September 1982]

361

HERPETOLOGICA

Two
Abronia aurita (1) GUATEMALA:BajaVerapaz:E
slope Cerro Quisis, 1829 m, KU 190851.
new lizards(generaAbroniaand Xenosaurus)from
the Los TuxtlasRange of Veracruz,Mexico. Trans. Abronia chiszari (1-holotype) MEXICO: Veracruz: 2.5 mi E Cuetzalapan, 360 m, UTA R-3195
KansasAcad. Sci. 64:123-132.
(that this is the exact provenance is doubtful).
Accepted: 3 March 1982
Abronia deppei (7) MEXICO:Guerrero:Omilteme,
Associate Editor: Stephen Tilley
2134 m, UTA R-4451, 5553, 5645-46, 5653-54; 1.6
km N Puerto del Gallo, 2621 m, UTA R-4151.
Museum of Natural History and De- Abroniafuscolabialis
(1) MEXICO:Oaxaca:5.8 km
partment of Systematics and Ecology,
W Totontepec, 2103 m, UTA R-9899.
The University of Kansas, Lawrence, KS Abronia lythrochila (1) MEXICO: Chiapas: 8 km
NW San Crist6bal de las Casas, 2385 m, UTA
66045, USA
R-3354.
Abronia mixteca (11) MEXICO:Oaxaca:Tejocotes,
2286 m, UTA R-5786,5790, 6126, 6228, 6825, 6827,
APPENDIX I
6989, 7737, 7825, 10277; 2377 m, KU 106303
In preparingthe descriptionof A. mitchelli, I have
(paratype).
had the benefit of examining the following speci- Abronia t. taeniata (2) MEXICO:Hidalgo: 3 km W
mens housed in the University of Kansas(KU) and
Xochicoatlan, KU 54055; Rio Chinameca, 8 km N
the University of Texas at Arlington (UTA) collecTianguistengo, 1020 m, KU 101144.
tions. The collecting localities and elevations (when Abronia t. graminea (5) MEXICO:Veracruz:3 km
available) are given.
W Acultzingo, KU 26484-87, 26827.
WERLER, J. E., AND F. A. SHANON. 1961.

Herpetologica, 38(3), 1982, 361-366

? 1982 by The Herpetologists' League, Inc.

A NEW SPECIES OF THE GENUS LEPIDOPHYMA

(SAURIA: XANTUSIIDAE) FROM MICHOACAN, MEXICO
ROBERT

L. BEZY, ROBERT G. WEBB, AND TICUL ALVAREZ

ABSTRACT: A new species of Lepidophyma is described from Michoacan, 250 km northwest

of the previously known northernlimit of the genus on the Pacific coast of Mexico. The species
resembles L. smithii in having low numbers of femoral pores and lateral tubercle rows, but
differs in having fewer dorsal and gular scales.

Key words: Reptilia; Squamata;Xantusiidae; Lepidophyma; Systematics; Mexico

THREE specimens of a species of Lepidophyma were collected at a locality in
Michoac'anwhich lies 250 km northwest
of the previously known northern limit of
the genus along the Pacific coast of Mexico. Comparisons with over 1200 specimens of all nominal taxa of the genus indicate that the material from Michoacian
represents a previously undescribed
species.
The specimens are deposited in the
collections of the Escuela Nacional de
Ciencias Biologicas, Instituto Politecnico
Nacional (ENCB) and of the Natural History Museum of Los Angeles County

(LACM). The nomenclature used in this

paper follows Bezy (1972, 1973, and unpublished); the terminology for scalation
is largely that of Savage (1963). Comparisons were made with topotypic material
of all nominal taxa (Bezy, 1973; Smith,
1973), except L. lipetzi (Smith and Alvarez del Toro, 1977) for which the original
description was used in conjunction with
specimens judged to represent the
species.
Lepidophyma tarascae

sp. nov.

Holotype.-ENCB 9221 (original field

number A.O. 6193) (Fig. 1), an adult male

362

HERPETOLOGICA

[Vol. 38, No. 3

CB 9221

FIG. 1.-Dorsal

and ventral views of the holotype (ENCB 9221) of Lepidophyma tarascae, new species.

obtained near Mexiquillo, Aquila District, Michoacan, Mexico, by Aurelio

Ocafia, on 10 July 1976. Mexiquillo
(18008'N, 103056'W) is a small town on
the coast of Michoacan, ca. 77 km (air)
WNW of L'azaro Cardenas and the mouth
of the Rio Balsas.
Paratypes.-Two,
ENCB 9222 (A.O.
6194) and LACM 134226 (formerly ENCB
9223, A.O. 6195), same locality, collector
and date as holotype.
Diagnosis.-The
hypodigm of L. tarascae has 16-18 total (both legs) femoral
pores; 16-17 well-developed lateral rows
of tubercles (axilla to groin) (Fig. 2); 2-3
dorsal interwhorls (separating large tail
whorls); four scales (total both sides) separating postocular from second postorbital supralabial; paravertebral rows heterogeneous, each with 16 large paravertebral tubercles (axilla to groin), usually
separated within-row by 1 intermediatesized tubercle and 0-1 small scales, and
between-rows by 2-3 mid-dorsal scales
(Fig. 2); 41-43 gulars (fold to second infralabials); 145-150 mid-dorsal scales (occiput to rump); and a distinct row of large

tubercles on the lateral nuchal area extending from the posteroventral tympanic margin to above the gular fold (Fig. 3).
L. tarascae differs from all other species
of the genus (except some L. smithii) in
having a distinct lateral nuchal row of
large tubercles; from all except L. smithii
in having less than 20 lateral tubercle
rows (16-17 vs. 20-46); and from all

except L. smithii and L. occulor in

having less than 20 total femoral pores
(16-18 vs. 20-44). In addition, L. tarascae differs from L. smithii and L. occulor
in having fewer dorsal scales (145-150 vs.
164-224 and 213-242, respectively) and
fewer gulars (41-43 vs. 44-59 and 59-71,
respectively).

Description of holotype.-Measurements (in mm): Snout-vent length, 93; tail

length, 110 (of which 52 is regenerated);
head length, 22.1; head width 15.1; head
depth, 11.3; orbit length, 3.9; fourth toe

length, 12,1.
The nasal and anterior labial regions
are slightly damaged on both sides of the
head, the integument partially missing,
exposing maxillae and teeth. The de-

September 1982]

HERPETOLOGICA

363

I-

FIG. 3.-Side of head of holotype (upper, ENCB

9221) of Lepidophyma tarascae, new species,
showing lateral nuchal row of large tubercles; and
FIG. 2.-Lateral (upper) and paravertebral(lower) of L. smithii (lower, LACM 130027, 1 km NW Puertubercle rows of the holotype (ENCB 9221) of Lep- to Marquez, Guerrero, M6xico).
idophyma tarascae, new species.

scription of paratype ENCB 9222 is sub-

stituted [in brackets], when a character is

not clear on the holotype. Nasals in contact posterior to rostral, followed by median frontonasal, two prefrontals (no median prefrontal), and two frontals (medial
suture of the frontals longer than that of
prefrontals); median interparietal (wider
anteriorly than posteriorly) touching both
frontals anteriorly and postparietals posteriorly, and separating lateral parietals;
position of parietal organ not discernible;
postparietals lacking anomalous sutures.
Nostril bordered by nasal, postnasal, and
first supralabial; postnasal followed by
anterior loreal (ca. same height as postnasal, its contact with frontonasal narrow,
especially on right side), and posterior loreal (slightly more than twice size of anterior loreal); very narrow vertically linear preocular. More or less complete
orbital ring of tiny scales; loreolabial (or
frenocular, between lower posterior part
of posterior loreal and fifth supralabial) in
broad contact with [fourth] supralabial;
three postoculars, lower largest, triangular; two upper postoculars, both touching
parietals, uppermost touching frontals.

Postoculars followed by two anterior

temporals (upper larger), very large median temporal (slightly larger than parietal), and large posterior temporal (ca.
one-fourth size of postparietal); single row
of two small scales separating large median temporal and seventh supralabial;
postocular separated from seventh supralabial by anterior lower temporal plus one
small scale; pretympanic area with four
(right)/three (left) enlarged scales (arranged in vertical row) followed by five/
six large auriculars. [Seven supralabials]
(integument of first [four] lacking), fifth
below eye, seventh smaller than sixth.
Large mental followed by four pairs of
infralabials, with first three pairs large and
fourth pair smallest (ca. one-fifth size of
third pair), and first two pairs having broad
common median sutures; gular scales
small, 41 along midline between fold and
second pair of infralabials; row of 14/15
large tubercles (_ three times size of adjacent scales) extending from posteroventral margin of tympanum to above gular fold.
Dorsal and lateral surfaces of body covered by small granules or scales of varying sizes, with interspersion of numerous

364

HERPETOLOGICA

enlarged, keeled tubercles. Paravertebral

rows of tubercles heterogeneous in size,
composed of large keeled tubercles
( -three times size of adjacent mid-dorsal
scales), each usually followed by one
small scale and one intermediate-sized
keeled tubercle (ca. twice size of dorsal
scales); 16 large tubercles in each paravertebral row between axilla and groin.
Vertebral area of uniformly small granules, three between paravertebral rows,
145 along vertebral line between occiput
and rump. Large tubercles on sides of
body arranged in vertical rows with six in
each row at midbody; 17 vertical rows between axilla and groin; two (mostly) or
three vertical rows of small granules
(midbody) between vertical rows of tubercles; dorsolateral areas of uniformly
small granules, three between paravertebral rows and dorsalmost tubercles of
lateral vertical rows (at midbody).
Ventral scales flat, mostly smooth,
quadrangular, in 10 longitudinal rows at
midbody; lateralmost row of ventrals elevated, keeled, each ca. two-thirds size
of adjacent ventrals, and not reaching axillary region; 35 transverse rows of ventral scales (several intercalary rows) from
gular fold to vent, including three rows
of preanals; two scales in first and second
rows of preanals, and four scales in last
(posteriormost) row; lateralmost preanals
of last row ca. one-fourth size of adjacent
medial preanals. Scales on surfaces of
limbs heterogenous in size, mostly
keeled; dorsal surface of hindlimbs with
scattered large keeled tubercles; 8/10
femoral pores; 23/22 fourth toe lamellae,
2/3 divided.
Tail encircled with whorls of large tubercular keeled scales, separated by two
or three interwhorls of smaller keeled
scales; 12 segments on unregenerated part
of tail; third (posteriormost)interwhorl of
each segment largest, ca. two-thirds
length of whorl; first (anteriormost)interwhorl never complete ventrally, consisting of small separated scales in first (proximalmost) segment, increasing in size in
posterior segments to form a dorsally

[Vol. 38, No. 3

complete interwhorl of 16 scales (in segment 12) subequal to those of the following (second) interwhorl.
Color dark brown on all dorsal surfaces; small white spots arrangedin paravertebral rows (just below paravertebral
row of tubercles), fading on posterior twothirds of body; dorsolateral row of very
small, faint spots on neck, fading posterior to axilla; lateral nuchal tubercle row
dirty white. Top and sides of head mostly
uniform brown (paler than body); lower
jaw dark brown with white bars on sutures between large scales (mental, infralabials); ventral surfaces dirty white,
individual scales white with brown pigment concentrated on anterior portion;
brown pigment least on gulars, forming
dark crescent on anterior one-third of
ventral body scales, and covering all but
posterior keels of ventral caudal scales.
Variation.-Sex,
6, c, Y (for ENCB
9221, LACM 134226, ENCB 9222, in that
order); snout-vent length (mm), 93, 66,
65; tail length/snout-vent length, unknown, 1.48, 1.45; total femoral pores, 18,
17, 16; lateral tubercle rows (axilla to
groin), 17, 16, 17; scales betwreen paravertebral rows, 3.0, 3.0, 2.0; total dorsal
caudal interwhorls (between first six
whorls), 11 (all); total complete ventral
interwhorls (between first six whorls),
10, 7, 10; total dorsal caudal annuli
(whorls + interwhorls), unknown, 127,
123; median prefrontals, 0 (all); scales
(total both sides) separating postocular
from second postorbital supralabial, 4
(all); gulars, 41, 43, 43; ventrals (gular to
vent), 35, 34, 34; ventrals across midbody, 10 (all); keeled ventrals across midbody, 2, 6, 6; fourth toe lamellae, 23, 22,
23; divided fourth toe lamellae, 2, 1, 2;
dorsal scales (occiput to rump), 145, 150,
145; dorsal scales in row above paravertebral tubercles (axilla to groin), 74, 74,
77; total (both sides) supralabials (anterior to second postorbital labial), unknown, 14, 12; second infralabial contact,
1, 0, 0; gulars contacting first infralabial,
0, 1, 1; anomalous postparietal sutures, 0
(all); large paravertebral tubercles sepa-

HERPETOLOGICA

September 1982]

rated within-row by a distance equal to

2.5 dorsal scales (all); scales in one paravertebral row (axilla to groin), (a) total,
45, 46, 42, (b) smaller than 1.5 dorsal
scales, 15, 15, 11, (c) larger than 1.5 dorsal
scales, 30, 31, 31, (d) larger than 2.0 dorsal scales, 24, 23, 28, (e) larger than 3.0
dorsal scales, 16 (all); width of posterolateral preanal/width of posteromedial
preanal, 0.41, 0.65, 0.71; height of postnasal/height of anterior loreal, 1.02, 0.99,
0.94; height of second postorbital supralabial/height of first postorbital supralabial, 0.83, 0.69, 0.74; length of frontal/
length of postparietal, 0.74, 0.86, 0.88;
length of postocular/length of orbit, 0.27,
0.43, 0.19; length of nasal/length of prefrontal, 0.45, 0.69, 0.64; prefrontal, length
along midline/length along lateral border, 0.49, 0.30, 0.29; interparietal, posterior width/anteriorwidth, 0.78, 0.84,0.96.
Distribution and ecology.-The three
specimens of L. tarascae were caught in
museum special traps set in a rocky ravine in tropical semi-deciduous forest.
Additional occurrences of the species may
be anticipated along the Pacific flank of
the Sierra de Coalcom'an from the Rio
Balsas to the Rio Coahuayana.
Etymology.-The species is named for
the Tarascan people whose vast empire
once extended from the Mesa Central to
the coast of Michoac'an (Stanislawski,
1947).
DISCUSSION

L. tarascae most closely resembles L.

smithii in having fewer femoral pores,
lateral vertical rows of tubercles, and dorsal caudal interwhorls than other species
in the genus. The lateral tubercles are
large and well-aligned into a relatively
few vertical rows. The paravertebral tubercles are concomitantly large and tend
to encroach upon the small scales in the
row, resulting in short runs of tubercles
(particularly in ENCB 9222) that are
nearly as juxtaposed as those of L. tuxtlae
and L. pajapanensis. In the latter two
species, the rows consist of uninterrupted juxtaposed tubercles of a relatively ho-

365

mogeneous intermediate-size, while in L.

tarascae, they contain 16 large tubercles
(' three times dorsal scales), 14-15 intermediate-sized tubercles (1.5-2.9 times
dorsal scales) and 11-15 small scales (0.51.4 times dorsal scales). A similar pattern
exists for the caudal scales of L. tarascae.
The large size of the whorls is associated
with a reduction in number of dorsal interwhorls, approaching the two that are
characteristically present in L. gaigeae,
L. radula and L. dontomasi. In these latter species, the whorls are relatively undifferentiated from the interwhorls, while
in L. tarascae they differ strikingly in size
and keel development.
The population of L. tarascae occurs
250 km northwest of the range of Lepidophyma smithii, which extends on the
Pacific versant from Guerrero (Mautz and
Lopez-Forment, 1978) to El Salvador.
Only three specimens of L. tarascae are
currently available, but these have been
compared with data from 264 L. smithii.
The degree of divergence in scalation between the two taxa is approximately
equivalent to that between L. tuxtlae and
L. pajapanensis, species that are sympatric in the Los Tuxtlas region of Veracruz.
Differences in scalation between the several disjunct populations (including three
named races) of L. smithii (Bezy, unpublished) are considerably less than between it and L. tarascae. For example, L.
tarascae is approximately as divergent in
dorsal scale number (x = 146.7, n = 3)

from L. smithii of the northernmost population in Guerrero (i = 186.6, SE = 1.51,

n = 11) as from those to the south on the

Isthmus of Tehuantepec (x = 189.0, SE =
2.03, n = 44). These data suggest that it
is unlikely that additional material from
the area between the ranges of L. smithii
and L. tarascae will bridge the morphological gap between the two species.
Acknowledgments.-We thank Sr. Aurelio Ocaiia
and Dr. Diaz Pardo for collecting the type series,
Dr. Carl Lieb and Mr. MartinRuggles for field assistance, Mr.John De Leon for photographyand Dr.
John W. Wright for reviewing the manuscript.

HERPETOLOGICA

366

[Vol. 38, No. `3

SMITH, H. M., AND M. ALVAREZDEL TORO. 1977.

LITERATURE CITED
BEZY, R. L. 1972. Karyotypicvariation and evo-

A new troglodytic lizard (Reptilia, Lacertilia,

Xantusiidae) from Mexico. J. Herpetol. 11:37-40.

lution of the lizards in the family Xantusiidae. STANISLAWSKI,D. 1947. Tarascan political geogContrib. Sci. 227:1-29.
raphy. Amer. Anthropol.49:46-55.
1973. A new species of the genus LepiAccepted: 22 February 1982
dophyma (Reptilia: Xantusiidae) from GuatemaAssociate Editor: Stephen Tilley
la. Contrib. Sci. 239:1-7.
1978.
MAUrZ, W. J., AND W. LOPEZ-FORMENT.
RLB: Section of Herpetology, Natural
Observations on the activity and diet of the cavMuseum of Los Angeles County,
History
ernicolous lizard Lepidophyma smithii (Sauria:
Los Angeles, CA 90007, USA; RGW: DeXantusiidae). Herpetologica 34:311-313.
SAVAGE, J. M. 1963. Studies on the lizard family
partment of Biological Sciences, UniverXantusiidaeIV. The genera. Los Angeles Co. Mus. sity of Texas at El Paso, El Paso, TX
Contrib. Sci. 71:1-38.
79968, USA; TA: Escuela Nacional de
SMITH, H. M. 1973. A tentative rearrangementof
the lizards of the genus Lepidophyma. J. Her- Ciencias Biolo'gicas, Instituto Polite'cnico Nacional, Me'xico17, D.F., Me'xico
petol. 7:109-123.

Herpetologica, 38(3), 1982, 366-374

? 1982 by The Herpetologists' League, Inc.

TWO NEW SPECIES OF POISON-DART FROGS

(COLOSTETHUS) FROM COLOMBIA
JOHN LYNCH
ABSTRACT: Two dendrobatids, provisionally assigned to Colostethus, are named from the
Cordillera Oriental of Colombia near Bogota, Colombia. Both species differ from other Colostethus in lacking the vocal slits and in having elongate anal sheaths. One species is cavemicolous and occurs at elevations above 3000 m (caves within paramos).The other species occurs
along mountain streams in areas once supportinghigh altitude cloud forests (elevations ca. 2400-

2800 m).

Key words: Amphibia; Salientia; Dendrobatidae; Colostethus; Colombia; High altitude;

Troglodytic; Relationships

DENDROBATID frogs of the genus Colostethus are considered the most primitive members of the poison-dart frog family (Edwards, 1974; Lynch, 1971; Noble,
1931). Edwards (1974) recognized 62
species distributed through forested (plus
paramo) environments of the Neotropics
from Costa Rica and Tobago south to Bolivia and to eastern and southeastern
Brasil. The frogs included in the genus
by Edwards have toe discs and pads, obvious scutes atop the digital pads, premaxillary/maxillarydentition, and usually drab patterns involving
pale

dorsolateral, lateral (= oblique), and/or

ventrolateralstripes. These features (most
of which are synapomorphies of the family or a more inclusive group) are combined with the absence of pumiliotoxin-C class alkaloids [(found in all
Dendrobates + Phyllobates; Myers et al.,
1978) = skin toxins auctorum] to "define" Colostethus.
In 1979, Sr. Vladimir Corredorand Dr.
Pedro M. Ruiz showed me some peculiar
frogs collected at various mountain sites
in the vicinity of Bogota by staff and students from the Instituto de Ciencias Na-