the Executive Editor. A list of available
issues will be sent upon request. Individual numbers in stock are $5.00 each,
unless otherwise noted.
The Society also publishes a Newsletter on a somewhat irregular basis.
These are distributed to the membership free of charge. Also published are
Maryland Herpetofauna Leaflets and these are available at $. 25/page.
Information for Authors
All correspondence should be addressed to the Executive Editor. Manu¬
scripts being submitted for publication should be typewritten (double spaced)
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Indicate where illustrations or photographs are to appear in text. Cite all lit¬
erature used at end in alphabetical order by author.
Major papers are those over five pages (double spaced, elite type) and
must include an abstract. The authors name should be centered under the title,
and the address is to follow the Literature Cited. Minor papers are those pa¬
pers with fewer than five pages. Author’s name is to be placed at end of paper
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Reprints are available at $.07 a page and should be ordered when manu¬
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and must return proof preferably within seven days.
The Maryland Herpetological Society
Department of Herpetology
Natural History Society of Maryland, Inc.
2643 North Charles Street
Baltimore, Maryland 21218
Bulletin of the Maryland Herpetological Society
page 71
Am-)
BULLETIN OP THE
US ISSN: 0025-4231
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f)ecpetological
0odety
(j?40
£fpt
DEPARTMENT OF HERPETOLOGY
THE NATURAL HISTORY SOCIETY OF MARYLAND, INC.
MDHS . A Founder Member of the Eastern
Seaboard Herpetological League
30 SEPTEMBER 2003 VOLUME 39 NUMBER 3
NOV 1 9 .
BULLETIN OF THE MARYLAND HERPETOLOGICAL SOCIETY
Volume 39 Number 3 September 2003
CONTENTS
A New Cryptic Species of Pseudoeurycea (Amphibia, Caudata:Plethodontidae) of
the Leprosa Group From Central Mexico
Guillermo Lara-Gongora . . . . . . . 21
Knobloch's King Snake (Lampropeltis pyromelana knoblochi ) of Mexico a Species
Julio A. Lemos-Espinal, David Chiszar and
Hobart M. Smith . . . . . . . . . . . . 53
Disltributional and Variational Data on the
Frogs of the Genus Rana in Chihuahua, Mexico, Including a New Species
Hobart M. Smith and David Chiszar . . . . . 58
BULLETIN OF THE
mbhs
JAN 1 5 2004
Utm/w
Volume 39 Number 3
September 2003
The Maryland Herpetological Society
Department of Herpetology, Natural History Society of Maryland, Inc.
President Tim Hoen
Executive Editor Herbert S. Harris, Jr.
Steering Committee
Frank B. Groves Jerry D. Hardy, Jr.
Herbert S. Harris, Jr. Tim Hoen
Library of Congress Catalog Card Number: 76-93458
Membership Rates
Membership in the Maryland Herpetological Society is $25.00 per year
and includes the Bulletin of the Maryland Herpetological Society. For¬
eign is $35.00 per year. Make all checks payable to the Natural History
Society of Maryland, Inc.
Meetings
Meetings are held monthly and will be announced in the "Herp Talk"
newsletter and on the website, www.naturalhistory.org.
Volume 39 Number 3
September 2003
A NEW CRYPTIC SPECIES OF
PSEUDOEURYCEA (AMPHIBIA, CAUDATA:
PLETHODONTIDAE) OF THE LEPROSA GROUP
FROM CENTRAL MEXICO
Guillermo Lara-Gongora
ABSTRACT
A morphological and ecological well differentiated population of Pseudoeurycea
belonging to the leprosa species group was collected at several localities in the cen¬
tral section of Eje Neovolcanico Transversal in central Mexico. It is named as a dis¬
tinct species : Pseudoeurycea tlilicxitl. A comparison of meristic, morphometric and
coloration characters and ecological preferences is made with all the other recog¬
nized species of the leprosa group, and a more detailed analysis is made on the 2
known sympatric species with which it occurs, namely : Pseudoeurycea leprosa and
Pseudoeurycea altamontana. Pseudoeurycea tlilicxitl differs from R leprosa as well
as from P. altamontana in its lower number of maxillary - premaxillary teeth, and
vomerine teeth, longer limbs, less webbing on hand and feet, coloration, body pro¬
portions, and ecological preferences. The status of P. altamontana is questioned.
INTRODUCTION
In spite of the strong attention the Bolitoglossini tribe of plethodontid sala¬
manders has received in the last 30 years (particularly the supergenus Bolitoglossa)
knowledge on systematics and ecology of many neotropical salamander taxa is still
partial or even unstable. See Brame (1968), Wake and Lynch (1976), Crump (1977),
Maxson and Maxson (1979), Maxson and Wake (1981), Hanken (1983), Lynch, Wake
and Yang (1983), Wake and Elias (1983), Papenfuss, Wake and Adler (1983), Elias
(1984), among others.
Most of the interest and work done has been concentrated on tropical southern
forms, and the accumulated knowledge on some more northern, boreal or transitional
autochthonous groups is scarce, as it is true for several central Mexico highland spe¬
cies of the genus Pseudoeurycea. (For the most recent taxonomic review of the genus
Pseudoeurycea and its status see Wake and Lynch, 1976.)
The leprosa group has not been a stable taxonomic entity since different au¬
thors do not agree on its species content (see Taylor, 1938, 1944 ; Wake and Lynch,
1976 ; Maxson and Wake, 1981; and Lynch, Wake and Yang, 1983). The 9 species
recognized in this group by Wake and Lynch (1976) are distributed in boreal, mesic
Bulletin of the Maryland Herpetological Society
page 21
Volume 39 Number 3
September 2003
forests, from above 2000 m asl. to tree line (except for Pseudoeurycea nigromaculata)
in central and southern Mexico. Four species occur on the Eje Neovolcanico Trans-
versal, 4 more occur in southern Sierra Madre Oriental, and 1 species occupies Sierra
Madre del Sur. All of them but P. leprosa are microendemic and are known from only
a few locations around their type locality.
Even though a diagnosis of the leprosa group has never been given, the group
is generally characterized and distinguishable from other groups at the morphologi¬
cal level by having small to medium size species, with both hands and feet mainly
unwebbed and having a dark brown background coloration, usually with light spots,
bands or reticulations. A glandular circular area above the hind limb insertion is present
in most species. The head is flat on its top, and the digits and toes are slender, taper¬
ing toward tip (except for P. juarezi, P. nigromaculata and P. robertsi in which are
flattened.) Limb size is much variable, from short (P. firscheni, P. leprosa, P. longicauda,
and P. mystax) to large or very large (P. anitae and P. juarezi). The number of maxil¬
lary and premaxillary teeth as well as vomerine teeth is also much variable, and goes
from few teeth (P. longicauda and P. robertsi) to many (P. firscheni, P. juarezi, and P.
nigromaculata). All species are terrestrial (semifossorial) but one is presumably some¬
what troglodytic (P. anitae), and another one scansorial (P. firscheni).
Pseudoeurycea leprosa (Cope, 1869) is represented by large series in museum
collections. It is an old and well known species, and the most abundant and wide
ranging species of its group. Nevertheless, it has a very unstable taxonomic history as
it is shown by the number of genera and species in which it has been placed. There
are at least 9 different names to which this species has been allocated: Spelerpes
leprosus (Cope, 1869), S. laticeps (Brocchi, 1883), S. orizabensis (Blatchley, 1893),
S. gibbicaudus (Blatchley, 1893), Oedipus leprosus (Dunn, 1918), O. cephalicus (in
part, Dunn, 1926), O. orizabensis (Taylor, 1938), Bolitoglossa leprosa (Taylor, 1941),
and finally Pseudoeurycea leprosa (Taylor, 1944).
Smith and Smith (1976) correlated the above mentioned characteristics (high
abundance, conspicuity, distinctiveness, and antiquity of knowledge) with a higher
accumulation of synonyms for Mexican lizards. The same could be applied to this
species, but also at least the following reasons may have contributed to the species
unstable taxonomic history : 1) high geographical, ontogenic, and sexual variability
within the species; 2) ambiguity and scarcity of good taxonomic parameters and cri¬
teria currently in use for this species; 3) Limited number of specimens studied in
several of the old species descriptions; and 4) Absence of statistical treatment and
data analysis.
page 22
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
September 2003
Lynch, Wake and Yang (1983) found a great genetic variability in peripheral
Pseudoeurycea leprosa populations. But because the high genetic variability and
morphological overlapping they found, they didn’t see fit to propose any taxonomic
changes.
From a study on spatial resources partitioning by plethodontid salamanders at
El Capulin region, in the state of Mexico (Lara and Ortega, 1980, unpublished) it was
found that 4 species inhabited sympatrically the same area, among which there was a
cryptic form of Pseudoeurycea leprosa, morphologically and ecologically distinct
from it.
This study is concerned with the taxonomic description of this new species of
the leprosa group, and its phyllogenetic relationships with the other group members,
with emphasis on P. leprosa and P. altamontana which are the other 2 sympatric spe¬
cies in the group.
MATERIALS AND METHODS
Sixty eight specimens were checked, 44 belonging to P. leprosa and 14 belong¬
ing to the new taxon herein described. All specimens were or had been fixed in 10%
formalin and preserved in 70% ethanol. Specimens are deposited in the 2 following
collections: Museo Alfonso L. Herrera, Facultad de Ciencias, Universidad Nacional
Autonoma de Mexico (MZFC), and Instituto de Biologia, Universidad Nacional
Autonoma de Mexico (IBH), both in Mexico city.
For the comparative analysis 57 parameters were taken into account:
MERISTIC: (4 parameters) maxillary-premaxillary teeth, vomerine teeth, sepa¬
ration of adpressed limbs, costal grooves.
MORPHOMETRIC: (28 parameters) snout to vent length (SVL), tail length
(TL), snout to arm length (SAL), armpit-groin length (AGL), head width (HW), head
length to gular fold (HLG), head length to quadratoyugal articulation (HLQ), foreleg
length (FLL), hind leg length (HLL), eye diameter (ED), snout length (SL), interor¬
bital width (IOW), intemarial width (INW), eyelid width (EW), TL/SVL ratio, FLL/
SVL ratio, HLL/SVL ratio, HLG/SVL ratio, HLQ/SVL ratio, SL/SVL ratio, AGL/
SVL ratio, HW/SVL ratio, SAL/SVL ratio, ED/SVL ratio, IOW/SVL ratio, INW/
SVL ratio, EW/SVL ratio, and separation of adpressed limbs/SVL ratio.
COLORATION: (6 parameters) maxillary-premaxillary teeth, dorsal body,
ventral body, lateral body, tail, and limbs.
Bulletin of the Maryland Herpetological Society
page 23
Volume 39 Number 3
September 2003
STATE CHARACTERS: (19 parameters) white lower eyelid, light parietal
spots, digits and toes shape, hand webbing, feet webbing, digit formula, toe formula,
first digit size, dorsal surface of body (texture), palate shape (arrangement of choa-
nae and vomerine-parasphenoid series of teeth), nasolabial groove, cephalic dorsal
musculature, digital pads, mental gland, dorsal head profile, lateral head profile, can-
thus rostralis, glandular area above insertion of hind limbs, and first transverse cephalic
groove.
The criteria for the analysis of many of these parameters are those as Taylor
(1938). All data were tabulated and 23 selected parameters were analyzed statisti¬
cally using the chi-square, t-student, U-Mann-Whitney, and simple linear regression
statistics, with a statistical significance of p=0.05 for the null hypothesis. Null hy¬
pothesis was that R leprosa and the new taxon described here belonged to the same
population and represented the same taxonomic entity. For the remainder 34 param¬
eters simple statistical concentration measures were calculated. Measurements and
counts were obtained in a uniform manner. Measurements were taken with a dial
caliper to the nearest 0. 1 mm.
For additional comparisons with Pseudoeurycea altamontana and P. leprosa) I
followed Taylor descriptions (1938.)
RESULTS
From the statistical analysis of the 23 aforementioned parameters between P.
leprosa and the new taxon hereby described, 11 proved to be of statistical signifi¬
cance at p=0.05 thus rejecting the null hypothesis and supporting the distinctiveness
of the new taxon and its taxonomic designation as a new species (refer to figure 1 for
the parameters characterization.)
The 1 1 statistically significant parameters were: TL/SVL ratio, maxillary-pre¬
maxillary teeth number, vomerine teeth number, FL/SVL ratio, type of palate, sepa¬
ration of costal folds when the limbs are adpressed (ADP-LIMBS), maxillary-pre¬
maxillary teeth color, hand webbing, feet webbing, first digit size, and the decreasing
sequence of toes size (toe formula.) (Refer to figure 3 for statistically significant
parameters.)
The SVL comparison showed no statistical differences between P. leprosa and
the new taxon, although the latter attains higher lengths than P. leprosa.
Linear regression analysis of individual characters as a function of SVL, par¬
ticularly TL/SVL ratio, FL/SVL ratio, and ADP-LIMB S/S VL ratio, proved to be good
discriminating parameters between the 2 taxa with almost no overlap in the FL/SVL
page 24
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
September 2003
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page 25
Volume 39 Number 3
September 2003
Third roc
Foot structures. Phafatwcs arc
numbered. Tarsai bones £\rc
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Figure 2. Schematic representation of foot structure and webbing in R leprosa
and P tlilicxitl
and ADP-LIMBS/SVL, which means that the new taxon has much longer feet than P.
leprosa (see figures 5, 6 and 7.)
In regards with the palate type, the new taxon shows a constant arrangement of
2 series of vomerine teeth slightly curved and separated medially. The parasphenoid
series are separated form the vomerine series by a distance approximately equal to
the length of one vomerine series. The choanae are slit-like. In contrast, P. leprosa has
a variable palate with choanae sometimes small and rounded and the vomerine series
closer to each other.
Teeth coloration is constant in P. leprosa. The teeth are conspicuously red-
tipped (RT), while in the new taxon they are variable in color, sometimes they are
entirely transparent, but they are never conspicuously red-tipped.
Hand webbing is variable in both taxa but the new taxon has higher values for
the vestigial character than P. leprosa (from base of first phalanx of second finger to
base of first phalanx of third finger.)
page 26
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
September 2003
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Bulletin of the Maryland Herpetological Society
page 27
Figure 3. Statistical characterization for those parameters with statistically significant values
Volume 39 Numbers
September 2003
Feet webbing is constantly vestigial in the new taxon (base of first phalanx of
second toe to base of first phalanx of third and fourth toes) and variable in P. leprosa,
but with low values for the vestigial character (see figure 2.)
First digit is small in P. leprosa as compared to vestigial (very small, with
almost no free tip) in the new taxon. The decreasing sequence of toes size (toe for¬
mula) is constant in R leprosa, but variable in the new taxon. In the latter third and
fourth toes are sometimes of equal size (see figure 2)
Five additional parameters showed significant differences between P. leprosa
and the new taxon. These parameters were 3 coloration characters: parietal spots,
lateral coloration pattern and contrast between dorsal body and dorsal tail coloration,
and 2 state characters: position of rictus oris in regards with the first transversal groove,
and the postorbital groove. All parameters showed significant differences (= 90%)
between both taxa (see figure 1 1 .)
The parietal spot is almost always present in P. leprosa as a lighter area that can
be creamish, pinkish or light brown. It is usually well evident. In the new taxon is
always absent.
The lateral coloration pattern in P. leprosa can be described as typically con¬
sisting of a gray “sparkling” pattern of tiny round dots. It is located in the upper
lateral area (laterodorsal) and it is markedly different from the lower lateral colora¬
tion (lateroventral) which is darker and without light dots. In the new taxon this
pattern is always absent and the lateral coloration is the same as the dorsal one, con¬
sisting or individual or fused brownish spots or blotches.
The tail dorsal coloration is noticeably lighter than the body dorsal coloration
in P. leprosa. It has more spots and light pigmentation. In the new taxon both colora¬
tion patterns are alike. The tail is not lighter or darker than the body or if lighter it is
not conspicuously so.
The rictus oris is always separated from the first head transversal groove in P.
leprosa (type A), while it is almost always in contact with the groove in the new taxon
(type B) (See figure 12.)
The 2 sections of the postorbital groove (anterior and posterior) in P. leprosa
form different angles. The anterior section has a greater angle and it is above the
posterior section. On the other hand, in the new taxon both sections form a continu¬
ous and almost straight line (see figure 12.)
In accordance with data presented by Taylor (1938) an attempt was made to
page 28
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
September 2003
analyze similarities and differences among the new taxon and O. altamontanus (=P.
altamontana), O. orizabensis (=P. leprosa), and O. leprosus (=P. leprosa) based on
their original descriptions. No attempt of statistical analysis was made because the
reduced number of specimens of O. altamontanus and O. leprosus. For the same
reason no variance analysis for the regression of characters was possible since F=0.
Neither did the analysis was made for O. orizabensis because it has been widely
recognized as a junior synonym of P. leprosa (Taylor, 1945, Smith and Taylor, 1948.)
Therefore, only numerical trends were observed. To fit this purpose I transformed the
real measurements given by Taylor to ratios and considered all characters as depen¬
dant or being a function of SVL in order to make possible a comparison of these 4
taxa (the new taxon, and O. altamontanus, O. orizabensis and O. leprosus.) These
data are presented in figure 4. The number of specimens checked is too low to be
conclusive. Although there are some evident and contrasting differences.
The new taxon has the lowest values for character 2, which means that its
limbs insertions are closer to each other. This same taxon has intermediate values for
character 6; i.e. it has longer hind limbs than O. orizabensis and O. leprosus, but
smaller than those of O. altamontanus. The new taxon also differs from the latter in
characters 8 and 10, having higher values and, thereby, longer snout (twice a long as
that of O. altamontanus), and more separated nostrils. The new taxon differs from O.
orizabensis in characters 1, 4, 5 and 3, and from O. leprosus in characters 5 and 3,
thus having longer snout, head and forelegs, and a wider head.
Pseudoeurycea tlilicxitl nov. sp.
(see figure 8)
The species epithet comes from 2 Nahuatl words (Nahuatl is one of the 2 most
important native Mexican languages spoken today): tliltic = black, and icxitl = foot,
in reference to the conspicuous “black feet” that typify this species.
Holotype.- MZFC-01461, adult male. Collected on July 13th, 1980 by Jose
Antonio Hemandez-Gomez and the author at 3 Km S of Laguna Quila, near Las
Trancas mountain brook, in Parque Nacional Miguel Hidalgo (Lagunas de Cempoala),
state of Mexico. Elevation: 2950 m asl. It was found under a log, in a humid fir forest
(Abies religiosa.) (see figure 9)
Paratypes.- MZFC-01461 -2 from same date and locality. MZFC-01253 (2 speci¬
mens) from El Capulin, state of Mexico. Collected on June 16th, 1979 by Jose Anto¬
nio Hemandez-Gomez and the author, at an elevation of 3050 m asl in a pine forest
(Pinus hartwegii.) MZFC-03985 to MZFC-03990 (6 specimens) from El Capulin
area (Cerro Cadena, C. Pelado and C. Malacatepec). Collected in August 1980 by
Bulletin of the Maryland Herpetological Society
page 29
Volume 39 Number 3
September 2003
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page 30
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
September 2003
Juan Jose Ortega Leon and the author in pine forests (Pinus leiophylla; 4 specimens)
and fir forest (Abies religiosa; 2 specimens) at 3000-3500 m asl. IBH-3683 from San
Rafael, state of Mexico. Collected on September 17th, 1977 by Gustavo Casas- Andreu
and Aurelio Ramirez-Bautista. Elevation 2700 m asl; collected in a pine-oak forest.
IBH-3683-2 from Western slope of Mountain Iztaccihuatl. Collected on September
17th, 1977 by Gustavo Casas- Andreu and Aurelio Ramirez-Bautista. IBH-3682 from
Cerro La Cima, Distrito Federal; collected on June 3rd, 1979 by Fernando Cervantez-
Reza. IBH-2715-3, from Desierto de Los Leones (Cruz Blanca), Distrito Federal.
Collected on June 29th, 1979 by Zeferino Uribe-Pena (see figure 10)
GEOGRAPHICAL RANGE
Central section of Eje Neovolcanico Transversal, at high elevations (above
3000 m asl) in Ocuilan, Ajusco, Las Cruces, and Sierra Nevada mountain ranges, in
the states of Mexico, Morelos and Distrito Federal (see figure 13), and very likely in
western Puebla.
DIAGNOSIS
A medium-sized species, SVL 30-64 mm, x 45.5 mm; maxillary-premaxillary
teeth number (both sides) 27-48, x 35.4; maxillary-premaxillary teeth color: trans¬
parent: 57.15% or slightly red-tipped: 42.85%; prevomerine teeth number (both sides):
9-24, x 18.3; choanae slit-like: 100%; hand webbing poorly developed: 42,8% or
vestigial: 57.16%, from the base of first phalanx of second finger to first phalanx of
third finger; feet webbing vestigial: 100%, from the base of first phalanx of second to
base of first phalanx of third to fourth toes; first finger rudimentary: 85.72% or small:
14.28%; toe formula (from biggest to smallest toe): 3425 1 : 64.29% or third and fourth
toe of equal size: 35.71%; TL/S VL ratio: 0.635-0.936, x 0.807; FL/SVL ratio: 0.193-
0.318, x 0.227.
Black body coloration with numerous cream or brownish rounded or elon¬
gated spots, sometimes fused and forming a reticulated pattern all over the back and
tail. Dorsal surfaces of arms and legs completely cream; hands and feet entirely black;
ventral surfaces gray-black, sometimes with few, scattered and conspicuous light
rounded spots (see figure 8.)
DESCRIPTION OF HQLQTYPE
Head flat between orbits, but surface roughened, deeply pitted; snout rounded
in dorsal profile; canthus rostralis rounded, but distinctive. A slight depression from
anterior angle of eye toward nostril; nostril almost at tip of snout; distance between
nostrils equal to interorbital width (1.9- 1.9 mm). Width of eyelid slightly less than
Bulletin of the Maryland Herpetological Society
page 31
Volume 39 Number 3
September 2003
SVL (mm)
Figure 5. TL/SVL ratio linear regression. Comparison between and br Ho: b2
Ha: bj=b2 t “student” statistic. T= 9.25, t005(2) 47= 2.012. H=rejected, HA=accepted.
p<0.001 . 11= values do not adjust to a line; Revalues adjust to a line. r2~proportion
of total variation in “Y” explained by the regression (determination coefficient).
S Vx“variance °f “Y” after taking into account dependence of “Y” over “X”. SY.X=
standard error of estimate or regression standard error.
interorbital distance (1.5 mm). Posterior ends of eyelids fitting under a fold. Length
of eye bigger than its distance from tip of snout (2. 8-2.0 mm). A deep longitudinal
groove that runs from eye back to the end of gular fold, nearly straight (postocular
groove as defined by Baird, 1951.) A transversal groove runs from a little behind
page 32
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
September 2003
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Bulletin of the Maryland Herpetological Society
page 33
Volume 39 Number 3
September 2003
ADF, LJMB5 (ccsfat Wds)
4
3
2
1
0
-t
y^2,444tc,ar7x
■ i i
bl ' \jV s 'hl 'JM
61 , 65
Sn. (nrn)
P. leprvsa JL
P. tliffcxitt JL
F^S AT
F^5,Q9
F0.05 a ). 143=4,06
F0 05 aX 1.12=4.75
He1 Rejected
Rejected
Actxipied
Ha: Accept ad
pz. 0.0005
0.025 LPi. 0.05 1
r^-O.lS
0.30
Sy.x=0.58 ccstci fatd
5y.x-l.ll costal feki
Figure 7. ADP.LIMBS/SVL ratio linear regression. Comparison between b1 and b2.
H: b = b2
Ha: b,? b2 t “student” statistic. T= 8.42, t005(2) 56= 2.003. H=rejected,
HA=accepted. p«0.001 Ho=values do not adjust to a line; HA-values adjust to a
line. r2=proportion of total variation in “Y” explained by the regression (determina¬
tion coefficient). S2y. ^variance of “Y” after taking into account dependence of
“Y” over “X”. Srx= standard error of estimate or regression standard error
rictus oris to dorsolateral portion of head, intersecting the longitudinal groove.
Mentolabial gland indistinct.
Costal folds 12-12 (left-right); a deep medial dorsal groove that goes from
posterior insertion of hind limbs to the transversal groove of head, where it bifurcates
and runs toward eyelids. Ten costal groves continue across venter; anal region very
slightly swollen; interior of cloaca papillated.
Arms and legs elongated, separated by half a costal fold when the limbs are
adpressed; a very slight indication of webbing; first finger extremely reduced (vesti¬
gial); the tip barely free; digital formula (from biggest to smallest): 3241; toe for¬
mula: 3425 1 ; webbing on hand from base of first phalanx of second finger to first
phalanx of third and fourth finger. Webbing on feet vestigial, from base of first pha¬
lanx of second toe to base of first phalanx of third toe to first phalanx of fourth toe;
tips of digits swollen underneath and tapering from base to tip.
page 34
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
September 2003
Figure 8. Live specimen of Pseudoeurvcea tlilicxitl from El Capulm, Mexico. Photo¬
graph by Jose Antonio Hemandez-Gomez
Skin of head and dorsal surface of body roughened and pitted; belly and limbs
smooth; the belly with fine transversal grooves; hedonic gland not visible externally;
no glandular area behind and above posterior insertion of hind limbs.
Vomerine teeth in 2 very slightly curved series of 9-10 teeth, separated medi¬
ally and extending to the interior borders of choanae; palatine teeth in 2 series of 2-3
longitudinal rows, contiguous anteriorly, diverging and widening posteriorly, sepa¬
rated from the vomerine series by a distance approximately equal to the length of the
vomerine series. Premaxillary teeth 6, small, curved, claw-like, and not piercing lip;
10-10 maxillary teeth; tongue boletoid, subcircular.
Length of snout 13.2 mm; armpit-groin length: 23.1 mm; head width: 7.2 mm;
head length: 8.6 mm; foreleg length: 10.7 mm; hind leg length: 11.5 mm; eye diam¬
eter: 2.8 mm; snot length: 5.4 mm; interorbital width: 1.9 mm; distance between
nostrils: 1.9 mm; snout to gular fold length: 10.7 mm; eyelid width: 1.8 mm; tail
length: 37.2; SVL: 45 mm.
COLORATION
(Preserved specimens) Body entirely black with cream to brownish spots, some
or all fused to form a reticulated pattern all over the back and dorsal surfaces of tail
and limbs. Hand and feet entirely black without light markings or with few, small
spots. This color contrasts sharply with light spots on upper portions of limbs. Ven¬
tral surfaces of body, tail and limbs are uniform gray black, sometimes with light
rounded spots more conspicuous in throat and gular regions. Sides of body entirely
black or with large light reticulation or spots.
Bulletin of the Maryland Herpetological Society
page 35
Volume 39 Number 3
September 2003
Figure 9. Holotype of R tlilicxitl. MZFC-01461, adult male from Lagunas de
Zempoala, Mexico. Photograph by J. A. Hernandez- Gomez
Figure 10. Comparison between R tlilicxitl (2 specimens on the left) and R leprosa
(3 specimens on the right). Photograph by I. Antonio Hemandez-Gomez
page 36
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
September 2003
Parameters
P leprosa
P. tlilicxitl
Parietal spots
Lighter than
surrounding areas;
pinkish, creamish
or brownish
Present: 90.24%
Absent (or not evident):
9.76%
Absent: 100%
Lateral coloration
pattern
Upper (laterodorsal) and
lower (lateroventral) areas
well differentiated
Upper consisting of a light
gray “sparkling” pattern
Lower same as ventral surfaces
(darker and without light
pigmentation) Present: 97.56%
Absent or inconspicuous: 2.44%
Upper and lower
sections alike.
Pattern consisting of
same dorsal coloration,
with brownish
individual or fused
spots or blotches in a
reticulated pattern,
contrasting with
uniform dark ventral
surfaces
100%
Contrast between
body dorsal
coloration and tail
dorsal coloration
Tail dorsal coloration
contrastingly lighter than
body dorsal surfaces
In 92.5% of specimens
No contrast in 7.5 %
No contrast between
body dorsal coloration
and tail dorsal
coloration or if lighter
not conspicuous
100%
Rictus oris
Always separated from
first head transversal groove:
100%
(type A) (see figure 12)
In contact with first
head transversal groove:
80% or separated from
it: 20%
(type B) (see figure 12)
Postorbital groove
Anterior and posterior sections
at different angles. Anterior
section at higher position than
posterior section, forming a
broken line:
100%
(type A) (see figure 12)
Anterior and posterior
sections at the same
angle and level,
forming a continuous
and almost straight line:
100%
(type B) (see figure 12)
Figure 1 1 . Additional significant coloration and state parameters between
P leprosa and R tlilicxitl
Bulletin of the Maryland Herpetological Society
page 37
Volume 39 Number 3
September 2003
Fig. 12 Schematic representation of head lateral profiles as characterized by
typical P leprosa (type A) and P. tlilicxitl (type B.) Notice main differences in terms
of anterior-posterior sections of cephalic groove and rictus oris reaching or not the
first transversal groove. Typical P leprosa have 2 sections of postorbital groove at
different angles, while typical P. tlilicxitl have a continuous and almost straight line.
Rictus oris is separated from the first transversal groove in P. leprosa , while it reaches
and is in contact with the groove in P. tlilicxitl.
HABITAT AND HABITS
R tlilicxitl prefers open and somehow dry pine forests with bunchgrass as the
herbaceous stratum. It can also be found in wetter environments coexisting
microsimpatrically with P. leprosa and Chiropterotriton chiropterus, in fir forests of
Abies religiosa, and mixed temperate forests of Pinus spp., Abies religiosa, Arbutus
glandulosa, Alnus jorullensis and Quercus spp., from 2700 m to 3500 m asl. It seems
to prefer logs for microhabitat as hiding daylight shelters. One specimen was found
under a soil mound and another under an extrusive volcanic rock.
P. tlilicxitl is an uncommon species and it has a relative abundance much lower
than its relative P. leprosa. A differential exploitation of forest macro and micro re¬
sources truly occurs between P. tlilicxitl and the 3 other microsimpatric species which
it lives together with: P. leprosa, P. cephalica and Chiropterotriton chiropterus. No
niche overlap occurs among them (Lara and Ortega, unpublished study.) Other am¬
phibians and reptiles that occur within P. tlilicxitl range are: Crotalus triseriatus, C.
transversus, Thamnophis scalaris, Storeria storerioides, Sceloporus mucronatus, S.
palaciosi, S. anahuacus, S. aeneus, Barisia imbricata, Phrynosoma orbiculare,
Pseudoeurycea cephalica, P. belli, and Chiropterotriton chiropterus, among others.
page 38
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
September 2003
i - 1 WO Km □ Oulf of Mexico □ 0- 200m HU WO- 1000m
miooo- 2ooom mmo- 3000m msooo^ooom
Figure 13. Geographical distribution of R tlilicxitl. The map shows Eje
Neovolcanico Transversal in central Mexico. Letters stand for highest mountain peaks:
A= Cerro Tancftaro, Michoacan, B= Nevado de Toluca, Mexico, C= Iztaccihuatl,
Mexico-Puebla, D= Popocatepetl, Mexico-Puebla, E= Malinche, Tlaxcala-Puebla,
F- Cofre de Perote, Veracruz, G~ Pico de Orizaba, Veracruz. Numbers show P tlilicxitl
localities: 1= Cruz Blanca (Desierto de Los Leones), Distrito Federal, 2= El Capulfn,
Mexico, 3= Lagunas de Zempoala, Mexico, 4= San Rafael, Mexico, 5= Western slope
of Iztaccihuatl, Mexico., 6= Cerro La Cima, Distrito Federal.
DISCUSSION
The data analysis shows clear and conspicuous differences among P. tlilicxitl
and all other species of the leprosa group (see figure 14), particularly as compared to
P. leprosa itself, its assumed closer relative (see also figures 1-7 and 11-12.) Superfi¬
cial similarities between P. tlilicxitl and P. leprosa, such as size, somehow similar
coloration and sympatry (the “mimetic” character as “cryptic species”), has led many
researchers to take P. tlilicxitl as “odd” P. leprosa specimens and include them both in
the same jar labeled as “P. leprosa”, thus obscuring even more their real differences.
Since 1945, Taylor proved conspecificity of Oedipus orizabensis and O. leprosus
with P. leprosa. This synonymy has remained unchallenged up to the present times
even though the original descriptions of the types of these taxa seem to differ in some
important respects. S. leprosus types (described by Dunn, 1926) have adpressed toes
either meeting or separated by 1 or 2 costal folds (as in P. tlilicxitl). O. gibbicaudus
Bulletin of the Maryland Herpetological Society
page 39
Volume 39 Number 3 September 2003
P. dtamontana
P. anitae
R ftrscheni
R tlilkxUl
R wared
A. BODY
SIZE
(Max.
SVL)
B. LEG
Small (48.6)
Small (50.0)
Medium (60.0)
Medium (64.0)
Small (50.0)
SIZE
**Small (slightly
Large (Touch
Small
Large-very large
Large-very large
larger than
or overlap
(separated by
(Slightly
(Slightly
P longicauda )
0.5)
slightly more
separated to
separated to
*Medium Large
(touching)
than 2)
overlap 3-5)
overlap 2)
C.
Present
?
Present
Present or
Present
GLAND¬
ULAR
Absent
AREA OVER
HINDLEG
D. FOOT
?
Large (Larger
Small
Large
Medium
SIZE
than P. mystax)
(Half the size
(Slightly larger than
of P leprosa)
R fmchani)
E. TOES &
?
Short
Long and slender
Long and slender
Short, broad
DIGITS
(But larger than
(But flat, not
Tapering toward
and flattened,
R mvstax): tapering
tapering
tips with well
slightly expanded
toward tip
conspicuously
developed pads
toward tips)
under tips
F.
♦Slightly
Very slightly
Slightly
Very slight to
Moderately
WEBBING
webbed
developed
developed
vestigial
developed
ON HANDS
(distal end
(to base of
(distal end of
(To base of
(ONLY ON
of first
second
first phalanx to
first phalanx)
THIRD DIGIT
r.
phalanx)
phalanx)
its base)
WEBBING
♦Slightly
Very slightly
Moderately
Vestigial
Moderately
ON FEET
webbed
developed
developed
(to base of
developed (to
(ONLY
(distal end
(to base of
first phalanx)
base of second
THIRD TOE)
of first
second
phalanx)
phalanx)
phalanx)
H. DIGITAL F: 34251
H: 3241
H: 3241
H: 3241
H: 3241
FORMULA
F: 34251
F: 34251
F: 34251 OR 43251
F: 43521 OR 3=4
(DECREAS¬
ING SIZES)
I. TAIL
Moderate
Moderate
Moderate
Small to Moderate
Moderate
SIZE
(**X=0.820;
(0.757-0.930,
(0.926, n-1 J)
(0.835-0.938
(0.949, n=lAF)
*X=0.967)
X=0.822)
X=0.807)
J. COSTAL
GROOVES
12
13
14
12
?
Figure 14. Comparative data among species of leprosa group
page 40
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3 September 2003
R leprosa
P. longicauda
Ptnystax
R nwromaculata
P. robertsi
P. tlahcuiloh
Medium (63.0)
Medium (65.3)
Small (47.5)
Small (56.5)
Small (57.0)
Medium (61.8)
Small (Smaller
than in P.
longicauda ;
separated by 2-5)
Small
(Separated)
Small
(Separated
by 2)
Medium
(Separated by
1-1.5, but
touching in
juveniles)
Medium
(Separated
by 1 or
touching in
juveniles)
Medium
Present
Present
?
Present
?
?
Small
(Smaller than
P. longicauda)
?
(Larger than in
P. leprosa)
Small
(Stout)
?
?
Large
Long and slender
More or less slender Short
Broadly, flattened
^Flattened, but
(?) (Apparently
Tapering toward
tip
but not long
Tapering
toward tip
and truncated at
tips. Digits very
broad
at tips which
are inflated
and rounded
vestigial or
slightly
developed
Very slight
(To distal end
of first
phalanx)
Very slightly to
vestigial (From
distal end of
third phalanx to
its base)
Very slightly
developed
(To distal end
of first phalanx)
Slightly developed
(To distal end
of first phalanx)
* Vestigial
(?) (Apparently
vestigial or
slightly
developed)
Very slight
(to distal end
of first phalanx)
Very slight
(to distal end
of first phalanx)
Very slightly
developed
(To distal end
of first phalanx)
Vestigial
(to base of
first phalanx)
"“Vestigial
(?) (Apparently
vestigial or
slightly
developed)
F: 34251
H: 3241
F: 34251 OR 43251
H: 3241
F: 34215
H: 3241
F: 34521 OR 43521
H: 3241
F: 34215
(?) Apparently
H: 3241
F: 34251
Moderate
Slightly large
Small to
Large
Moderate
Moderate to
(small to slightly
large) (0.875-1.091,
X=0.864)
(X=0.980)
moderate
(0.642-785,
X=0.718)
(1.500-1.289
X=1.232)
(**0.800)
(*0.889-0.991,
X=0.911)
slightly large
(0.810-1.030,
X=0.950)
12
13
? Continuous over 13
dorsum,
projecting
anteriorly at
middorsum)
13
?
Bulletin of the Maryland Herpetological Society
page 41
Volume 39 Number 3
September 2003
P altamontana
P. antiae
P. firscheni
R tlilkxiii
R marezi
K. MAX-
Moderate
Moderate
Many
Few
Many
PREMAX
TEETH
X=47
59-85
84-100
27-48, X=35.42
74-100
(ADULTS)
L
Moderate
Moderate
Many
Few
Moderate
VOMERINE X=25
24-26
32-42
9-24, X=18.5
24-30
TEETH
Series widely
Short feebly
Curved, barely
Slightly curved
(ADULTS)
separated medially curved series,
separated
series; separated
curving
perpendicular to
median axis; widely
separated
medially
M.
BG: Dark (?)
BG: Moderately
BG: Moderaly dark
BG: Very dark
BG: Moderately
DORSAL
(Purplish-violet
dark (Dark
(Grayish or
(Black or very dark
dark to dark (Light
BODY
S: Light (Cream
brown-reddish
brownish-gray
brown) S: Moderately brown to blue-black)
COLOR-
flecks and
brown) S: Dark
S: Dark (Few very
light (Lighter than
S: Light A light red
ATION
brownish-lavendar
(2 series of dark
small black
body; small,
or orange middorsal
clouding)
irregular
rounded spots.
brownish and
band
dorsolateral spots
Occasionally
abundant but usually
or bars)
some light
many of them fused
irregular spots
to form a
may be present)
reticulation
R TAIL
BG:?
BG:?
BG: Same as body
BG: Very dark
BG: Dark
COLOR¬
S:?
S: Dark dorsal
(Tip of tail reddish-
(Black or very
(Same as on body)
ATION
spots or bars
brown)
dark brown)
S: Light (Same as
S: Large, irregular
S: Same as on body
on body but with
few to moderately
but lighter
more “black holes” on
abundant light
(Brownish to
it; size and shape
spots (Silvery-
creamish and
irregular)
white to pinkish)
more extensive;
more or less
continuous over
the tail as a band)
0.
BG: Dark
BG: Light
BG: Moderately
BG: Dark
BG: Light
VENTRAL
(Purplish-
(Whitish)
dark (Gray or
(Gray to black)
(Creamy-yellow)
SURFACES violet)
S: Nearly
blackish; a darker,
S: Light
S: Dark
COLOR¬
S: Immacute
immaculate
ventrolateral band
(Only occasional
(irregular spotting
ATION
conspicuously
separated lateral
from ventral
S: Light
(Minute, whitish,
abundant; even
more under tail)
rounded and light
of dark pigment)
Figure 14. Continued
page 42
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
P. leprosa
R lomncauda
Moderate
Few
Moderate
45-84, X-64.25
X=41
57-66
Moderate
Few
Moderate
12-31, X=22.72
X=19
Holotype=30
BG: Very dark
BG: Dark to
Rows curved
inward and
backwards;
scarcely
separated
BG: Moderately
(Dark brown, slate
very dark
dark (Slate-gray)
or black) S: Lighter (Gray-brown
S: Dark
(Extensive irregular to nearly black)
(Small dark
light brown to
S: Light A more
blotches in a
reddish-brown;
or less indistinct
more or less
manyof them fused) (sometimes pale
linear
middorsal stripe
arrangement
BG: Very dark
BG: Dark
BG: Moderately
(But often totally
(Same as on
dark
covered by
body)
(Same as on
spotting)
S: Light
body)
8: Lighter
(Same as on
S: Light
(Same as on
body but more
(Pale gray to
body but lighter;
extensive)
white blotches
sometimes
on its sides)
creamish or
pinkish more
extensive)
BG: Dark
BG: Slightly light
BG: Very dark
(Gray-black)
to slightly dark
(Dark gray or
S: ?
(Pale gray to slaty)
almost black)
S :?
S: Absent
September 2003
R meromacalaiu
R roberisi
P. tlahcuiloh
Many
*Few to **Very
Moderate
94-105
few *4248
**X-28
46=58 X=43.6
Moderate
Few
Moderate
26-30
Long, curved
backwards;
scarcely
separated
17
Curved series;
separated
27
BG: Slightly dark
BG: Light
BG: Dark
to dark
S: Darker (A
(black)
(Gray-black or
well developed
S: Dark brown
dark lavendar)
tan, red-brown or patches, and
S: Very dark
fawn middorsal
numerous tiny
(Black spots; some
veriagated
green flecks.
occasional silvery
flecks on head)
stripe)
Light areas on
the occiput.
BG: Light
BG: Dark
BG: Dark
(Grayish-cream
(Gray-black)
(Black)
or pinkish-cream)
S: Light
S: Dark brown
S: Light
(Light silvery
flecks)
(Same as on
body or stripe
broken into
spots)
blotches
BG: Light
(Or lighter
than dorsum
S: Light
(Few light shades)
BG: Dark
(Lead color)
S: ?
BG: Dark
Bulletin of the Maryland Herpetological Society
page 43
Volume 39 Number 3
September 2003
P altamontana
P aniiae
R fmcheni
P tttlkxM
P mured
P. CHIN-
BG: Dark
BG: Light
BG: Moderately
BG: Light
BG: Light
THROAT-
(Same as on
(Whitish)
light (At least
(Lighter than
(Same as on body
ED
venter)
S: Nearly
lighter than venter;
ventral body
ventral surfaces)
COLOR-
S: Light
immaculate
grayish)
surfaces)
S: Absent
ATION
(Creamy flecks,
S: Light (Same
S: Absent
not very
as on venter
conspicuous)
but more
numerous
Q,
BG: Light
BG: Light
BG: Dark
BG: Very dark
BG: Dark (Same as
LATERAL
(Cream-Iavendar)
(Light gray)
(Dark gray to
(Same as on body)
on body, at least
BODY
S: Immature
S: Absent
almost black;
S: Moderately
posteriorly)
COLOR-
“sparkling”
light (Same as on
S:?
ATION
pattern of minute
body, at least on
silver-white dots)
S: Dark (Few, if
any; dark as on
body)
dorsolateral areas)
R. LIMBS
BG: Dark (Same
BG: Moderately
BG: Moderately
BG: Very dark
BG: Dark
COLOR¬
as on body)
dark
dark (Grayish
(Same as on
(Same as on body)
ATION
S: Light (Creamy
(Brownish)
as on body)
body)
S: Light (As on
areas or spots on
S: Dark (Few)
S: Light (Large
S: Light (Large,
body; merging with
dorsal proximal
or one continuous
extensive only
those on dorsum)
parts; absent or
area; pinkish or
at front of femora
less conspicuous
reddish-brown to
and humera; absent
on distal areas)
cream; only on
on hands and feet;
back of femora
and humera
conspicuous)
conspicuous)
S.
Microendemic
Microendemic
Microendemic
Microendemic
Microendemic
GEOGRA¬
Central portion
Sierra Madre del
Southern tip of
Central Portion of
Southern tip of
PHICAL
of Eje
Sur
Sierra Madre
Eje Neovolcanico
Sierra Madre
DISTRI¬
NeovoScanico
OAX
Central
Transversal
Oriental in isolated
BUTION
Transversal
VER, PUE
DIF, MEX, MOR
Sierra de Juarez
MEX, MOR
OAX
T.ALTI-
TUDONAL
DISTRI¬
BUTION
(m asl)
3000 up
2100
2200-2600
2700-3500
2000-2500
Figure 14. Continued
page 44
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
September 2003
P. leprosa
R lonpeauda
Rmjgax
P. nmmamkta
R rohertsi
P. tkhcuibh
BG: Light
BG: Light
BG: Slightly light BG:?
BG: ?
BG: Dark
(Or at least
(Or at least
to slightly dark
S: With white
S: ?
S: Few tiny
lighter than
lighter than
(Light gray to
silverly flecks
lighter areas
ventral body
ventral body
dark gray)
Gular fold bears
surfaces)
S: ?
surfaces)
S:7
S: ?
a light edge
BG: Very dark
BG: Slightly light
BG: Moderately
BG: Dark
BG: Dark
BG: Dark
(Brown, slate or
to slightly dark
dark (Same as
S: Dark
(Brownish-
(Black)
black)
(Light gray to
on body)
(Black spots)
lavender)
S: Light
S: Light (A
medium gray)
S: Dark (As on
S: Light
metallic brown
“sparkling”
S: Light (A
body, costal
(Orange-brown;
pattern of minute
“sparkling”
grooves darker
few)
rounded, very
pattern of minute,
than folds)
abundant, silver-
rounded, very
white dots)
abundant, silver-
white dots)
BG: Dark (Same
BG: Dark (Same
BG:?
BG: Light or
BG:?
BG: Dark
as on body)
as on dorsum)
S: Light
dark
S:?
(black)
S: Light (Few, if
S: Light (Same
(Mottled with
(Depending on
S: Small dark
any; inconspicuous) as on body but
light pink or
body coloration)
brown blotches
small, few, and
light reddish-
S: Present
(Digits light
more or less
indistint)
brown spots)
brown)
Endemic
Microendemic
Microendemic
Microendemic
Microendemic
Microendemic
Central and
Central portion of
Southern tip of
Atlantic slope of
Central
Central
Eastern portions
Eja Neovolcanico
Sierra Madre
Central Sierra
portion of
highlands of
of Eja Neovolcanico Transversal in
Oriental, in the
Madre Oriental
Eja Neovolcanico
Sierra Madre
Tansversal
Sierra de
Sierra de Villa
VER
Transveral in
del Sur
DIF, MEX, MOR
Temascaltepec
Alta
Nevado de Toluca
GUE
TLA, PUE, VER
orAngangueo
OAX
MEX
MEX, MICH
2500-3600
2650-3000
2050
ca. 1000
3320-3600
2400-3000
Bulletin of the Maryland Herpetological Society
page 45
Volume 39 Number 3
September 2003
P. altamontana P. anitae
P. ftrscheni
P. tlilicxitl
P. iuarezi
U. HABI- Pine & fir ?
Oak, pine and
Pine, oak-pine,
Clouds and pine
TAT forests
cloud forests
and fir forests
forests
V. MICRO¬
HABITAT
Under logs
Under rocks
Epiphytic
bromeliads
Under logs, rocks,
or mounds of soil
Under bark of
fallen trees or rocks
W.
Terrestrial
Presumably
Scansorial
Terrestrial
Terrestrial
HABITS
(Semi-fossorial)
troglodytic
(Arboreal)
(Semi-fossorial)
(Semi-fossorial)
X.
SOURCES
AND
*Taylor, 1938 (2)
**Lynch et al,
1983 (not
Bogert, 1967 (3)
Shannon and
Werler, 1955 (4);
Regal, 1966 (2);
This study (14)
Regal, 1966 (25)
VOUCHER specified)
this study (4)
SPECI¬
MENS
(n)
Figure 14. Continued
page 46
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
P. leprosa
R loneicauda
R mvstax
Pine, pine-oak
Pine, pine-oak, fir,
Pine and oak
and fir forests
and cloud forests
forests
Under longs, bark
Under logs, wood
Under rocks
of logs and
chips, and bark
stumps, or rocks
of logs
Terrestrial
Terrestrial
Terrestrial
(Semi-fossorial)
(Semi-fossorial)
(Semi-fossorial)
Lynch etal. 1983
Lynch etal. 1983
Bogert, 1987 (2)
(107); this study
(90); this study
(44)
(5)
September 2003
P. nipromaculata
P. robertsi
P. tlahcuiloh
Tropical evergreen
Pine forests
Pine-oak and
and, maybe cloud
mixed forests
forests
(pine-oak, fir,
birch)
?
Under stones
Under logs and
under bark of
standing trees
Terrestrial
Terrestrial
Terrestrial
(Semi-fossorial)
(Semi-fossorial)
Taylor, 1941 (33);
Taylor, 1938 (21);
Adler, 1966
Regal, 1986(3);
Taylor, 1944;
(ID
Shannon and
**Lynch et al, 1983
Werler (not
specified)
(not specified)
Bulletin of the Maryland Herpetological Society
page 47
Volume 39 Number 3
September 2003
(another synonym of P. leprosa) has adpressed toes separated by 4 costal folds (as in
P. leprosa) while O. orizabensis types have the adpressed toes separated by 3 costal
folds (as in R leprosa) Even though P. tlilicxitl can be clearly set apart from the types
of junior synonym taxa of R leprosa based on most of the body ratios presented
previously, some differences among these types may lead us to think they are com¬
posite and represent more than one single taxon. The possibility of R tlilicxitl being
included in those types series cannot be totally ruled out until a close examination of
all type specimens be conducted (types are deposited in the collections of Kansas
University.) However this does not challenge or compromise the taxonomic validity
and distinctiveness of R tlilicxitl as described here.
In some morphological values, such as the low number of maxillary-premaxil¬
lary teeth, low number of vomerine teeth, big feet, and less webbing on toes, R
altamontana as referred by Taylor (1938), seems to show a close relationship to R
tlilicxitl. For some other characters the latter seems to have intermediate values be¬
tween R leprosa and R altamontana. R altamontana and R tlilicxitl differ, among
other characters, in several body ratios, vomerine teeth, and body coloration (see
figure 4.) Specimens of R altamontana in Mexican collections checked by myself
have proven to be either R tlilicxitl or R leprosa. In fact, it seems that there are no R
altamontana specimens in Mexican collections at all (!) In spite of extensive field
work in R altamontana type locality and presumable suitable habitat no specimens
have ever been found. Even the “rare” microsimpatric Crotalus transversus has been
collected several times in the area. All this makes me tend to think that R altamontana
is either extremely rare or that it doesn’t exist as a separate and distinctive taxon. US
alcoholic specimens in museums seem to be scarce too and their allocation some¬
times doubtful. Lynch, Wake and Yang (1983) reported R altamontana specimens
from “Zempoala district, 20 Km N of Tres Cumbres, Morelos”. This locality is con¬
fusing because Lagunas de Zempoala are in the state of Mexico, not Morelos, and the
precise locality (20 Km N of Tres Cumbres) would lie in Canada de Conteras formed
by Magdalena stream, in Distrito Federal. Nevertheless they pinpointed the locality
in their map on the southern slope of Sierra del Ajusco in the state of Morelos. For
their locality 7 (in the text) they referred R leprosa (“Zempoala district; 10 Km N of
Tres Cumbres, Morelos) but in the map they showed R altamontana. Later on, Lynch
et al (ibid.) suggested that (I quote): “Because R altamontana and R robertsi are so
similar genetically and are allopatrically distributed, a case should be made
synonymyzing them. However, because the species are distinct in color and body
proportions we continue to recognize them as separate taxa to the present” (end of
quote). So far I personally cannot say what R altamontana really is, but R robertsi is
as different from R altamontana original description as any other Pseudoeurycea spe-
page 48
Bulletin of the Maryland Herpetological Society
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September 2003
cies is. Genetic similarity by itself doesn’t prove conspecifity, let alone allopatry of
populations. I am absolutely positive that nothing similar to P. robertsi occurs in the
P. altamontana geographical range. I also can say that P. tlilicxitl and R robertsi are
not the same, based on morphometric, meristic, coloration and ecological data (see
figure 14.) But the real point here would be to show the controversial specific status
of P. altamontana, as well as its geographical distribution.
Bogert ( 1 967) related the character of long legs to scansorial habits. According
to his statement, we could think of P. altamontana and P. tlilicxitl as scansorial forms.
But the fact that all specimens of both P. altamontana and P. tlilicxitl have been col¬
lected on the ground (under logs) and the lack or scarcity of epiphytes in their habitat
makes this view extremely unlikely, even though that it would explain the reduced
number of specimens collected because of being looked for in the wrong microhabi¬
tat. The alternative explanation of possible extinction doesn’t seem to be very likely
but cannot be disregarded as impossible either. Habitat alteration has occurred in the
type locality area that has affected drastically another microsimpatric salamander
populations: Rhyacosiredon zempoalensis. Being microendemic, this species has been
driven to the verge of extinction. Another possible explanation would be an extreme
rareness of P. altamontana due to a very narrow range of tolerance to environmental
parameters as it is the case of P. tlilicxitl, but even with narrower ranges (being a truly
stenoecious specialist.)
Until a detailed analysis of the 2 known specimens of the type series of P.
altamontana is made (they are at the University of Kansas) I recommend that the
specific status of this salamander be considered as doubtful. Because of the herein
proven distinctiveness of P. tlilicxitl from P. altamontana, this stand would neither
compromise nor challenge the validity of P. tlilicxitl.
A final comment should be made about a recently described new Pseudoeurycea
from Guerrero: P. tlahcuiloh (Adler, 1996.) The author says that it superficially re¬
sembles R leprosa in general proportions and color pattern. Besides the differences
shown in figure 14 between P. tlahcuiloh and all other leprosa group species, Adler
adds that R tlahcuiloh has thinner and longer tail than P. leprosa, a larger head and a
more pronounced neck. P. tlahcuiloh differs from P. tlilicxitl at least in having shorter
legs, more premaxillary-maxillary teeth, more vomerine teeth, longer tail, and differ¬
ent coloration (dorsal, ventral, lateral tail and limbs). I wanted to include P. tlahcuiloh
within the leprosa group only because of its alleged resemblance to P. leprosa. Adler
(op. cit.) didn’t want to venture to allocate any of his 5 new species to any
Pseudoeurycea group. The real kinship between R tlahcuiloh and other Pseudoeurycea
species is still unknown.
Bulletin of the Maryland Herpetological Society
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Volume 39 Number 3
September 2003
LITERATURE CITED
Adler, Kraig.
1 996. The salamanders of Guerrero, Mexico, with descriptions of 5 new
species of Pseudoeurycea (Caudata: Plethodontidae.) Univ. Kan¬
sas Nat Hist Mus. Occ. Pap. 177: 1-28, 5 figs.
Baird, Irwin L.
1951 An anatomical study of certain salamanders of the genus
Pseudoeurycea. Univ. Kansas Sci. Bull. 34(6): 221-265, pi XXVI-
XXVIII.
Blatchley,
1893. On a collection of batrachians and reptiles from Mount Orizaba,
Mexico, with descriptions of two new species. Proc. U. S. Nat.
Mus., 26: p. 38.
Bogert, Charles M.
1 967. New salamanders of the plethodontid genus Pseudoeurycea from
Sierra Madre del Sur, Mexico. Amer. Mus. Novit, 2314: 1-27,
figs. 1-8.
Brame, A. H. Jr.
1968. Systematics and evolution of the Mesoamerican salamander ge¬
nus Oedipina. I. Herpetol. 2: 2-64.
Brocchi, P.
1883. Etudes des batraciens de FAmerique Centrale. In Mision
scientifique au Mexique et dans FAmerique Centrale. Paris.
Imprimerie Nationale, pt. 3, sect. 2, livr. 3, pp. 109-110, planche
xvm-xix.
Cope, Edward D.
1 869. A review of the species of the Plethodontidae and Desmognathidae.
Proc. Acad. Nat. Sci. Philadelphia, 93-118.
Dunn, E. R
1926. The salamanders of the family Plethodontidae. Smith College,
Nort-hampton, Mass., vii + 441 p.
page 50
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
September 2003
Elias, P.
1984. Salamanders of the northwestern highlands of Guatemala. Nat.
Hist. Mus. Los Angeles Co. Contrib. Sci. 348: 1-20.
Hanken, I.
1983. Genetic variation in a dwarfed lineage, the Mexican salamander
genus Thorius (Amphibia: Plethodontidae): taxonomic, ecologic
and evolutionary implications. Copeia 1983: 1051-1073.
Lara and Ortega
1 980 Diferential spatial resources exploitation by 4 sympatric species
of salamanders, (unpublished) Paper presented at IV Congreso
Nacional de Zoologia, Ensenada, B.C.
Lynch James F., David B. Wake and Suh Y. Yang.
1983. Genic and morphological differentiation in Mexican
Pseudoeurycea (Caudata: Plethodontidae), with a description of
a new species. Copeia (4): 884-894.
Maxson, Linda R. and Richard D. Maxson.
1 979. Comparative albumin and biochemical evolution in plethodontid
salamanders. Evolution 33(4): 1057-1062.
— and David B. Wake.
1981. Albumin evolution and its phyllogenetic implications in the
plethodontid salamander genera Pseudoeurycea and
Chiroptero triton. Herpetologica 37(2): 109-117.
Papenfuss, T. J., David B. Wake and Kraig Adler.
1 983. Salamanders of the genus Bolitoglossa from Sierra Madre del Sur
of southern Mexico. J. Herpet. 17:295-307.
Shannon, Frederick A. & John E. Werler.
1955. Report on a Small collection of amphibians from Veracruz, with a
description of a new species of Pseudoeurycea. Herpetologica
11(2): 82-84, figs. 1-2.
Regal, Philip, J.
1 966. A new plethodontid salamander from Oaxaca, Mexico. Amer. Mus.
Novit. 2266: 1-8, figs. 1-3.
Bulletin of the Maryland Herpetological Society
page 51
Volume 39 Number 3
September 2003
Smith, Hobart M. and Rozella B. Smith.
1976. Synopsis of the herpetofauna of Mexico, Vol IV: Source analysis
and index for Mexican amphibians. John Johnson, North
Bennington, Vermont.
Taylor, Edward H.
1 938. Concerning Mexican salamanders. Univ. Kansas Sci. Bull. 25 ( 14):
259-313, pl.XXV-XXIX.
1941. New amphibians from the Hobart M. Smith Mexican collections.
Univ. Kansas Sci. Bull. 27(8): 141-145, fig. 1.
1944. The genera of plethodontid salamanders in Mexico. Pt. I. Univ.
Kansas Sci. Bull. 30, pt. (12): 189-231.
1 945 . Summary of the collections of amphibians made under the Walter
Rathbone Bacon traveling scholarship. Proc. U. S. Nat. Mus. ,
95(3185): 521-545.
Wake, David B. and Paul Elias.
1983. New genera and new species of Central American salamanders,
with a review of the tropical genera (Amphibia, Caudata,
Plethodontidae.) Contrib. Sci., 345: 1-19.
— and James R Lynch.
1 976. The distribution, ecology and evolutionary history of plethodontid
salamanders in tropical America. Nat. Hist. Mus. LosAngeles
County Sci. Bull., 25: 1-65
Guillermo Lara-Gongora
Manantiales de Agua Zarca 389-A, Campeste de Los Manantiales
Morelia , Michoacdn , Mexico
grammicus @ hotmail. com
Received
Accepted
page 52
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
September 2003
Knobloch’s King Snake (Lampropeltis pyromelana
knoblochi) of Mexico a Species
Julio A. Lemos-Espinal, David Chiszar and Hobart M. Smith
Abstract.
Comparison of three Lampropeltis p. pyromelana with one L. p. knoblochi , all
from Chihuahua, Mexico, indicates that the latter taxon is of species rank.
A multi-redspotted king snake from western Chihuahua was described as
Lampropeltis knoblochi by Taylor (1940). It continued to be so recognized at least
until 1953 (Maldonado-Koerdell), although Bogert and Oliver (1945) suggested that
it might be a subspecies of Lampropeltis pyromelana (Cope, 1867). The name last
appeared at specific rank in Marx (1976), but that was in a list of type specimens,
using the original name. In 1953 Tanner formally placed the taxon as a subspecies of
L. pyromelana, and it has been so designated ever since.
The type locality was Mojarachic, Chihuahua, and Tanner (1985) noted speci¬
mens subsequently taken at Yoquiro and 25 mi S Creel, Chihuahua, and Yecora,
Sonora. Campbell and Lamar (1989) recorded it from above Cascada de Baseachic,
Rio Durazno. Probably far more specimens have been taken for herpetohusbandry
than have found their way into museums, to judge by the numerous (at least 21 to our
knowledge) references on the subject (e.g., Assetto, 1984; Grumbeck, 1991; Markel,
1978, 1990; Mara, 1994; Martin, 1975). However, it is likely that some references
are in error. Markel (1990), for instance, has a painting that is partially incorrect (the
lateral blotches and sublateral light lines are not depicted, and the color is shown as
red, rather than orange), and the two photographs supposedly of L. knoblochi are of
L. pyromelana. The geographic ranges shown for the two species is also incorrect.
Mara (1994) likewise illustrates L. knoblochi with a specimen actually of L.
pyromelana. Much of the difficulty in such misidentifications is the absence of accu¬
rate locality data.
We here report another voucher specimen, UBIPRO 10575, taken by JALE at
Arroyo del Agua, near Maguarichi (27°54'45.6"N, 107°58,49.2,,W), at 2083 m, 25
September 2002. It is an adult female with the typical spotted pattern of the taxon
(Fig. 1), measuring 776 mm TTL, TL 124 mm. The ventrals are 228, anal entire,
subcaudals 66 (missing a few terminal scales), scale rows 23-25-19. There are 7-7
supralabials, 8-9 infralabials, 1-1 preoculars, 1-1 loreal, 2-2 postoculars, temporals
1-1 although a second one is narrowly separated from the postoculars.
Bulletin of the Maryland Herpetological Society
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Volume 39 Number 3
September 2003
Figure 1. Dorsal view of Lampropeltis knoblochi, UBIPRO 10575.
Figure 2. Ventral view of same specimen as in Fig. 1.
page 54
Bulletin of the Maryland Herpetological Society
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September 2003
Dorsal orange spots on body 38 (or 48 if fused ones are counted separately);
15 spots on tail, all but the terminal three orange-centered; spots on body black-
bordered, 8-11 scale rows wide, 2-5 (usually 3-4) scales long, bordered laterally by
an irregular white line 1 scale wide, and separated by white interspaces 1 scale long.
A series of rounded, orange, extensively fused, black-bordered spots, 2-3 scales
in diameter, lateral to lateral light line; an irregular sublateral light line ventral to the
lateral spots, mostly along the 1st and 2nd scale rows. Ends of more or less alternate
ventrals orange; scattered, more or less square black spots, about one ventral in length,
scattered on venter (Fig. 2).
Snout light tan; a roughly doughnut-shaped, black-bordered orange spot on
head, frontal to parietals, extending to lower eye level, and surrounding a light cen¬
tral spot on posterior tip of frontal; a light tan ring, the first of the interspaces between
the spots on body, crossing the posterior ends of the parietals; anterior black border
of 1st dorsal spot extending laterally no farther than the level of the 7th supralabial,
and posterior to it; no markings on either supralabials or infralabials.
This specimen is sharply different from L. p. pyromelana, of which three adults
were collected by JALE and Matthew J. Ingrassi in the summer of 2002. They are
UBIPRO 10172 from km 38 on the Chihuahua-Namiquipa highway (29°4'38.3"N,
106°36'30.4''W), 2300 m, 9 August; UBIPRO 10190, km 17.1, Chihuahua-Namiquipa
highway (29°5'28.7HN, 106°28’51.4"W), 1743m, 9 August; UBIPRO 10955, middle
of Canon del Oso (31°16T7.7"N, 108°437.0"W), 1661m, 18 September. The head
scales in these specimens are the same as in L. p. knoblochi, except that the anterior
temporals are 2-2 in two, 1-1 in one. The ventrals are 217-225, subcaudals 70-72 (all
males); scale rows 23-23-19 in two, 23-23-17 in one. The dorsal spots are bright red,
black-bordered and open-sided, the red portion extending across the abdomen or
well onto the sides of the ventrals. The light interspaces between the dorsal spots
number 38, 43, 43, on tail 9, 10, and 16 (counting the anal interspaces). The black,
spots on the ventrals tend to be more rectangular than square, in numerous cases
extending across much or all of the scales. The dorsal blotch on head is jet black, not
light-centered; there are one or two black streaks on infralabial sutures; the anterior
black border of the first body ring reaches to or ventral to the 7th supralabial, border¬
ing or crossing it. The largest specimen measured 963 mm TTL.
Although no differences in scutellation of these two subspecies are evident,
there are numerous differences in color and pattern. L. p. knoblochi differs from L. p.
pyromelana in having orange (vs red) markings on body; dorsal markings spotlike
(vs ringlike), a lateral light line (vs none); a row of small, orange, black-bordered
spots below the lateral light line (vs none); a sublateral light line below the lateral
Bulletin of the Maryland Herpetological Society
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Volume 39 Number 3
September 2003
spots (vs none); orange on ends of ventral scales independent of more dorsal spots (vs
connected, and more extensive); and dorsal blotch on head doughnut shaped, orange-
centered, enclosing a tan central dot (vs wholly jet black).
These distinctions appear to be categorical. Tanner (1953) regarded two speci¬
mens, from northern Sonora and southeastern Chihuahua as intergrades, but they are
far from the range of L. p. knoblochi, and later were apparently assumed to be vari¬
ants of L. p. pyromelana by Tanner (1983, 1985).
Furthermore, the range of L. p. knoblochi appears to be dichopatric relative to
the range of the other subspecies. It is limited to the spectacularly deep canyons on
Pacific slopes of extreme southwestern Chihuahua, whereas L. p. pyromelana is lim¬
ited to the Atlantic slopes of the Sierra Madre, except perhaps in the extreme north.
For these reason, we revive the rank originally assigned to L. knoblochi (Tay¬
lor, 1940). It is obviously related to L. pyromelana , but appears to be on an indepen¬
dent evolutionary tangent.
Acknowledgments.
We are much indebted to Dr. E. A. Liner for help with the literature, to Mat¬
thew J. Ingrassi for field assistance and to Dr. Ned Friedman and William Gallup for
the photographs. CONAB 10 kindly provided support for field work by JLE under
projects U003, X004 and AE003.
Literature Cited.
Assetto, R.O., Jr.
1984 Captive reproduction in Lampropeltis pyromelana. Bull. Phila¬
delphia Herp. Soc. 30: 25-26.
Bogert, C.M. and J.A. Oliver.
1945. A preliminary analysis of the herpetofauna of Sonora. Bull. Am.
Mus. Nat. Hist. 83: 297-426.
Campbell, J.A. and W. W. Lamar.
1989. The venomous reptiles of Latin America. Ithaca, New York,
Cornell Univ. xiii, 425 pp.
Cope, E.D.
1 867. Fifth contribution to the herpetology of tropical America. Proc.
Acad. Nat. Sci. Philadelphia: 18: 300-314 (1866).
page 56
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
September 2003
Grumbeck, S.
1991. Mexican mountain kingsnake (Lampropeltis pyromelana
knohlochi). North Texas Herp Soc. Newsl. 1991 (Febr.): 1, 5.
Maldonado-Koerdell, M.
1953. [Amphibians and reptiles], pp. 107-133 in Beltran, E. (ed.), Vida
silvestre y recursos naturales a lo largo de la carretera
panamericana. Mexico, D. F. Inst. Mexicano de Recursos Natu¬
rales Renovables, A. C. [v], 354 pp.
Mara, W.R
1994.
Milk snakes: every species and sub-species in brilliant color. Nep¬
tune City, New Jersey, T.F.H. 63 pp.
Markel, R. G.
1978.
Keeping kingsnakes in captivity, with a checklist of the genus
Lampropeltis. Herpetology 9(4): 1-5.
1990.
Kingsnakes and milksnakes. Neptune City, New Jersey, T.F.H.
144 pp.
Martin, B.E.
1975.
Notes on the Sonora mountain kingsnake. Herp 11(3-4): 9-15.
Marx, H.
1976.
Supplementary catalogue of reptiles and amphibians in Field
Museum of Natural History. Fieldiana Zool. 69: 33-94.
Tanner. W.W.
1953.
A study of taxonomy and phylogeny of Lampropeltis pyromelana
Cope. Gt. Basin Nat. 13: 47-66.
1983.
Lampropeltis pyromelana. Cat. Am. Amph. Rept. (342): 1-2.
1985.
Snakes of western Chihuahua. Gr. Basin Nat. 45: 615-676.
Taylor, E.H.
1940.
A new Lampropeltis from western Mexico. Copeia 1 940: 253-
255.
Bulletin of the Maryland Herpetological Society page 57
Volume 39 Number 3
September 2003
JALE: Laboratorio de Ecologia, UBIPRO, Facultad de Estudios Superiores
Iztacala, Tlalnepantla, Estado de Mexico, 54090 Mexico.
DC and HMS: University of Colorado Museum, Boulder, Colorado 80309-0334.
Received: 15 May 2003
Accepted: 27 May 2003
page 58
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
September 2003
Disltributional and Variational Data on the
Frogs of the Genus Rana in Chihuahua, Mexico,
Including a New Species
Hobart M. Smith and David Chiszar
Abstract.
Seven species of Rana are known to occur in Chihuahua: R. berlandieri, R.
catesbeiana, R. chiricahuensis, R. forreri, R. magnaocularis, R. lemosespinali sp.
nov., and R. tarahumarae. Locality and variational data on these species are recorded
from material of the genus recently collected in the state by Julio A. Lemos-Espinal.
Material of the genus Rana collected in recent years by Julio A. Lemos-Espinal
(JLE) include representatives of all but one of the known species of that genus in the
state, as well as a new species. Material reported here for the first time is in the
collection of Unidad de Biologia, Tecnologia y Prototipos (UBIPRO), Faultad de
Estudios Superiores Iztacala, Tlalnepantla, Mexico. All catalog numbers refer to that
collection.
Rana berlandieri Baird
Current knowledge of distribution of this species in Chihuahua, as represented
by Conant and Collins (1998) indicates restriction to the eastern half of the state. The
present material confirms its existence also in the southwestern mountains. Speci¬
mens at hand include 6962-70, Ojo Rancho del Cuervo (30° 14'29.5"N,
105°19'52.1"W), 1134m; 6676, 6825, Ejido Mesa El Zorrillito, Guadalupe y Calvo
(26°3'34.8"N, 106°57'28.8"W), 2595m; 8079, Rio Papigochi at La Junta, mpio
Guerrero (28°2754.0HN, 107°19,39.4,,W), 2103m; 8525, Basigochi de Aboreachi,
mpio Guachochi (27°5'26.3"N, 107°14’40.3"W), 2400m; 8531-3, 8537-40, Guazarare,
mpio Guachochi (27°3’47.2"N, 107°12'0.5"W), 2300m; 8025, river 1 km WBalneario
Division del Norte, mpio Jimenez (25°53,32.8"N, 104°22T7.8"W), 1360m; 9133-4,
km 209.5 Creel-Guachochi (27°5,23.1,,N, 107°14,59,,W), 2350 m. Webb and Baker
( 1 984) reported (as ((Rana pipiens group ”) what is presumably the same as the west¬
ern Chihuahua R. berlandieri here reported, from 7.7 rd mi SW El Vergel, 3002m,
and 15 rd mi NE Atascaderos, 2900 m, in Chihuahua; others were reported from
adjacent Durango.
The western samples probably represent a taxon different from that of the east¬
ern sample; the two populations appear to be dichopatric. The eastern samples al-
Bulletin of the Maryland Herpetological Society
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Volume 39 Number 3
September 2003
ways have considerable dark pigmentation on the lower jaw, at least at the sides, and
the rear surfaces of the thighs are boldly reticulated, black on white. In the western
samples, with one exception, there is little or no pigmentation on the lower jaw, and
the rear surfaces of the thighs are not at all, or but weakly, reticulated. The exception
is a transformling 28mm SVL from La Junta. Nos. 9133-4 are partially transformed
tadpoles 78-82 mm TTL, tail 50-52 mm, hind leg length 29-39 mm, no forelegs. The
tail is mottled black.
In all samples, the dorsolateral folds often are not noticeably displaced poste¬
riorly. No. 6825 is a male with external vocal sacs, associated with a clutch of eggs.
Rana catesbeiana Shaw
Lemos-Espinal et ai (2000) reported this species (UBIPRO 2169) from Ejido
Ojo de Enmedio, at the base of the Sierra de Samalayuca.
Rana chiricahuensis Platz and Mecham
The range of this species slightly overlaps that of R . berlandieri: 8495-7,
Basigochi de Aboreachi, mpio Guachochi (27°12T2.2"N, 107°22'45.0HW), 2409 m;
10380, km 95, Flores Magon-Buenaventura (29°54T2.9"N, 107° 17*49. 1"W), 1591
m; 8854-5, tadpoles, Pacheco (30°5*L7,,N, 108°20'29.4HW). It is our opinion that the
uRana sinaloae ” (-liana pustulosa) of Dominguez et ai (1977) is actually Rana
chiricahuensis ; it is from 13 km W Matachic; the latter is at 28°51'N, 107°45'W.
The series of four adults varies in SVL 45-52 mm. The most striking character¬
istic shared by all four, and differing from all other species of Rana in the state, is the
nearly or quite uniformly dark posterior surfaces of the thighs, except for a tiny white
dot capping each of the numerous tubercles.
The dorsolateral folds are bright white in two, dull-colored in two; they are not
noticeably displaced posteriorly. The upper lip is prominently reticulated black and
white in three, only marginally in one. The prominent subtympanic white stripe ends
anteriorly below the eye. There are 11-17 dark spots between the dorsolateral folds,
one on each eyelid. In only one is there a dark spot anterior to the anterior border of
the eyes.
The two tadpoles are unusual in having the musculature of the tail and dorsum
of the body prominently evident, whitish in color. The whitish epaxial muscles ex¬
tend anteriorly almost to the eyes, and sharply contract with the dark skin elsewhere.
page 60
Bulletin of the Maryland Herpetological Society
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September 2003
Ranaforreri Boulenger
This species was reported for the first time from Chihuahua by Lemos-Espinal
et al (2002), UBIPRO 5923, at Arroyo El Camuchil, Batopilas (27°01'34.rN,
107°45,44.5"W), 435 m, 17 July 2000. The present collection includes UBIPRO 9025
(tadpoles) and 9398 from the same locality, coordinates and altitude as the preceding;
and 10605-6, Arroyo Las Borregas, Chmipas (17023'4.3"N, 108o32,21.1"W), 470 m.
The transformed specimens available are 59, 48 and 27 mm SVL; the posterior
surfaces of the thighs are very dimly or not retiuclated; the dorsolateral folds are dull-
colored, embrace 10 or 11 large dark spots, and are not noticeably displaced posteri¬
orly.
There are about 35 tadpoles under no. 9025; their assignment to R. forreri is
conjectural. The breeding seasons of the sympatric R. forreri and R. magnaocularis
are thought to be different (Frost and Bagnara, 1977); and the specimens obtained of
the latter species are all transformed, whereas the tadpoles available are quite small.
Regardless, the tadpoles (about 35) are unusual in their small size (11.5 mm
maximum body length), opaquely black body, and the presence of 7 mm hind legs on
the largest specimens. The tail fin at midtail is little more than half the diameter of the
tail musculature at that point. There are two equally long rows of denticles in the
upper labium, but the inner row is widely split about half the length of either end. The
lower labium has three complete rows of equal length of denticles,
Rana magnaocularis Frost and Bagnara
This species is partially sympatric with R. ferreri , and appears to be much
more abundant. It has not been reported before from Chihuahua, although known
localities of occurrence are very close. Available are 9244, 9427-34, Arroyo El
Camuchil, Batopilas (27°01'34.rN, 107°45,44.5,,W), 435 m; 9069-70, 9294-9316,
9336-8, 9440, Batopilas, same coordinates and altitude as the preceding; 10607, Ar¬
royo Las Borregas, Chmipas (17°2314.3”N, 108°32'21.1MW), 470 m.
All specimens available are juveniles; the maximum SVL is 46 mm, and the
smallest is a transformling with a tail remnant 7 mm in length, and the SVL is 26 mm.
The dorsolateral folds are white except in the smallest specimens, in which they are
dull-colored. The posterior surfaces of the thighs are brightly mottled. Dorsal dark
spots are not evident in the transformling; apparently further development is required
before their appearance is triggered, because fully transformed individuals of the
same size as the transformling have well-developed spots.
Bulletin of the Maryland Herpetological Society
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September 2003
The dark spots between the dorsolateral folds, anterior to the end of the wo
style but excluding the head, vary 11-21 (M=16.7). The dorsal cranial dark spots vary
considerably, but never number more than three (only one has that number). Occa¬
sionally (20% of 24 in which spots are reliably discerned) there are no spots on top of
the head at all
The displacement of the dorsolateral folds posteriorly is regularly evident.
Rana lemosespinali sp. nov .
Holotype. LE-UBIPRO 8236, adult male, between Creel and San Rafael,
mnicipio Urique (27°31T8.2"N, 107°50,50.5,’W), 2313 m, 8 September 2001, Julio
A, Lemos-Espinal collector. Paratypes. Seven topotypes, LE-UBIPRO 8233-5, 8237-
40. Referred material . LE-UBIPRO 9495, 1 km N Humira (27°25'43"N,
107°29'24.6"W), 1900 m, 13 July 2002.
Diagnosis . A member of the Rana pipiens group having 25-46 sharply out¬
lined, white-bordered dark spots between the dorsolateral folds; dorsolateral fold
prominent and continuous; dorsum weakly ridged as well as pustular; sides of body
strongly granular; venter unpigmented.
Description of holotype. Male, 54mm SVL; 46 light-bordered dark spots be¬
tween dorsolateral folds from behind eyes to near the end of the urostyle, tending to
be elongate; a weak longitudinal ridge in the center of some spots; dorsal skin some¬
what pustular, not smooth; dorsolateral fold light tan, not white, continuous to rear
level of urostyle, curving slightly medially over sacrum; sides of body strongly granu¬
lar, upper parts with numerous dark spots like those on dorsum, lower parts whitish,
black-reticulated; venter white, unpigmented, granular posteriorly.
Two small dark spots on each eyelid, two between and two on snout; a bright
white stripe from axillary level to below eye, indistinguishable anteriorly; upper lip
black-reticulated on white, trending longitudinally; a few black spots posteriorly on
lower lip.
Posterior surfaces of thighs granular on lower third, smooth on upper part,
reticulated black on white; dark crossbars on hind legs distinctly narrower than spaces
between.
Variation. The paratypes are remarkably similar to the holotype. They vary 53-
56 mm SVL; 1-2 dark spots on snout, anterior to orbits; 2-4 small supraorbital spots
(total both sides); 1-3 spots between orbits; interdorsolateral fold spots 28-46 (M=37).
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Bulletin of the Maryland Herpetological Society
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September 2003
The referred specimen from near Humira is somewhat different from the type
series, having but 25 interdorsolateral fold spots, and the dark crossbars on hind legs
equal in width or wider than their interspaces. The locality is about 40 km east of the
type locality, and both are on Pacific slopes.
Comparisons. All of the other six species of Rana known from Chihuahua are
readily distinguished from R. lemosespinali by having fewer than 25, or no, dorsal
spots. Other conspicuous differences occur in R. catesbeiana and R. tarahumarae,
both lacking well-defined dorsolateral folds (vs present) and no spotting about (vs
spotted); R. chiricahuensis has a uniformly dark, white dotted posterior surface of
thigh (vs reticulated) and a dark-pigmented ventral surface (vs white); R. berlandieri ,
R. forreri and R. magnaocularis all have smooth sides of the body (vs granular).
R. chiricahuensis appears to be most similar to R. lemosespinali , sharing size,
granular skin, weak ridges on the dorsum, etc. The two may be dichopatric in Chi¬
huahua, the latter to the west of the Continental Divide, the former to the east.
Etymology. The species is named for the collector, Dr. Julio A. Lemos-Espinal,
who has studied and sampled the herpetofauna of the previously rather neglected
state of Chihuahua far more than anyone else.
Rana tarahumarae Boulenger
Although this species is well known from the same area in which R.
lemosespinali was taken (Tanner, 1989; Zweifel, 1968; other records in Dominguez
et al., 1977; Webb and Baker, 1984; Webb, 2000), JLE has not found it, perhaps
because of its specialized, highly aquatic habitat. It is likewise surprising that no
others have collected R. lemosespinali where JLE found it.
This species is almost twice the size of R. chiricahuensis and R. lemosespinali,
and is readily distinguished from all other species of Rana in Chihuahua, except R.
catesbeiana, by the absence of or very weak dorsolateral folds. The latter species is
readily distinguished from the former by its larger, smooth-surfaced tympanum, nearly
as long as eye or longer, and its very large size, maximum 203 mm (vs a tubercular
tympanum about half as long as eye, and a SVL not exceeding about 102 mm). R.
catesbeiana does not occur in the mountainous habitat of the R. tarahumarae.
The records for R. tarahumarae in Chihuahua and adjacent Sonora are all
in the southern corners of the states, widely isolated from the records in the
United States and northern Sonora. Electrophoretic comparisons would be of consid¬
erable interest.
Bulletin of the Maryland Herpetological Society
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Volume 39 Number 3
September 2003
Key to the Species of Rana In Chihuahua
Because of the frequently encountered difficulty of determining whether the
posterior ends of the dorsolateral folds are displaced or not, despite frequent use in
keys, we here mostly resort to other characters that we regard are more readily deter¬
mined.
1 A. Dorsolateral folds absent or dimly evident ........................................... 2
B. Dorsolateral folds clearly evident ......................................................... 3
2A. Diameter of tympanum about half that of eye; SVL not exceeding
~102MM ............................................................................ tarahumarae
B. Diameter of tympanum about equal to eye diameter, or (in males) much
greater; SVL to 203 mm ...................................................... catesbeiana
3 A. Posterior surfaces of thighs more or less uniformly dark except
for a tiny white dot on each of the numerous, small tubercles ...............
......................................................................................... chiricahuensis
B. Posterior surfaces of thighs patterned differently ................................. 4
4 A. Dark spots between dorsolateral folds 25 or more; sides of body
strongly granular ............................................................... lemosespinali
B. Dark spots between dorsolateral folds 23 or fewer; sides of body
smooth .................................................................................................. 5
5A. Southwestern comer of the state, at altitudes less than 500m .............. 6
B. East of the continental divide, at altitudes greater than 1000m ..............
.............................................................................................. berlandieri
6A. Posterior surfaces of thighs not or very dimly reticulated; dark
dark spots between dorsolateral folds fewer (10-11 in available
material); dorsolateral folds not displaced posteriorly ................ forreri
B. Posterior surfaces of thighs strongly reticulated; dark spots between
dorsolateral folds more numerous (11-21, M=16.7); dorsolateral folds
usually clearly displaced posteriorly .............................. magnapcularis
Acknowledgments.
We are grateful to Robert G. Webb for critical counsel.
page 64
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
September 2003
Literature Cited.
Conant, R. and J. T. Collins.
1998. Reptiles and amphibians: eastem/central North America. New
York, Houghton Mifflin, xviv, 616 pp.
Dominguez, P., T. Alvarez and P. Huerta.
1 977. Coleccion de anfibios y reptiles del noroeste de Chihuahua Mexico.
Revta. Soc. Mex. Hist. Nat. 35: 117-142 (1974).
Frost, J.S. and J. T. Bagnara.
1977. An analysis of reproductive isolation between Rana
magnaocularis and Rana herlandieriforreri (Rana pipiens com¬
plex). J. Exp. Zool. 202: 291-305.
Lemos-Espinal, J., D. L. Auth, D. Chiszar and H. M. Smith
2002. Geographic distribution: Rana forreri. Herp. Rev. 33:63.
_ _ _ _ , H. M. Smith and D. Chiszar.
Distributional records of anurans in Chihuahua, Mexico. Bull.
Chicago Herp. Soc. 35: 162-163.
Tanner, W.W.
1989. Amphibians of western Chihuahua. Great Basin Naturalist 49:
38-70.
Webb, R.G.
2001 . Frogs of the tarahumarae group in western Mexico. Pp. 20-43 in
Johnson, J.D., R.G. Webb and O.A. Flores-Villela (eds.),
Mesoamerican herpetology: systematics, zoogeography, and con¬
servation. Univ. Texas El Paso, Spec. Publ. (1.) iv, 200 pp.
_ _ _ and R. H. Baker.
1 984. Terrestrial vertebrates of the Cerro Mohinora region, Chihuahua,
Mexico. Southw. Nat. 29:243-246.
Zweifel, R.G.
1968. Rana tarahumarae Boulenger. Cat. Am. Amph. Rept. (66:) 1-2.
Bulletin of the Maryland Herpetological Society
page 65
Volume 39 Number 3
September 2003
Received
Accepted
HMS, DC: Museum of Natural History, University of Colorado ,
Boulder, CO 80309-0334.
9 May 2003
30 May 2003
page 66
Bulletin of the Maryland Herpetological Society
Volume 39 Number 3 September 2003
page 68 Bulletin of the Maryland Herpetological Society
Volume 39 Number 3
News and Notes
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News and Notes
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Bulletin of the Maryland Herpetological Society
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US ISSN: 0025-4231
(QL
bH-0
8
BULLETIN Or THE
'/flacylanb
fSecpetologkal
0odety
DEPARTMENT OF HERPETOLOGY
THE NATURAL HISTORY SOCIETY OF MARYLAND, INC.
MDHS . A Founder Member of the Eastern
Seaboard Herpetological League
31 DECEMBER 2003
VOLUME 39 NUMBER 4
BULLETIN OF THE MARYLAND HERPETOLOGICAL SOCIETY
Volume 39 Number 4 December 2003
CONTENTS
Adelphicos quadrivirgatum (Serpentes: Colubridae) in Hidalgo, Mexico, with
Comments on its Relationships to A. Visoninum
Fernando Mendoza Quijano, Jose Ismael Campos Rodriguez,
Juan Carlos Lopez Vidal, Hobart M. Smith and
David Chiszar . . . . . . . . 77
Analysis of Phenotypic Variation in the Lizard Sceloporus cautus and Adjacent
Populations of Related Species
George M. Ferguson, Hobart M. Smith and David Chiszar . . 85
2001-2002 Anurans, Exclusive of Rana, from Durango and Chihuahua, Mexico
Julio A. Lemos-Espinal, Hobart M. Smith and
David Chiszar . . . . . 92
The Madrean Alligator Lizard of the Sierra del Nido, Chihuahua, Mexico (Reptilia:
Sauria: Anguidae)
Hobart M. Smith, Julio A. Lemos-Espinal, David Chiszar and
Matthew J. Ingrasci . .99
Book Review: Amphibians and Reptiles of Delmarva
Harlan D. Walley . . . . . . . . . . 103
Book Review: Introduction to Horned Lizards of North America
Harlan D. Walley . . . . 105
Errata . . . . . . . . . . . . . 107
BULLETIN OF THE
mbt)8
Volume 39 Number 4 December 2003
The Maryland Herpetological Society
Department of Herpetology, Natural History Society of Maryland, Inc.
President Tim Hoen
Executive Editor Herbert S. Harris, Jr.
Steering Committee
Frank B. Groves Jerry D. Hardy, Jr.
Herbert S. Harris, Jr. Tim Hoen
Library of Congress Catalog Card Number: 76-93458
Membership Rates
Membership in the Maryland Herpetological Society is $25.00 per year
and includes the Bulletin of the Maryland Herpetological Society. For¬
eign is $35.00 per year. Make all checks payable to the Natural History
Society of Maryland, Inc.
Meetings
Meetings are held monthly and will be announced in the "Herp Talk"
newsletter and on the website, www.naturalhistory.org.
Volume 39 Number 4
December 2003
Adelphicos quadrivirgatum (Serpentes: Colubridae)
in Hidalgo, Mexico, with Comments on its
Relationships to A. visoninum
Fernando Mendoza Quijano, Jose Ismael Campos Rodriguez, Juan Carlos Lopez '
Vidal , Hobart M. Smith and David Chiszar.
Abstract.
Adelphicos q. quadrivirgatum is recorded for the first time from Hidalgo,
Mexico. Comparison of variation in this and other populations of the species indicate
that A. q. newmanorum is identifiable only by pattern. The two taxa may be closely
associated in the vicinity of Xilitla, San Luis Potosi. Known localities for A.
quadrivirgatum sensu stricto are listed, and problematic records of that species are
summarized. Populations on Atlantic slopes eastward from the Isthmus of Tehuantepec
in Veracruz and Oaxaca represent A. visoninum. Those westward from Guatemala, in
southern Veracruz, adjacent Oaxaca, northern Chiapas, Tabasco and Belize, repre¬
sent A. visoninum, but are subspecifically distinct from A. v. visoninum of Guatemala
and Honduras, as A. v. acutirostris Bocourt (1883).
The genus Adelphicos embraces two complexes. The A. veraepacis complex
occurs in the western highlands of Guatemala, Chiapas and Oaxaca. The A.
quadrivirgatum (gender fide laDuc, 1995) complex extends on Atlantic slopes from
Monterrey, Nuevo Leon (David Lazcano, pers. comm.) to Honduras, and on Pacific
slopes in Chiapas and Guatemala. The former complex differs from the latter in lack¬
ing expansion of the anterior part of the anterior chinshields toward the lip; it was
monographed by Campbell and Ford (1982), and an additional species was described
by Campbell and Brodie (1988). We here deal solely with the A. quadrivirgatum
complex.
The entire A. quadrivirgatum complex was considered a single species (e.g.
Campbell and Ford 1982), until Smith et al. (2001) proposed that the three subspe¬
cies formerly recognized in the species (Smith, 1942) are separate species, A.
quadrivirgatum and A. visoninum Cope (1866) on Atlantic slopes, and A. sargii
(Fischer, 1885) on Pacific slopes of Chiapas and Guatemala. We are here primarily
concerned with the former two.
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Adelphicos quadrivirgatum Jan
We here report three specimens of this species from Hidalgo, the first known
from the state (only mentioned by Canseco-Marquez et al. , in press). They are of
special interest also because they are from an area between the known ranges of the
nominotypical subspecies and A. q. visoninum (Taylor, 1950). They are as follows:
ENCB (Escuela Nacional de Ciencias Biologicas, IPN) 11498, San Felipe, munici¬
pality of Orizatlan (21°10’25"N, 98°35,23”W), ca. 160 m elev., April, 1982; ITAH
(Institute Tecnologico Agropecuario de Hidalgo) 775, Coyolapa, mpio Atlapexco,
ca. 2.5 km N Atlapexco (21°08'N, 98°20'50"W), ca. 160 m elev., 2001; and ITAH
Table 1. Comparison of Selected Characters of Hidalgo Adelphicos quadrivirgatum
with Those Previously Recorded
A.q. quadrivirgatum A. q. newmanorum Hidalgo
Female Subcaudals
32
41-44
36, 37
Male Subcaudals
36-49
47-50
tail inc.
Female Ventrals
124-138
141-155
141, 145
Male Ventral
131-146
139-142
139
Stripes
continuous
absent or
interrupted
continuous
776, Chalahuiyapa, mpio Huejutla (20°09'N, 98°25'W), 170 m elev., 2001. All were
collected in disturbed tropical deciduous forest (Rzedowsky, 1981). The two ITAH
specimens were found dead on a gravel road.
In determining the subspecific identity of the Hidalgo specimens, it should be
noted that the only alleged differences between the two subspecies are in number of
ventrals and subcaudals, and in pattern (Martin, 1955; Taylor, 1950; Smith, 1942).
Table 1 compares known data for these characters.
The variation in scalation of the Hidalgo specimens does not support distinc¬
tion of the two subspecies on that basis: number of female ventrals, female subcaudals
and male ventral are all intermediate. The only consistent difference is in pattern,
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December 2003
which in the present specimens corresponds with that of A. q. quadrivirgatum, to
which we assign the Hidalgo sample. These are the northernmost specimens known
of the subspecies, except perhaps for one from the Xilitla region, San Luis Potosi
(Taylor, 1949, 1950), where Taylor (op. cit.) also recorded A. q. newmanorum, other¬
wise known only northward. The occurrence of a specimen referable to each subspe¬
cies in the Xilitla region (fide Taylor, op. cit.) may reflect altitudinal separation (inas¬
much as the area is of rugged topography), intergradation, or sympatry of two spe¬
cies. Additional material will be required to resolve that problem.
South of Hidalgo, the range of A. q. quadrivirgatum extends into northeastern
Puebla, as indicated by the following material. EBUAP 1094, Zacatipan, 600 m
(20°02'N, 97°26'W); EBUAP 112, Finca San Jose, 4 km W Santiago Yacuictlalpan,
360 m (20°03'N, 97°26'W); EBUAP 1095, 1 km SW Yohualichan, 750 m (20°03'N,
97°28'W); EBUAP 1096, 1 .5 km E Yohualichan, 725 m (20°03'N, 97°30’W); EBUAP
1097, Tatahuitaltipan, 3 km SW Yohualichan, 550 m (20°04,N, 97°30’W); EBUAP
1098, Octimaxal Norte, 930 m (20°02'N, 97°30’W); EBUAP 1099, 1338, 1 km N El
Paraiso, 720 m, mpio Huitzilan de Serden (Canseco-Marquez et al. , 2000). All ex¬
cept 1099 and 1338 are from mpio Cuetzalan del Progreso.
In the state of Veracruz, A. q. quadrivirgatum is known to occur at Jicaltepec
(Smith, 1942), to which the original type locality of “Mexico” was restricted by Smith
and Taylor (1950). No other confirmed record for the state is known to us, although
probably some of those cited as A. quadrivirgatum in Perez-Higareda and Smith (1991)
are actually of this species. However, all specimens of A. quadrivirgatum cited in that
work from the Los Tuxlas area actually are A. visoninum (Perez-Higareda, pers.
comm.), confirming the same identification recorded by Ramirez-Bautista ( 1 977) for
material from that area.
In northeastern Oaxaca, confirmed records of A. q. quadrivirgatum are avail¬
able from the Sierra de Juarez at 2 km NE Vista Hermosa, 1 800 m (MZFC 6522);
mpio Juan Yave (MZFC 1596); and Metates, mpio Valle Nacional, 800 m (MZFC
2239). Cadle (1984) recorded a specimen from hwy 185, 8 km S Veracruz state line.
Apparently the southern limit of the range of A. q. quadrivirgatum is represented by
these specimens.
The specimen from Pochutla, Oaxaca (Smith, 1942) is clearly A.
quadrivirgatum, but the locality is in error, as neither that species nor A. visonium is
known from Pacific slopes; both are limited to Atlantic slopes.
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Adelphicos visoninum Cope
Near the Isthmus of Tehuantepec, in Veracruxz and Oaxaca, A. visoninum re¬
places A. quadrivirgatum. The known Veracruz records are cited in a preceding para¬
graph.
Several specimens from Oaxaca in LJCM are clearly A. visoninum, all having
the anterior chinshields separated from the lip by a very narrow third infralabial. The
localities represented are 12 de Julio, mpio Mixe (39891-2); mpio Mixe (52516, prob¬
ably from the vicinity of 12 de Julio); Palomares, mpio Juchitan (49321); Lag Muelles,
Palomares, mpio Juchitan (39893); and Vista hermosa, mpio Ixtlan (39894, 52385).
All were collected by Thomas MacDougall. Occurrence in northern Chiapas was
also confirmed at Palenque (Smith, 1942).
The confirmed records of specimens with no third infralabial (the character of
A. quadrivirgatum) from within the range of A. visoninum have been assigned to A .
quadrivirgatum on that basis (e.g. Smith, 1942), in error. They are widely scattered,
with no geographic continuity. The third infralabial in Mexican A. visoninum is very
narrow - distinctly narrower than in Guatemale specimens of the same species - and
may on that account be subject to some variation, contrary to the populations in
Guatemala. Included among these aberrant specimens are those recorded by Smith
(1942) from Ocozucoautla, Chiapas, and Silkgrass Creek, Belize, and ENCB 1959,
14 km W Raudales, Chiapas. Two others are UCM 39890 from Palomares, mpio
Juchitan, Oaxaca, and UCM 60077, from “Oaxaca.” Both UCM specimens were
collected by Thomas MacDougall, hence the latter specimen is probably from the
same region as the former. At least the Palomares specimen is definitely from the
same locality as others that conform with A. visoninum (see above). That only one
taxon is involved is indicated also by the fact that male ventrals of A. visoninum are
only 117-129, including the large series reported by Wilson and Meyer (1985) from
Honduras, as opposed to 131-146 in male A. quadrivirgatum. All male ventrals for
Mexican and Belize “A. quadrivirgatum” (those without the third infralabial), fall
within the range of A. visoninum. The number of female ventrals overlaps in the two
species.
Variability in occurrence of the third infralabial apparently does not exist in
populations of A. visoninum (type locality “Honduras”) from Guatemala and Hondu¬
ras; the third infralabial is regularly present (and larger than in Mexican A. visoninum)
in the 66 UCM specimens available from Guatemala (Sayache, Peten Tamahu and
Beleu, Alta Verapaz), and in the 62 reported from Honduras by Wilson and Meyer
(1985).
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December 2003
The Mexican specimens of A. visoninum differ from the Guatemala/Honduras
populations in (1) the very small size of the third infralabial, even occasionally ab¬
sent (vs larger, regularly present); (2) the light dorsal ground color, on which the
typical triple dark stripes are clearly evident (vs ground color quite dark, sometimes
obscuring completely the dark stripes); (3) absence of ventral pigmentation even on
the head (vs usually extensive ventral pigmentation, always on head); and (4) small
size, maximum 351 mm total length (vs 526 mm). The differences are striking, and
may well indicate specific rank for the two populations. At least they justify resurrec¬
tion of A. v. acutirostrum Bocourt (1883) for the Mexican and Belize populations. Its
original type locality “Mexico” is here restricted to Palenque, Chiapas, where the
taxon has previously been reported (Smith, 1942). Presumably Duges’ (1896) record
for “Tabasco,” and Muller’s (1882) for Tenosique, Tabasco, are referable to A. v.
acutirostrum .
Key to the Species and Subspecies of the
A, quadrivirgatum Complex
1 A. Subaudals 24-29 in females (3 1 ), 29-35 in males (33); third infralabial always
present ............................................................................................... A. sargii
B . Subcaudals 3 1 or more in females, 36 or more in males; third infralabial present
or absent ........................................................................................................ 2
2A. Third infralabial absent, or, if present, male ventrals
131-146.......................................... A. quadrivirgatum ............................... .3
B. Third infralabial present or, if absent,
male ventrals 117-129 ............................... A. visonum ................................ 4
3A. Stripes on body absent, broken or weak ............................A. q. newmanorum
B. Stripes on body continuous, prominent ......................... A. q. quadrivirgatum
4 A. No ventral pigmentation, even on head; ground color light, triple
dark stripes prominent; third infralabial extremely narrow,
occasionally absent ............................................................. A. v. acutirostrum
B. Ventral pigmentation usually extensive, always present on head;
ground color dark, the triple dark stripes usually partially obscured;
third infralabial a little wider, always present ........................ A. v. visoninum
Bulletin of the Maryland Herpetological Society
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Volume 39 Number4
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Acknowledgments .
We thank Alejandra Ramirez Hernandez for her assistance in the laboratory,
and Hector Tovar Tovar and Baltazar Hernandez Hernandez for their help in the field.
Drs. Jonathan Campbell, Oscar Flores Villela, Linda Ford, Kenny Krysko, Adrian
Nieto Montes de Oca and Gonzalo Perez Higareda, as well as Mario Mancilla Moreno,
Alan Resetar and Greg Schneider, furnished vital information from specimens in
their collections.
Literature Cited
Bocourt, M. F.
1 883. Livr. 9, pp. 529-592, in Mission Scientifique au Mexique et dans
1'Amerique Centrale... Recherches zoologiques. Part 3, Sect. 1.
Etudes sur les reptiles. Paris, Imprimerie Imperiale. 1012 pp.
Cadle, J. E.
1 984. Molecular systematics of neotropical xenodontine snakes: II. Cen¬
tral American xenodontines. Herpetologica 40: 21-30.
Campbell, J. A. and E. D. Brodie, Jr.
1988. A new colubrid snake of the genus Adelphicos from Guatemala.
Herpetologica 44: 416-422.
Campbell, J. A. and L. Ford.
Phylogenetic relationships of the colubrid snakes of the genus
Adelphicos in the highlands of Middle America. Occ. Pap. Mus.
Nat. Hist. Univ. Kansas (100): 1-22.
Canseco-Marquez, L., G. Gutierrez-Mayen and J. Salazar-Arenas.
2000. New records and range extensions for amphibians and reptiles
from Puebla, Mexico. Herp Rev. 31: 259-263.
Canseco-Marquez, L., F. Mendoza-Quijano and M. Gutierrez-Mayen.
In press Analisis de la distribution de la herpetofauna de la Sierra Madre
Oriental. 29 pp.
Cope, E. D.
1 866. Fourth contribution to the herpetology of tropical America. Proc.
Acad. Nat. Sci. Philadelphia 18: 123-132.
page 82
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Volume 39 Number 4
December 2003
Dimes, A. A. D.
1896.
Reptiles y batracios de los Estados Unidos Mexicanos. Naturaleza
(2) 2: 479-485.
Fischer, J. G.
1885.
Ichthyologische und herpetologische Memerkungen V.
Herpetologische Bemerkungen. Jb. Hamb. Wiss. Anst. 1884: 82-
119.
Jan. G.
1862.
Enumerazione sistematico della specie d/ofidi del gruppo
Calamaridae. Archo. ZooL Anat. Fis. 2: 1-76.
LaDuc, T. J.
1995.
The nomenclatural status and gender of Adelphicos. J, Herp. 291 :
141.
Martin, P S.
1955,
herpetological records from the Gomez Farias region of southern
Tamaulipas, Mexico. Copeia 1955: 173-180.
Muller, F.
1892.
Siebenter Nachtrag zum Katalog der herpetologischen Sammlung
des Easier Museums. Verb. Naturf. Ges. Basel 10: 195-215.
Perez-Higareda, G. and H. M. Smith.
1991. Ofidiofauna de Veracruz: analisis taxonomico y zoogeografico.
Mexico, D. R, Univ. Nac. Aut. Mexico, Publ. Espec. 7: 1-122.
Raxnfrez-Bautista, A.
1 977. Algunos anfibios y reptiles de la region de “Los Tuxtlas,55 Veracruz.
Xalapa, Veracruz, Univ. Veracruzana. Biologo Tesis. 145 pp.
Rzedowsky, J.
1981.
Vegetation de Mexico. Mexico, Mexico, Ed. Limusa. 432 pp.
Smith, H. M.
1942.
A review of the snake genus Adelphicos. Proc. Rochester Acad.
Sci. 8: 175-195.
Smith, H. M., D. Chiszar and M. Mancilla-Moreno.
2001. Nomenclature of the earth snakes ( Adelphicos ) of the A.
quadrivirgatus complex. Bull Maryland Herp. Soc. 37: 39-41.
Bulletin of the Maryland Herpetological Society
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Volume 39 Number4
December 2003
Smith, H. M. and E. H. Taylor.
1950. Type localities of Mexican reptiles and amphibians. Univ. Kan¬
sas Sci. Bull. 33:313-380.
Taylor, E. H.
1 949. A preliminary account of the herpetology of the state of San Luis
Potosi, Mexico. Univ. Kansas. Sci. Bull. 33 (2): 169-215.
Taylor, E. H.
1950. Second contribution to the herpetology of San Luis Potosi. Univ.
Kansas. Sci. Bull. 33: 441-457.
Wilson, L. D. and I. R. Meyer.
1985 The snakes of the Honduras. 2nd edition. Milwaukee, Wisconsin,
Milwaukee Public Mus. x, 150 pp.
FMQ, Institute Tecnologico Agropecuario de Hidalgo , Carr. Huejutla-
Chalahuiyapa, Apartado Postal 94, Huejutla de Reyes, Hidalgo, 43000 Mexico.
JICR and JCLV, Laboratorio de Cordados Terrestres, Escuela Nacional de
Ciencias Biologicas, Instituto Politecnoco Nacional, Plan de Ayal, s/n, Esquina con
Prolongacion Manuel Carpio, A. P. 31-186, Casco de Santo Tomas, Mexico D. F.,
11340 Mexico.
MHS, DC, University of Colorado Museum, Boulder, Colorado, 80309-0334,
USA.
Received 15 May 2003
Accepted 29 May 2003.
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December 2003
Analysis of Phenotypic Variation in the Lizard
Sceloporus cautus and Adjacent Populations of
Related Species
George M. Ferguson, Hobart M. Smith and David Chiszar.
Abstract.
Data on external morphology of Sceloporus cautus confirm its allospecificity
from S. olivaceus and S. undulatus/edbelli. On the basis of these data S. cautus is
referred to the olivaceus group of Wiens and Reeder. S. cautus is reported for the first
time from the Mexican state of Tamaulipas.
In 1982 GMF was awarded a Master of Science deree at the University of
Texas at El Paso, in part based on a dissertation analyzing the external morphology of
the then controversial nominal species Sceloporus cautus in comparison with the
adjacent, related populations of S. olivaceus and S. “undulatus” (Ferguson, 1982).
At that time S. edbelli was not known, hence the “undulatus” populations sampled
actually represented both S. edbelli and S. undulatus consobrinus. We here maintain
the usage of “ undulatus ” as in the dissertation, with however the understanding that
in this context the name refers to both taxa. The differences here noted between S .
cautus and S. “ undulatus ” apply euqally well to S. edbelli and S . undulatus
consobrinus , the adjacent subspecies. Indeed, S. edbelli was originally thought to be
a subspecies of S. undulatus .
Although some 21 years have passed since the dissertation was completed, the
identity and relationships of S. cautus remain enigmatic. The latter and S. olivaceus
clearly belong to the same group, as shown hereinafter, based on phenetic similarity
so great that they have been suggested as synonyms by various workers (Bussjaeger,
1971; Hall, 1973). Confusion of S. cautus with S. “undulatus” and S. olivaceus is
evident in the literature as well as in museum identifications.
Therefore we here exhume the critical morphological data given in the disser¬
tation, comparing the three species. Twelve characters were regarded as the most
significant, here numbered as in Table 1 .
1. Dorsals. Number of transverse scale rows counted middorsally from inter¬
parietal to the level of the posterior margins of the hind limbs (directly above the
vent).
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2. Scales around the midbody. Number of longitudinal scale rows counted
around the middle of the body.
3. Femoral pores. Combined number of femoral pores along the posterior bor¬
der of the thighs.
4. Interfemoral pore scales. Fewest number of scales in a row separating me¬
dial ends of the pore series.
5. Dorsal body blotches. Number of dorsal body blotches (or crossbars) along
medial edge of dorsolateral light stripe, combined for the two sides. The character
could not be determined where the anterior blotches were fused longitudinally.
6. Dorsolateral stripe scales. The combined number of scale rows included
within the width of the dorsolateral light stripes. One row consisted of the area be¬
tween the keels of adjacent scales.
7. Scales between dorsolateral stripes. Number of longitudinal scale rows be¬
tween the dorsolateral light stripes. One row consisted of the area between the keels
of adjacent scales.
8. Gular semeion separation (males only). Fewest number of scales between
medial edges of posterior blue gular semeions, disregarding the black border.
9. Abdominal semeion separation. Fewest number of scales between the me¬
dial edges of the abdominal semeions, including the black borders where present.
10. Tail length divided by snout- vent length (TL-SVL).
11. Gular semeion extent (males only). Gular semeion extension anterior to
level of ear openings (2) or not (1).
12. Converging postocular stripes. Two dark, narrow lines, one originating at
the upper, the other at the lower border of eye. Fusion of the lines in temporal re¬
gions, character state 1; absence of fusion, character state 2.
Table 1 shows that the more diagnostically significant differences between S.
cautus and S. ” undulatus ” are 1, 3, 4, 10, 11 and 12, and those between S. cautus and
S. olivaceus 1, 2, 7, 10 and 12. There are no categorical distinctions of S. cautus from
S. “undulatus” (although 4 is nearly so), and none from S. olivaceus. Several less
diagnostically significant differences exist between S. cautus and S. “undulatus”
alone (2, 7, 9), S . olivaceus alone (5), and both (6, 8).
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December 2003
Table 1 . Analysis of variance comparing S. cautus with S. “undulatus” and S.
olivaceus for each of 12 morphological characters (sexes combined except for male
characters). Ranges are followed by the deviation (SD). F-values significant at the
0.01 level are indicated by *.
No.
S. cautus
S. “ undulatus ”
S. olivaceus
1
33-41 (36.85)
N=188SD=1.81
36-48 (41.60)
N=116 SD 2.55
F=358.53*
28-36 (31.49)
N=116SD=1.31
F=771.40*
2
36-45 (39.62)
N=187 SD=1.83
36-48 (41.96)
N-117 SD-2.49
F=88.80*
32-40 (35.43)
N=110SD=1.86
F=361.03*
3
20-31 (24.68)
N=185 SD-2.03
27-42 (33.41)
N=116 SD=2.32
F=1 177.76*
20-30 (25.25)
N=115SD=1.96
F=5.76
4
8-14(10.76)
N=185 SD-1.28
4-8 (5.52)
N=115 SD=1.01
F=1381.07*
7-12(9.40)
N=114SD=1.2
F=81.76*
5
14-23(19.16)
N=142 SD=1.38
10-22(19.78)
N=18 SD-2.84
F=2.35
14-22(16.76)
N=63 SD=1.38
F=1 32.65*
6
1-2(1.14)
N=123 SD=0.24
1-2(1.50)
N=62 SD=0.33
F=70.16*
1-2 (1.77)
N=90 SD=0.27
F=308.46*
7
6-8 (6.62)
N=123 SD-0.52
5-8 (7.00)
N=62 SD=0.65
F=18.65*
3-6 (4.30)
N=91 SD=0.53
F-1029.54*
8
1-9 (5.44)
N=84 SD=1.53
0-7 (2.49)
N=61 SD=1.76
F— 1 15.33*
5-11 (8.26)
N=43 SD=1.35
F=104.02*
9
3-8 (5.54)
N=95 SD=0.90
0-7 (4.60)
N-59 SD=1.30
F=28.38*
4-8 (5.87)
N=47 SD-0.97
F=4.17
10
1.04-1.58 (1.30)
N=87 SD=0.10
1.19-2.11 (1.62)
N=49 SD=0.15
F=224.44*
1.38-2.12(1.76)
N=75 SD=0.16
F=495.31*
11
1-2(1.05)
N=86 SD=0.21
1-2(1.89)
N=61 SD=0.32
F=363.64*
1 (1.00)
N=43 SD=0.0
F=2.07
12
1-2(1.17)
N=175 SD-0.37
2 (2.00)
N=117 SD=0.0
F^585.01*
1-2(1.95)
N=lll SD=0.23
F=393.13*
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Volume 39 Number4
December 2003
Data were taken on seventeen other characters, as follows, all of which showed
at least some level (p=< 0.01) of statistical significance, except for two (E, I).
A. Neck scales. Number of longitudinal rows of dorsales across neck between
uppermost auricular lobules.
B. Supraoculars. Combined number of supraocular scales, including smaller
scales if obviously associated with supraoculars rather than with adjacent scales.
C. Lorilabial rows. Fewest number of scales in horizontal rows between
subocular scales and supralabials (combined).
D. Auricular lobules. Number of enlarged scales along anterior margin of ear,
combined.
E. Fourth toe lamellae. Combined number of scales along ventral surface of 4lh
toe.
F. Ratio, 4th toe length/hind leg length (HL).
G. Ratio, foreleg length/HL.
H. Ratio, HL/snout-vent length (SVL).
I. Ratio, snout-axilla length (SA)/SVL.
J. Ratio, head width, maximum at anterior ear border (HW)/SA.
K. Ratio, HW/snout to anterior edge of ear.
L. Ratio, Head width between lateral edges of superciliaries (HS)/HW.
M. Ratio, HS/snout to occiput.
N. Ratio, interparietal length/width between lateral edges of parietals.
O. Head pattern (excluding S. “ undulatus ”) of dark stripes.
R Postrostral-subnasal in contact below nasal.
Q. Prefrontals in contact (excluding S. “undulatus”) or separated by one azy¬
gous scale.
Statistically significant differences exist between S. cautus from both of the
other species in characters A (the most significant in this set), C, G, J, K, L, M, N and
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P; from S. “undulatus” alone in B and D; and from S. olivaceus alone in F, H, O and
Q. No significant differences were found in E or I.
Geographic range is an important character also. That of 5. cautus is west from
the western edge of the Sierra Madre Oriental, barely entering Tamaulipas on the
northeastern edge of the Mexican Plateau, and extending south from southeastern
Coahuila and central Nuevo Leon to southern San Luis Potosi, east from the southern
half of Zacatecas. It is known only from the Mexican states mentioned. A narrow
zone of apparent sympatry of S. cautus and the S. edbelli component of S. “undulatus ”
occurs in Zacatecas. S. olivaceus is narrowly dichopatric (possibly parapatric) with
S. cautus , occurring to the north of its range and east of the Sierra Madre at their
common latitudes.
Subjectively, both S. cautus and S. “undulatus ” are categorically different from
S. olivaceus in behavior and habitat (terrestrial vs arboreal behavior, and non-for-
ested vs forested habitat, respectively). There is a significant difference (p=<0.001)
between the distributional elevation of S. olivaceus (12-1525 m, M=530 m) and that
of S. cautus (1000-2200, M=1860) in the study area. Altitudes recorded for “S.
undulatus” were 100-2200 m (M=1130).
The aggregate of differences between these three taxa leaves no doubt that
they are allospecific. Their relationships remain in question. Sympatry between S.
cautus and S. “undulatus, ” and their nearly categorical difference in at least one
character, suggests that they are more distantly related to each other than S. cautus is
to S. olivaceus . The ranges of the latter two species are allopatric (possibly parapatric)
and none of their differences are categorical. On these bases we regard S. cautus as a
member of the olivaceus group of Wiens and Reeder ( 1 997), rather than the undulatus
group, in which it has generally been placed (e.g. Smith, 1939; Sites etai, 1991), or
the spinosus group (Larsen and Tanner, 1975).
The following key distinguishes the three taxa in external morphology.
1 A. Interfemoral pore scales 4-7 . . . . . . . . . 2
B. Interfemoral pore scales 8-14 . . . . 3
2A. Interfemoral pore scales 4-7 (96.5%); femoral pores 29-42
(99.9%); dorsals 37-48 (98.3%); fused postocular dark
lines 0% . . . . . . . S. “undulatus”
B. A combination of interfemoral pore scales 7; femoral pores less than 29; and
dorsals less than 37 . . . . . S. olivaceus (<4%)
Bulletin of the Maryland Herpetological Society
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3A. A combination of interfemoral pore scales 8, femoral pores
>30, and dorsals >39 . . . . . S. “undulatus (<4%)
B. Interfemoral pores scales >8, OR, if interfemoral pore scales 8,
then femoral pores not >30 and dorsals not >39 . . . . . 4
4A. Interfemoral pore scales 8-14 (100%), 9.2% 13 or more; dorsals
34-41 (97.9%); scales around body 38-45 (89.3%); scales between dorsolat¬
eral light lines 6-8 (100%); fused postocular dark
lines (83.4%) . . . . . . . S. cautus
4B. Interfemoral pores scales never more than 12; dorsals 28-33
(92.2%); scales around body 32-37 (84.5%); scales between
dorsolateral light lines 3-5 (97.8%); fused postocular dark lines
(5.4%) . . . S. olivaceus
Acknowledgments,
We are much indebted to Dr. Robert G. Webb for his guidance of this study,
and to the curators of the following museums for loan of study material now in their
collections: AMNH, CAS, CM, FMNH, KU, LACM, LSUMZ, MCZ, MVZ, TCWC,
TNHC, TU, OCM, UMMZ, USNM, UTA, and UTEP. Many friends and colleagues
contributed to the completion of the dissertation, and are acknowledged in it.
Literature Cited.
Bussjaeger, L. J.
1971. Phylogenetic significance of the comparative ethology of the
spinosus group of Scloporus (Iguanidae). Norman, Univ. Okla¬
homa, Ph.D. Diss.
Ferguson, G. M.
1982. Distribution, variation and phenetic relationships of the lizard
Sceloporus cautus Smith in northeastern Mexico. El Paso, Texas
Univ. Texas at El Paso. MS dissertation, xiii, 195 pp.
Hall, W. P.
1973. Comparative population cytogenetics, speciation, and evolution
of the iguanid lizard genus Sceloporus. Cambridge, Massachu¬
setts, Harvard Univ., Ph. D. Diss.
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Volume 39 Number 4
December 2003
Larsen, K. R. and W. W. Tanner.
1 975. Evolution of the sceloporine lizards (Iguanidae). Great Basin Nat.
35: 1-20.
Sites, J. W., jr., J. W. Archie, C. J. Cole and O. Flores Villela.
1992. A review of phylogenetic hypotheses for lizards of the genus
Sceloporus (Phrynosomatidae): implications for ecological and
evolutionary studies. Bull. Am. Mus. Nat. Hist. (213): 1-110.
Smith, H. M.
1939. The Mexican and Central American lizards of the genus
Sceloporus . Zool. Ser., Field Mus. Nat. Hist. 26: 1-397.
Wiens, I. J. and T. W. Reeder.
1997. Phylogeny of the spiny lizards (Sceloporus) based on molecular
and morphological evidence. Herp. Monogr. (11): 1-101.
GMF: Department of Ecology and Evolutionary Biology, University of Arizona,
Tucson, AZ 85721 (georgef@email.arizona.edu).
HMS: Department of EPO Biology, University of Colorado, Boulder, CO 80309-
0334 ( hsmith @buffmail. Colorado, edu).
DC: Department of Psychology, University of Colorado, Boulder, CO 80309-0345
( chiszar@clipr. Colorado, edu ).
Received: 23 May 2003
Accepted: 3 June 2003
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2001-2002 Anurans, Exclusive of Rana, from
Durango and Chihuahua, Mexico
Julio A. Lemos- Espinal, Hobart M. Smith and David Chiszar
Abstract.
Fifteen anuran species are reported, including a first state record (Pachymedusa
dacnicolor) as well as numerous new locality records for the state of Chihuahua,
Mexico.
The material reported here was obtained in the summers of 2001 and 2002, all
from Chihuahua except for the few, so indicated, from Durango. The salamanders
and Rana collected at the same time have been reported elsewhere (Lemos-Espinal)
et al., in press a, b). All specimens are in the collection of the Unidad de Biologfa,
Tecnologia y Prototipos (UBIPRO), UNAM, to which catalog numbers refer.
Bufo cognatus Say. 7396, 8001-14, betw Ejido San Dionisio and Sierra La
Campana, mpio Tlahualilo, Durango (26012'9.1MN, 103°41'47.2"W), 111 m, ^Sep¬
tember; 10287, pasture E side Ojo Laguna, betw Ejido and lago (29°36'47.8"N,
106°10'34.1"W), 1540 m, 15 September. This latter specimen is 22 mm SVL, taken
with a large series, 10201-10363.
The species was previously reported from Ejido San Dionisio by Lemos-Espinal
etal (2001).
At both sites Bufo debilis was also collected. Stebbins (2003) mapped the range
of B. cognatus into the Sierra Madre Occidental, where it does not occur.
Bufu debilis insidior Girard. 6792, Llano El Victorio (29°55'15.9"N,
104°39,58.7"W), 1282 m, 9 April; 7598-7602, Rancho La Bamba (30°5TL7MN,
105°24'30.9"W), 1390 m, 30 July; 7609-10, Rancho El Setenta (31°11T6.2"N,
106°30'20.7"W), 1334 m, 1 August; 7892, betw Ejido San Dionisio and Sierra la
Campana, mpio Thahualilo, Durango (26°12,9.1,,N, 103°41'47.2"W), 1111 m, 1 Sep¬
tember; 10364-8, 10384, pasture E side of Ojo Laguna, betw Ejido and lago
(29°26'47.8iEN, 106°19,34.1MW), 1540 m, 15 September.
Bufo marinus (Linnaeus). 8579-8729, Batopilas (2706'53.8”N,
107°39'52.4"W), 867 m, 16 June. All are transformlings ~19 mm SVL, some with
vestige of the tail.
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The species was first reported from Chihuahua by Tanner (1989).
Bufo mazatlanensis Taylor. 7327, Satevo (26°59'25.8"N, 107°25'52.9"W) 567
m, 18 June; unassigned no., Batopilas (27°T34.T'N, 107°45'44.5"W), 435 m, 18
June; 10608, Arroyo Las Boregas (27°23'4.3MN, 108°32’21.r'W), 470 m. All have a
conspicuously deep valley between the interorbital crests, and a small, vertically oval
parotoid gland. A vertebral light stripe is complete in two, anterior half only in one.
Bufo mexicanus Brocchi. 8074-8, Rio Pagigochi at La Junta, mpio Guerrero
(28°27'54.0"N, 107°19'39.4"W), 2103 m, 6 September; 8148, Arroyo Seco, km 16.5
hwy 127 (28°15'45.4"N, 107°29,31.5'W), 2191 m, 7 September; 8299-8300, Cusarare,
small stream (27°37'22.4"N, 107°32'38.4"W), 2302 m, 9 September; 8419, Napuchis,
mpio Guerrero (27°18T9.1MN, 107o3r40.2"W), 2179 m, 11 September; 8494,
Basigochi de Aboreachi, mpio Guachochi 927°12T2.2MN, 107°22'45.0"W), 2409 m,
11 September; 8541, San Pablo Balleza (26°55I48.3MN, 106°20'57.3"W), 1638 m, 11
September; 9534-40, km 7.8, San Juanito-Basaseachi (27058'41.1"N, 107°39,43.0"W),
2402 m, 15 July; 9593-4, km 12, San Juanito-Basaseachi (27°58’17.9MN,
107°49T5"W), 2375 m, 15 July; 9650, 13 km N Maguarichi (27°53’58.8HN,
107°56'35.1"W), 1923 m, 19 July; 9671-2, Gurichivo-Basogachi (27°59T1.3"N,
107°51'55.6"W), 2127 m, 19 July; 9675-6, 2 km S jet San Juanito-Basaseachi and
hwy to Maguarichi (2801'8.1"N, 107°48'57.4"W), 2313 m, 19 July; 10579-80, km
1 14 Creel-Guachochi 927°35’32.6,,N, 107°32,56.9"W, 2139 m, 26 September; 10943-
64, km 7.6 San Juanito-Maguarichi (27°58,41.1UN, 107°39'43.0"W), 2402 m, 12
October.
A few adults are very dark, somewhat obscuring the diagnostic light band across
the eyelids. Webb (1972), although concerned primarily with distinguishing B .
speciosus from B . mexicanus, provides important data distinguishing the latter spe¬
cies from the sympatric B. occidentalis. The latter occurred with B. mexicanus, B.
woodhousii and Scaphiopus multiplicata at km 7.6, San Juanito-Basaseachi.
Bufo occidentalis Camerano. 8074-8, Rio Papigochi at La Junta (28°27'54.0"N,
107°19'39.4"W), 2103 m, 6 September; 8730-93, km 121.7, hwy 127, nr detour to
Tejaban, mpio Guachochi (27°33T1.5"N, 107°31,47.3,,W), 2332 m. 10 September;
8794-7, Rio Papigochi at Guerrero (28°40'6.8MN, 107°34,5.5”W), 1979 m, June 15;
9511-33, km 7.8, San Juanito-Basasacheachi (27°58'41.1"N, 107°39,43.0"W), 2402
m, 15 July; 9553, 9621, Huerta de Manzana y Durazno, N Rancho Mojarachi
(27°51’46.4,!N, 107o55'47.0"W), 2211 m, 15-16 July; 9662-3, Aserradero
(27°55'5.2MN, 107°54T5.4"W), 2172 m, 19 July; 9690-1, Huevachic (28°6,26.2,,N,
108°0'03.rw), 2115 m, 20 July; 9702-5, km 82, San Juanito-Basaseachi (28°9,25.3I,N,
108°6I26.9"W), 2320 m, 20 July; 10563, Arroyo del Agua, nr Maguarichi
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(27°54'45.6"N, 107°58'49.2"W), 2083 m, 25 September; 10960, km 7.6 San Juanito-
Maguarichi (27°58,41.1"N, 107°39'43.0”W), 2402 m, 12 October.
Nos. 8730-93 are transformlings -12 mm SVL, with a vestigial tail.
Relatively few previous records for Chihuahua exist (Tanner, 1989, as Bufo
simus). Riemer (1955) first reported the species for the state, at Rio Gavilan, 7 mi SW
Pacheco, 5700 ft. Van Devender and Lowe (1977) and Van Devender et al (1989)
reported it from other northern localities. The present localities considerably aug¬
ment knowledge of the distribution of the species in the state.
Bufo punctatus Baird and Girard. 6751-2, 7024-5, Rancho El Escondido
(29°55T2.6"N, 105°32'9.8MW), 1426 m, 8 April, 9 June; 7539, Rancho El Gatunozo,
mpio Camargo (26°6’50.7"N, 104°5'51.4,,W), 1325 m, 28 July; 9166, Canon de
Balleza, mpio Balleza (26°57'45.7"N, 106°26,9.6"W), 1699 m, 12 May; 9847, 1 km
S Red Rock, Rio Riedras Verdes (30°22T1.4"N, 108°14,8.7”W), 1682 m, 24 July;
10130-1, plains base Sierra del Nido, nr La Providencia (29°40'22.3 "N, 106°37'4.4"W),
1569 m, 6 August; 10378-9 pasture E side Ojo Laguna, betw Ejido and lago
(29°26'47.8"N, 106°19'34.1"W), 1540 m, 6 August.
All lack a vertebral light line, have light-tipped warts and a black-speckled
chest, thus comforming with characteristics of populations in the eastern part of the
range of the species, but different from the southwestern specimen reported by Lemos-
Espinal et al (2001). No. 10131 is 69 mm SVL.
This species was taken with B. cognatus, B. debilis, Scaphiopus couchii and
Spea stagnalis between Ojo Laguna and Ejido.
Bufo woods housii australis Shannon and Lowe. 7208-96, 8561-78, Rio Verde,
Pacheco (30°51T.7MN, 108°20'29.4"W), 1949 m, June 14; 10015, middle Canon de
la Madera, Sierra de San Luis (31013'4LrN, 108°44’5.5MW), 1638 m, 27 July. The
specimens from Pacheco are all transformlings, 10-13 mm SVL.
Hyla arenicolor Cope. 6701-2, Ejido Dolores, Guadalupe y Calvo
(25°58'57.6"N, 107°10T1.5"W), 954 m, 3 April; 8080-82, Rio Pagicochi at San Pedro,
mpio Guerrero (28°23'35.9,,N, 107°26'8.6"W), 2081 m, 6 September; 8084-8147,
Arroyo Seco, km 16.5 hwy 127 (28°15,45.4"N, 107°29,35.1"W), 2191 m, 7 Septem¬
ber; 8276-9, hwy 127 , mpio Bocoyna (27°39,51.2,,N, 107°33'53.3"W), 2313 m, 9
September; 8288-96, 8313-5, Cusarare, small stream (27°37'22.4MN, 107°32’38.4nW),
2302 m, 9 September; 9024, 9054-7, 9246, Arroyo El Camuchil (27°1,34.1"N,
107°45'44.5"W), 435 m, 8 May; 9490-3, 1 km N Humira (27°25,43,,N,
107°29'24.6"W), 1906 m, 13 July; 9664, Aserradero 500 m N detour Mojarachi
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(27°55'5.2f,N, 107°54,15.4,,W), 2172 m, 19 July.
Most are small, 24 mm SVL or less; others are 29-38 mm SVL. The specimens
from El Camuchil were taken at a surprisingly low elevation. No water was in the
stream bed, but the frogs were in a quite wet cave - a typical habitat.
Hyla wrightorum Taylor. Seven: 6701-2, Ejido Dolores, Guadealupe y Calvo
(26°58'57.6"N, 107°10,1L5,,W), 954 m, 3 April; 8241-2, km 48 Creel-San Rafael,
mpio Urique (27°31'18.2MN, 107°50,50.5"W), 2313 m, 8 September; 8301-6, Cusarare,
small stream (27°37'22.4S,N, 107°32,38.4"W), 2302 m, 9 September; 8417-8,
Napuchis, mpio Guerrero (27°18T9.1nN, 107°31,40.2,,W), 2179 m, 11 September;
8535, Guazarare, mpio Guachochi (27°3'47.2"N, 107°12'5.0WS!), 2300 m, 11 Sep¬
tember; 9542 [9542-46], km 7.6, San Juanito-Basaseachi (27°58'41.1"N,
107°39'43.0"W), 2402 m, 15 July 9590-2, km 12, San Juanito-Basaseachi
(27°58T7.9"N, 107°49T5.0"W), 2375 m, 15 July; 9673-4, Gurichivo-Basogachi
(27°59T1.3MN, 107°51I55.6,,W), 2127 m, 19 July; 10564, Arroyo del Agua, nr
Maguarichi (27°54'45.6,,N, 107°58'49.2nW), 2083 m, 25 September.
Pachymedusa dacnicolor (Cope). A single, slightly injured, partially trans¬
formed larva (9068) 28 mm SVL, tail -10 mm, is from Ejido la Junta, 203 km NW
Batopilas (27°1,34.1,,N, 107°45'44.5,,W), 435 m, 19 May. Its identification, prima¬
rily due to its small size, is based on elimination. It is not Hyla arenicolor, of which
specimens of much the same size are available from nearby El Camuchil, because of
its smooth skin and different pattern. No other hylids are known from southwestern
Chihuahua, although Pachymedusa dacnicolor, Smilisca baudinii and Pternohyla
fodiens are known from nearly Sonora and Sinaloa. It differs from both of the latter
two, totally lacking a pattern. Its toe webbing is much shorter than in S. baudinii, and
the metatarsal tubercles are much smaller than in P. fodiens. All of these features
correspond with the characters of P dacnicolor, with which it agrees in pigmentation.
The body is rather densely and uniformly pigmented, and the limbs are much more
lightly pigmented (appearing almost white to the naked eye) but also uniformly pig¬
mented. The species is known from very near Chihuahua, 13 km ESE Alamos, Sonora
(Due 11 man, 2001), but the present specimen is the first known of the species from
Chihuahua.
The specimen was taken from the Rio Batopilas in an area of dense aquatic
vegetation, where numerous Rana magnaocularis were seen. The frogs were very
wary, leaping into the water upon close approach; only three small Rana (28-42 mm
SVL) could be caught, along with the Pachymedusa.
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Scaphiopus couchii Baird. 7591, Cd. Coyame (29°27’48.2"N, 105°511.2MW),
1270 m, 29 July; 7732 llano 4 km N Villa Ahumada (30°41,20.7”N, 106°30,57.2,,W),
1202 m, 2 August; 7791-3, Rancho El Vergel, nr Samalayuca, mpio Juarez
(31°13'4.5"N, 106°37,51.7"W), 1009 m, 1 August; 7893-5, 7897-8, Ejido San Dionisio,
mpio Tlahualilo, Durango (26° 1 2’9. 1 ”N, 103o41'47.2"W), 1111m, 1 September;
10128-9, plains at base of Sierra del Nido, nr La Providencia (29°40'22.3"N,
106°37'4.4"W), 1569 m, 6 August; 10369-77, pasture E side of Ojo Laguna, betw
Ejido and lago (29°26’47.8"N, 106°19,34.1"W), 1540 m, 15 September; 10505, Ojo
de Agua, Estacion Guzman (31°13T9.7"N, 107C278.2"W), 1449 m, 21 September;
10634, Jaco (27°5734.1MN, 103°57T6.0"W), 1283 m, 13 October.
Spea bombifrons (Cope). 7611-4, Rancho El Setenta (3 1 ° 1 1'16.2"N,
106°30’20.7"W), 1334 m, 1 August. Two adults and two transforming larvae, 28 mm
SVL, tail 27, 29 mm, partially larval mouthparts.
Spea multiplicata (Cope). 9534-40, km 7.6 San Juanito-Basaseachi
(27°58'41 .1"N, 107°39'43.0"W), 2402 m, 15 July; 10130-1, plains base of Sierra del
Nido, nr Providencia (29°40'22.3HN, 106o37'4.4"W), 1569 m, 6 August.
Spea stagnalis (Cope). 7732, 4 km N Villa Ahumada (30°4r20.7"N,
106°30'57.2"W), 1202 m, 2 August; 7791-3, Rancho El Vergel, nr Samalayuca
(31°ir58.8"N, 106°35'44.3"W), 1248 m, 1 August; 10369-77, 10382-3, pasture E
side Ojo Laguna, betw Edjido and lago (29°26'47.8"N, 106°19,34.1"W), 1540 m, 15
September.
Acknowledgments.
We are much indebted to CONABIO for support under projects U003, X004
and AE003.
Literature Cited.
Conant, R. and J. T. Collins.
1998. A field guide to reptiles and amphibians: eastern and central north
America. Third edition, expanded. New York, Houghton-Mifflin.
xix, 616 pp.
Degenhardt, W. G., C. W. Painter and A. H. Price.
1 996. Amphibians and reptiles of New Mexico. Albuquerque, Univ. New
Mexico, xix, 616 pp.
page 96
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December 2003
Duellman, W. E.
2001. Hylid frogs of Middle America. Ithaca, New York, Soc. Study
Amph. Rept., Contru. Herp. 18 (2 vols.).
Lemos-Espinal, J. A., D. L. Auth, D. Chiszar and H. M. Smith.
2001. Year 2000 amphibians taken in Chihuahua, Mexico. Bull. Mary¬
land herp. Soc. 37: 151-155.
Lemos-Espinal, J. A., D. Chiszar and H. M. Smith.
In press, a. Observations on ambystomatid salamanders of Chihuahua,
Mexico. Bull. Chicago Herp Soc.
Lemos-Espinal, J. A., D. Chiszar and H. M. Smith.
In press, b. Distributional and variational data on the frogs of the genus Rana
in Chihuahua, Mexico, inlcuding a new species. Bull. Maryland
Herp. Soc.
Riemer, W. J.
1955.
Comments on the distribution of certain Mexican toads.
Herpetologica 11: 17-23.
Stebbins, R. C.
2003.
A field guide to western reptiles and amphibians. Boston,
Houghton Mifflin, xv, 533 pp.
Tanner, W. W.
1989.
Amphibians of western Chihuahua. Gt. Basin Nat. 49: 38-70.
Van Devender, T. R., R A. Holm and C. H. Lowe, Jr.
1989. Life history notes: Pseudoeurycea belli sierraoccidentalis. Herp
Rev. 20: 48-49.
Van Devender, T. R. and C. H. Lowe, Jr.
1977. Amphibians and reptiles of Yepomera, Chihuahua, Mexico. J.
Herp. 11:41-50.
Webb, R. G.
1972. Resurrection of Bufo mexicanus Brocchi for a highland toad in
western Mexico. Herpetologica 28: 1-6.
DC and HMS: University of Colorado Museum, Boulder, Colorado,
80309-0334 USA
Bulletin of the Maryland Herpetological Society
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Volume 39 Number4
December 2003
Received:
Accepted
2 June 2003
10 June 2003
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December 2003
The Madrean Alligator Lizard of the Sierra del Nido,
Chihuahua, Mexico (Reptilia: Sauria: Anguidae)
Hobart M. Smith , Julio A. Lemos-Espinal, David Chiszar and Matthew J. Ingrasci
Among specimens collected by JLE in Chihuahua during the summer of 2002
are eleven examples of the genus Elgaria. Ten, from the central western Sierra Madre
Occidental, are typical E. k. kingii. The other specimen is from the isolated Sierra del
Nido, close to the north central part of the state, farther east than the genus has ever
been taken before in Chihuahua. That specimen exhibits pattern features not known
in E. kingii, the only other species of the genus in Chihuahua. It is here named
Elgaria usafa sp. nov.
Holotvpe.
UBIPRO (Unidad de Biologfa, Tecnologia y Prototipos, UNAM) 10177, taken
at the ruins of Rancho El Mesteno Chiquito (27051'46.5"N, 107°55'47.0"), Sierra del
Nido, 2223 m, on August 8 by JLE and MJI.
Diagnosis and Definition.
A member of the Elgaria kingii species complex with dim, dark tan interspaces
mostly one scale long between uniformly dark brown or black crossbars; numerous,
irregular black spots and vertical streaks, interspersed with light scales, on sides of
body; most of lateral fold dark; dorsal surfaces of head and neck anterior to the inter¬
space on the anterior border of the crossband at foreleg level nearly uniformly dark,
lacking any light marks except for scattered small light flecks. Reduced intemasals
and lateral supraoculars remain to be evaluated with additional material.
Description.
The only known specimen, the holotype, is 130 mm SVL - about the same as
the maximum recorded for E. kingii (Stebbins, 2003). Dorsal scale rows 16; ventral
scale rows 12; dorsals posterior to interparietal 53; ventrals mental to preanals inclu¬
sive 69.
Two supranasals broadly in contact medially posterior to rostral; no intemasal
on left, a small, elongate one on right at the side of the frontonasal; latter large,
partially fused with right prefrontal; frontal contacting interparietal, separating the
two frontoparietals; parietals 1-1.
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Five large medial and two small lateral supraoculars on each side; 5-5
superciliaries; 2-2 postnasals; 2-2 loreals; 1-1 canthals above anterior loreal and con¬
tacting posterior loreal, upper postnasal and frontonasal (and, on the right side, the
posterior end of the intemasal). Preoculars 1-1, suboculars and postoculars 2-2;
supralabials 10-10; anterior temporals 3-3; posterior temporals 4-4, none excluded
from an anterior temporal; infralabials 10-10; postmentals 1-1, in medial contact and
contacting the first two infralabials; three large chinsheilds on each side, anterior pair
in medial contact, all separated from infralabials. Tail incomplete.
Dorsal color very dark, the light interspaces only dimly distinct from the 9(10)
dark crossbands on body, the first at foreleg level. Light interspaces between crossbands
mostly one scale long, with some narrow interruptions by adjacent dark bands; latter
3-4 scales long, and little if any darker on their posterior borders than anterior to
them; sides of body with irregular, vertical rows of dark or dark-edged scales inter¬
spersed among white scales; lateral fold mostly dark, with scattered, small light spots,
each covering a few granules; lateral three rows of ventrals with dark edges on nu¬
merous scales; ventral surfaces of head, body and tail unmarked. No light marks on
dorsal surfaces of the black head and neck anterior to the light border of the crossband
at foreleg level, except for tiny, scattered light flecks; supralabial area strongly barred.
Comparisons.
The ten examples previously mentioned of E. k. kingii are the chief bases for
comparison with E. usafa, augmented by the guides and revisions by Good (1988),
Stebbins (2003), and Webb (1962, 1970). Pattern differences are categorical, consis¬
tent in the material available and in the literature, and therefore indicative of species
rank. In E. kingii, light stripes and other markings on a dark head and neck are regu¬
larly present except where the entire area is light; the interspaces between the
crossbands are sharply distinct and two or more scales long; the crossbands are dark
brown with black posterior borders; the sides of the body are less extensively and
less irregularly mottled; and the lateral fold has considerably more extensive light
areas.
The quantitative differences in scales (single, reduced intemasal, and two lat¬
eral supraoculars) of the single E. usafa from E. kingii are subject to variation in the
latter species and likely will not prove with further material to be taxonomically
useful. Among the 10 E. kingii examined, one has no intemasals, one has a tiny one
on one side, and one has the two scales well separated medially. Six of the 10 have 3-
3 lateral supraoculars; three have 2-3; and one has 2-2. However, the latter and one
with 2-3 lateral supraoculars have an anomalous relationship with the superciliary
series, which in both cases are interrupted, having 3+ 1-3-1 and 2+1-5 scales respec-
page 100
Bulletin of the Maryland Herpetological Society
Volume 39 Number4
December 2003
lively, instead of the usual continuous series of 4 to 6 scales. In E. usafa the
superciliaries are 5-5 in an uninterrupted series.
Etymology.
The specific name usafa is an acronym for United States Air Force Academy,
the personnel of which, particularly of the Department of Biology, made the semester
spent there by one of us (DC) a cherished memory. USAFA also generously sup¬
ported the work that led to the discovery of E. usafa.
Remarks.
A few decades ago, this taxon would probably be regarded as a subspecies of
E. kingii, to which it is obviously closely related. An example of such points of view
is the endemic Crotalus willardi amabilis Anderson of the Sierra del Nido. Current
understanding, however, regards categorically distinct, isolated populations of any
species group as evolutionarily independent, hence species. The present sample ex¬
hibits a pattern unlike that of any other taxon of the E. kingii complex, and is widely
isolated from others; we therefore conclude that it represents a species.
Acknowledgments.
We are much indebted to CONAB 10 for support to JLE under projects U003,
X004 and AE003.
Literature Cited.
Good, D. A.
1988. Phylogenetic relationships among gerrhonotine lizards: an analy¬
sis of external morphology. Univ. California Publ. Zook 121: i-x,
1-139.
Stebbins, R. C.
2003. A field guide to western reptiles and amphibians. Third edition.
New York, Houghton Mifflin, xv, 533 pp.
Webb, R. G.
1 962. A new alligator lizard (genus Gerrhonotus) from western Mexico.
Herpetologica 18: 73-79.
Webb, R. G.
1970. Gerrhonotus kingii. Cat. Am. Amph. Rept. (97): 1-4.
Bulletin of the Maryland Herpetological Society
page 101
Volume 39 Number 4
December 2003
HMS: Department of Ecology and Evolutionary Biology, University of Colorado,
Boulder, Colorado 80309-0334.
JLE: Laboratorio de Ecologia, UBIPRO, Facultad de Estudios Superiores
Iztacala, UNAM, Apartado Postal 314, Avenida de los Barrios, No. 1, Los Reyes
Iztacala, Tlalnepantla, Estado de Mexico, 54090 Mexico.
DC: Department of Psychology, University of Colorado, Boulder, Colorado 80309-
0345.
MJ1: Department of Biological Sciences, University of Notre Dame, Notre Dame,
Indiana 46556.
Received 1 3 June 2003
Accepted 2 July 2003
page 102
Bulletin of the Maryland Herpetological Society
Volume 39 Number 4
December 2003
News and Notes
Book Review
Amphibians and Reptiles of Delmarva, by James F. White, Jr. and Amy
Wendt White, 2002, xvi + 248 pp. + 96 pis. Tidewater Publishers, Centreville, Mary¬
land. ISBN 0-87033-543-X, (paperback) $14.95.
The authors have provided the first field guide to the amphibians and rep¬
tiles of the Delmarva Peninsula which consists of the entire state of Delaware and
portions of the states of Maryland and Virginia. The first naturalists having pub¬
lished on the herpetofauna of Delmarva were Henry W. Fowler and Roger Conant.
In the early part of the 12th century, Fowler collected extensively, and published his
findings in Copeia. In 1936 Roger Conant began his systematic and comprehensive
Delmarva investigation into the herpetofauna, which was followed by Clyde F. Reed
studies in the 1950s.
The book opens with a brief introduction and historical account of the
Delmarva Peninsula along with a short introduction on the physiography of the area.
The Atlantic Coastal Plain and Piedmont Provinces cover the major portion of the
territory separated by the Fall Line. The Fall Line serves as a vital boundary for
many of the Delmarva amphibian and reptile species. This is followed by a short
discussion on how to find the organisms in addition to guidelines for handling and
precautions regarding collecting herptiles , followed by addresses for reporting un¬
usual or rare species found within the Delmarva Peninsula.
Chapter 5 relates to conservation of the Delmarva herpetofauna with a short
discussion on causes of amphibian and reptile declines, conservation and manage¬
ment, species of special concern, and an appendix for recording notes on species
cited within the province.
This well- illustrated field guide size (185 x 44 cm) volume covers seventy-
three species with each account containing a detailed description for the species,
followed by comparisons of similar species, and information on the geographical
distribution, reproduction and development, and comments on population declines.
This volume contains 96 color plates centrally located within the text, and all are of
unusual or excellent quality.
The bibliography has only one fault, as it fails to cite any references from the
Catalogue of American Amphibians and Reptiles. Otherwise the authors cite 312
references, which is amazing considering the size of the territory involved.
Bulletin of the Maryland Herpetological Society
page 1 03
Volume 39 Number4
December 2003
News and Notes
This landmark field guide will certainly be a welcome addition to the litera¬
ture, and will hopefully educate and stimulate younger generations to appreciate
these awesome creatures. The price will certainly make it available to anyone inter¬
ested in the herpetofauna of this area, and any other bibliomaniac friends.
Harlan D. Walley, Department of Biology, Northern Illinois University,
Dekalb, Illinois 60115. hdw@niu.edu
Received 21 June 2003
page 104
Bulletin of the Maryland Herpetological Society
Volume 39 Number 4
December 2003
News and Notes
Book Review
Introduction to Horned Lizards of North America, by Wade C. Sherbrooke,
2003. California Natural History guide Series Number 64, xiii +178 pp. Illus. with
164 color photographs, university of California Press, Berkeley. 0-520-22827-8 $16.95
(paper), 0-520-22825-1 $35.00 (cloth).
The author has published extensively on the Horned Lizards of North
America, with his first major review appearing in 1981, and his PhD thesis on the
Integumental Biology appearing seven years later.
This book is a concise introduction to the natural history and evolution of
the homed lizards, and their impact on ancient Indian rock art through modem art.
The book opens with a short historical sketch along with remarks on diversity of
form in relationship to the environment. This is followed by a unique flow
diagrammatical key for the identification of the 13 species found within Mexico,
Southwestern North America and Canada.
The book is extremely well illustrated with photographs of each species, and
habitat. This is followed with a short discussion of convergent evolution between the
horned lizards of the New World family Iguanidae, and the Australian Thorny Devil
(Moloch horridus) of the Old World lizard family Agamidae. This is well illustrated
with photographs of both genera in defensive display, and feeding and drinking be¬
havior.
The Natural History section covers the activity cycle, enemies, defense, repro¬
duction and human impact, and is well illustrated with photographs on morning emer¬
gence, orientation to morning sun, burrowing behavior, removal of sand particles
from nasal and eye areas, sleeping behavior, mimicry, color changes during thermo¬
regulation and pigmentation. The physiological section is highly informative, with
excellent photographs of cryptic coloration, defensive behavior (blood squirting),
and predation. This is followed by an excellent chapter on reproduction. The last
chapter relates to the homed lizard in relation to man. the cliff-dwelling Anasazi and
Hohokam of the southwestern United States have numerous pottery artifacts illus¬
trating the horned lizard design for the part 2000 years, this tend continues in stone
and wood carvings and ceramic pottery of the Zuni Indians of modem times.
Overall, the book is extremely well written, and the major portion of the pho¬
tographs are of high quality, although the plate on page 49 of Phrynosoma solare
Bulletin of the Maryland Herpetological Society
page 105
Volume 39 Number4
December 2003
News and Notes
should have been taken with a flash to eliminate the dark backgrounds, and dull
appearance.
The book is actually written in field guide fashion, as the author provides a
selected reference section with 91 author citations, although none of these references
are cited within the text. I would highly recommend this book to anyone interested in
the saurology of North America, and the price is certainly reasonable.
Harlan D, Walley, Department of Biology, Northern Illinois University,
Dekalb, Illinois 60115. hdw@niu.edu
Received 28 July 2003
page 106
Bulletin of the Maryland Herpetological Society
Volume 39 Number4
December 2003
News and Notes
Errata:
In the last issue, Vol. 39, No. 3 on p. 52 the Received and Accepted dates were
accidentally omitted and are as follows...
Received: 27 May 2003
Accepted: 17 August 2003
Bulletin of the Maryland Herpetological Society
page 107
Volume 39 Number4
December 2003
News and Notes
Reptile and
Amphibian Rescue
410-580-0250
We will take as many unwanted pet reptiles and
amphibians as space allows.
Leave a message with your name and number to
give up an animal for adoption;
or to volunteer to help with our efforts.
OUR CURRENT NEEDS:
• Station Wagon or Van
• UV Lights • Power & Hand Tools • Bleach
• Equipment & Food • Paper Towels
www.reptileinfo.com
page 108
Bulletin of the Maryland Herpetological Society
Volume 39 Number 4
December 2003
News and Notes
Bulletin of the Maryland Herpetological Society
page 109
Society Publication
Back issues of the Bulletin of the Maryland Herpetological Society, where
available, may be obtained by writing the Executive Editor. A list of available
issues will be sent upon request. Individual numbers in stock are $5.00 each,
unless otherwise noted.
The Society also publishes a Newsletter on a somewhat irregular basis.
These are distributed to the membership free of charge. Also published are
Maryland Herpetofauna Leaflets and these are available at $. 25/page.
Information for Authors
All correspondence should be addressed to the Executive Editor. Manu¬
scripts being submitted for publication should be typewritten (double spaced)
on good quality 8 1/2 by 11 inch paper with adequate margins. Submit origi¬
nal and first carbon, retaining the second carbon. If entered on a word proces¬
sor, also submit diskette and note word processor and operating system used.
Indicate where illustrations or photographs are to appear in text. Cite all lit¬
erature used at end in alphabetical order by author.
Major papers are those over five pages (double spaced, elite type) and
must include an abstract. The authors name should be centered under the title,
and the address is to follow the Literature Cited. Minor papers are those pa¬
pers with fewer than five pages. Author’s name is to be placed at end of paper
(see recent issue). For additional information see Style Manual for Biological
Journals (1964), American Institute of Biological Sciences, 3900 Wisconsin
Avenue, N.W., Washington, D.C. 20016.
Reprints are available at $.07 a page and should be ordered when manu¬
scripts are submitted or when proofs are returned. Minimum order is 100
reprints. Either edited manuscript or proof will be returned to author for ap¬
proval or correction. The author will be responsible for all corrections to proof,
and must return proof preferably within seven days.
The Maryland Herpetological Society
Department of Herpetology
Natural History Society of Maryland, Inc.
2643 North Charles Street
Baltimore, Maryland 21218
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Bulletin of the Maryland Herpetological Society
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