Journal of Herpetology, Vol. 46, No.

4, 473479, 2012
Copyright 2012 Society for the Study of Amphibians and Reptiles

A New Golden Toad (Bufonidae: Incilius) from Northwestern Guatemala and

Chiapas, Mexico
JOSEPH R. MENDELSON III,1,2,3 DANIEL G. MULCAHY,4 SARA SNELL,2,5 MANUEL E. ACEVEDO,6

AND JONATHAN

A. CAMPBELL7

Department of Herpetology, Zoo Atlanta, 800 Cherokee Avenue Southeast, Atlanta, Georgia 30315 USA
2
School of Biology, Georgia Institute of Technology, 301 Ferst Drive., Atlanta, Georgia 30332 USA
4
National Museum of Natural History, Smithsonian Institution, MRC 162, P.O. Box 37012, Washington, DC 20013-7012 USA; E-mail: mulcahyd@si.edu
6
Escuela de Biologia, Universidad de San Carlos de Guatemala, Ciudad Universitaria, Zona 12, Guatemala, Guatemala; E-mail: manuelaceved@gmail.com
7
Department of Biology, University of Texas at Arlington, Arlington, Texas 76019 USA; E-mail: campbell@uta.edu

ABSTRACT.We describe Incilius aurarius sp. nov., a new species of toad known from several localities on the humid Caribbean slopes of the
Sierra de los Cuchumatanes in Huehuetenango, Guatemala, and adjacent highlands of Chiapas, Mexico. This species previously has been
confused with populations of Incilius valliceps and Incilius macrocristatus. The new species is morphologically similar to the Mexican species I.
macrocristatus but differs by having less prominent crests, a distinctive golden coloration in the males, and the absence of vocal slits.

The taxonomic status of Bufo valliceps macrocristatus Firschein

and Smith, 1957 was reviewed by Mendelson (1997) who
recognized the taxon as a full species occurring in the
Chimalapas region of Chiapas and Oaxaca, Mexico, the Selva
Negra region of Northern Highlands of Chiapas, Mexico, the
Atlantic slope of the northwestern portion of the Sierra de las
Cuchumatanes, Huehuetenango, Guatemala, and the nearby
Lagos de Montebello area of Chiapas, Mexico. This is a rarely
encountered species in the Forest Toad clade recognized by
Mendelson et al. (2011) that is represented by few specimens,
many of which are juveniles or poorly preserved. Since
Mendelsons (1997) review, additional specimens and photographs from Huehuetenango, Guatemala, have been collected,
indicating that this population differs markedly from Incilius
macrocristatus (sensu stricto). Based on these differences, here we
describe the toad as a new species of Incilius.
MATERIALS

AND

METHODS

General terminology and measurements follow those of

Mendelson (1997). Foot-webbing formulae follow the system
of Savage and Heyer (1967), as modified by Myers and
Duellman (1982) and Savage and Heyer (1997). The general
format of the description and diagnosis is formatted similar to
that of Mendelson et al. (2005). Specimens examined are listed in
(Appendix I). Museum abbreviations are those proposed by
Leviton et al. (1985).
Incilius aurarius sp. nov.
Figures 13
Suggested Standard English name: Cuchumatan Golden Toad
Bufo macrocristatus.Flores-Villel\a, 1993 (in part; for records
representing Tropical Highlands of Chiapas, Mexico, and
Guatemala); Mendelson, 1997 (in part; for specimens from
southeastern Chiapas, Mexico, and Sierra de los Cuchumatanes,
Guatemala).
Bufo valliceps.Flores-Villela, 1993 (in part; for records
representing Tropical Highlands of Chiapas, Mexico, and
Guatemala).
3
5

Corresponding Author. E-mail: j.mendelson@zooatlanta.org

E-mail: sara.snell@gatech.edu

DOI: 10.1670/11-140

Holotype.UTA A-52591 (original number MEA 1391), an

adult male from Guatemala: Huehuetenango: Nenton,

Aldea
Yalambojoch, Ro Sancapech, Finca San Francisco, 1,270 m
(approximately 15858 0 07.16 00 N, 91833 0 45.90 00 W; 1,270 m) collected
by Manuel E. Acevedo on 25 May 1998.
Paratypes.UTA A-52597, adult female with locality data same
as holotype. CAS 163782, adult female, CAS 163783, adult male,
both from Mexico: Chiapas: 4.8 km (3 miles) east of Lago Tsikoa,
Montebello National Park (elevation not recorded), collected by
D. E. Breedlove on 14 May 1973.
Referred Specimens.See Appendix 1.
Diagnosis.The new species is allocated to the genus Incilius
(sensu Frost et al., 2009) by the presence of a full complement of
cranial crests and the presence of a descending row of enlarged
lateral tubercles on the body. A moderately sized species of
Incilius (males to 67.5 mm SVL; females to 79.5 mm SVL), having
the following combination of characters: (1) tympanum evident,
round, smaller in diameter than the orbit; (2) canthal, supraorbital, supratympanic, postorbital, parietal, pretympanic, supralabial, preorbital crests present in females; all crests present in
males, except pretympanic absent, preorbital and supralabial
being reduced; (3) cranial crests moderately developed in males,
lacking rounded protuberance at junction of parietal, supraorbital, and postorbital crests, females bearing hypertrophied, thin
crests rising well above level of orbit; (4) tibia long, about 42%
SVL; (5) feet long, about 44% SVL; (6) dorsal skin of females
mostly smooth with scattered subequal rounded tubercles,
becoming more concentrated posteriorly, and distinctly spiculate
on flanks and limbs; dorsal skin of males granular with densely
concentrated subequal rounded turbercles, becoming moderately
spiculate on flanks and limbs; (7) ventral skin in females with
densely concentrated small distinctly spiculate tubercles; ventral
skin in males uniformly granular; (8) descending row of enlarged
lateral tubercles in females appear as sharply defined row of
distinctly spiculate tubercles, in males these appear as moderately defined row of slightly enlarged weakly pointed tubercles;
(9) vocal slits absent; (10) snout shape bluntly rounded in lateral
view, sharply pointed in dorsal view; (11) parotoid glands
elongate, subtriangular; (12) skin between cranial crests on top of
head in females revealing striated texture of underlying bone,
smooth in males; (13) dorsal coloration uniformly pale golden in
males, pale or dark brown in females with variable dark brown
or black markings.

474

J. R. MENDELSON ET AL.

FIG. 1. Incilius aurarius in life. (A, B) Male holotype (UTA A-59591), SVL = 67.5 mm; (C) adult female from Guatemala: Huehuetenango: San Jose
Maxbal photographed by T. Pierson on 9 April 2011, SVL unknown; (D) adult female paratype (UTA A-52597), SVL = 79.5 mm, visible tympanum is
perforated; (E) Ventral aspect of female paratype (left) and male holotype (right); (F) juvenile female (UTA A-52599), SVL = 32.4; (G) tadpole at Gosner
Stage 32 (UTA A-61033); (H) tadpole at Gosner Stage 44 (UTA A-61032). Color reproduction supported by the Thomas Beauvais Fund.

NEW GOLDEN TOAD (INCILIUS) FROM GUATEMALA AND MEXICO

475

FIG. 2. Preserved specimens of Incilius aurarius, females top and males situated below. Top left: adult female UTA A-52597 (SVL = 79.5 mm); top
right: adult female (MVZ 143380 (SVL = 67.5 mm); bottom left: adult male MVZ 143365 (SVL = 54.5 mm); bottom right: adult male holotype UTA A52591 (SVL = 67.5 mm). Color reproduction supported by the Thomas Beauvais Fund.

Incilius aurarius shows marked sexual dimorphism; however,

this species can be distinguished from most species of Incilius by
the unique uniform golden coloration of the males, whereas the
females show a more plesiomorphic pale or dark brown
coloration with variable dark brown or black markings (e.g., a
pattern popularly referred to as dead leaf pattern). Females
additionally have enlarged cranial crests raised well above level
of the orbit, whereas those in the males are lower. This species is
most similar to I. macrocristatus, but it differs by having the
distinctive golden coloration in males, lacking dark brown or
black markings including on the legs and feet (dull brown or

yellowish-tan, usually with dark brown or black markings

especially on the legs and feet in I. macrocristatus), and by
having smaller cranial crests in the males, and lacking a distinct
bony knob at the junction of the parietal, supraorbital, and
postorbital crests in both males and females (forming a distinct
bony knob at the junction of the parietal, supraorbital, and
postorbital crests in I. macrocristatus); I. aurarius further differs
from I. macrocristatus by lacking vocal slits (present, small,
bilateral in I. macrocristatus).
Incilius aurarius differs from Incilius campbelli, also occurring
in Guatemala, by having the distinct golden coloration in males

476

J. R. MENDELSON ET AL.

FIG. 3. Comparison of cranial crests between adult males and females of Incilius aurarius and Incilius macrocristatus. (A) Female of I. aurarius (UTA
A-52597); (B) female of I. macrocristatus (MVZ 99521); (C) male of I. aurarius (UTA A-52591, holotype); (D) male of I. macrocristatus (UMMZ 123994).
Note the bony knob at the junction of the parietal, postoborbital, and orbital crests is more developed in I. macrocristatus (B, D) than in I. aurarius (A,C);
this distinction is more evident in females than in males.

(mostly brown in I. campbelli) and by having distinctly enlarged

crests, especially in the females (low, thin crests in both sexes in
I. campbelli); but note that a population referable to I. campbelli
(D. G. Mulcahy and J. R. Mendelson, unpubl. data) in the Maya
Mountains of Belize may have relatively large crests. Males of I.
luetkenii, known from the subhumid lowlands in Guatemala, are
yellowish-green but differ from I. aurarius by having reduced
parietal crests and small rounded parotoid glands (vs. prominent parietal crests and elongate parotoid glands); additionally,
females of I. luetkenii are nearly uniform dull brown, lacking the
distinctive dead leaf pattern described herein.
Males of the Golden Toad of Costa Rica (Incilius periglenes) are
more golden orange than the golden yellow coloration of I.
aurarius, whereas the females of I. periglenes bear a distinctive
brilliant coloration consisting of bold yellow, black, and red
markings that is in no way similar to a dead leaf pattern;
furthermore, the cranial crests of I. periglenes are greatly
reduced, lacking the preorbital and parietal crests entirely.
Color images of both Incilius luetkenii and I. periglenes were
presented by Savage (2002).
Description of the Holotype.Body robust; head wider than
long, width 37.5% SVL, length 33% SVL; snout sharply pointed in
dorsal view, sharply pointed in profile, rostral keel absent;
canthal, supraorbital, parietal, pretympanic, supratympanic, and
postorbital crests present; all cranial crests well developed except
pretympanic crest; junction of parietal postorbital, and supraorbital crests not forming distinct knob; skin on top of head coossified; nostril protuberant, directed dorsally; canthus rostralis

forming distinct, raised, canthal ridge; loreal region concave; lip

indistinct; suborbital ridge present, indistinct, extending from
angle of jaw anteriorly to level of anterior margin of orbit; notch
at symphysis of upper jaw present, distinct; eyenostril distance
about 110% of diameter of orbit; tympanum distinct, round;
tympanic annulus distinct, dorsal margin obscured by supratympanic crest. Forelimbs short, moderate; hand thin, with long,
thin fingers; relative length of fingers: II < IV < I < III; webbing
absent; tips of fingers not expanded, smooth dorsally; palmar,
pollical, subarticular, and supernumerary tubercles present,
distinct; palmar tubercle ovoid, approximately twice size of
pollical tubercle; nuptial excrescences discrete, brown granular
patch covering most of dorsal surface of Finger I, present but less
developed on dorsal surface of Finger II. Hind limbs relatively
long, slender, tibia length 44.5% SVL; foot length 43.2% SVL;
tarsal fold absent; outer metatarsal tubercle small, ovoid, slightly
smaller than inner metatarsal tubercle; toes long, thin. Relative
length of toes: I < II < V < III < IV; webbing thin, webbing
formula I21II21III22IV1.51V; tips of toes not expanded,
smooth dorsally, tips of Toes IVV demarcated distally by distinct
dermal fold on dorsal surface; subarticular tubercles present,
small, round; supernumerary tubercles present, very small.
Skin on dorsum of body granular with scattered, slightly
larger, low, rounded tubercles; larger dorsal tubercles bearing
small clusters of finely keratinized points; parotoid glands
distinct, ovoid; descending row of enlarged lateral tubercles
distinct, comprising slightly enlarged, weakly pointed tubercles;
dorsal surface of head between cranial crests smooth; dorsal

NEW GOLDEN TOAD (INCILIUS) FROM GUATEMALA AND MEXICO

477

TABLE 1. Morphometric variation in Incilius aurarius and Incilius macrocristatus. Mean 6 SD above, range in parentheses below; all measurements in
millimeters.
Incilius aurarius
Males, N = 5

Variable

Snoutvent length
Tibia length
Foot length
Head length
Head width
Orbit diameter
Tympanum diameter
Supratympanic crest length
Parotoid gland length

60.0
26.2
27.4
21.5
22.5
8.1
3.5
4.8
8.7

6
6
6
6
6
6
6
6
6

5.0
1.8
3.0
2.2
1.9
0.4
0.5
0.7
1.4

(54.567.5)
(24.228.8)
(24.331.6)
(18.424.0)
(19.424.1)
(7.78.8)
(3.14.4)
(3.65.5)
(7.611.0)

Incilius macrocristatus
Females, N = 5

65.1
27.3
28.2
24.5
26.5
8.7
3.7
5.8
10.5

6
6
6
6
6
6
6
6
6

surfaces of limbs covered with densely arranged weakly

pointed tubercles of various sizes; skin on throat and venter
uniformly granular.
Choanae large, transversely elliptical, widely spaced; teeth
and odontoids absent; tongue long, posterior sixth of length
free; vocal slits absent.
Measurements of the Holotype (in Millimeters).SVL 67.5, head
length 24.0, head width 24.1, tibia length 28.8, foot length 31.6,
orbit diameter 8.8, tympanum diameter 4.4, supratympanic crest
length 5.5.
Coloration of Holotype (Alcohol after Formalin).Uniform dull
brown, dorsal surface and limbs grey-brown; ventral surfaces
dull cream with a distinct dark brown mottling posterior to
pectoral girdle.
Coloration in Life.Males are a uniform golden yellow, with the
color becoming paler ventrally (Fig. 1). This color is not observed
in specimens stored in alcohol. All dorsal surfaces are golden
yellow, and cranial crests are dark brown. The ventral surface is
cream with indistinct grey mottling. The iris is copper with fine
black flecking.
Females are brown dorsally, becoming grey laterally. They
exhibit a lateral row of enlarged tubercles that are grey, and the
lateral surfaces below the tubercles are distinctly dark brown.
The suborbital area is cream becoming dark brown over the
tympanum. The dorsal surfaces of the limbs are grey-brown
with indistinct brown markings, and the tips of the digits are
distinctly yellow. The ventral surfaces are dark brown with
distinct cream mottling becoming more pronounced posteriorly
and on the ventral surfaces of the legs. There is a pale grey
middorsal stripe and black interorbital bar. The iris is dark
brown with indistinct copper flecking.
Variation.Morphometric variation is summarized in Table 1.
Variation in color pattern and sexual dimorphism, especially of
the cranial crests, are evident in Figures 13.
Etymology.The specific epithet aurarius is Latin meaning
golden and is used in reference to the distinctive coloration of
the male of this species.
Distribution and Ecology.This species is known from a few
localities in adjacent regions of Huehuetenango, Guatemala, and
Chiapas, Mexico, on the humid Caribbean versant at elevations
between 1,100m and 1,798 m (Fig. 4). The few observations of this
species in the wild, and its phylogenetic placement among the socalled Forest Toads (Mulcahy and Mendelson, 2000; Mendelson
et al., 2011; as I. sp. nov.), suggest that this species is associated
closely with primary cloud-forest habitat and likely intolerant of
even minor alterations to such forest (see Mendelson et al., 1999).
The species is known from within the Lagos de Montebello
National Park, in Mexico, where its habitat receives a certain level

10.4 (53.179.5)
4.9 (20.934.4)
4.5 (22.134.6)
4.8 (20.132.0)
5.1 (21.333.6)
1.3 (7.410.8)
0.8 (2.84.9)
1.5 (4.27.9)
1.9 (9.013.8)

Males, n =20

59.4
25.8
26.9
20.7
21.9
7.6
3.2
4.3
8.9

6
6
6
6
6
6
6
6
6

3.8
1.8
1.5
1.5
1.7
0.9
0.3
0.6
1.4

(51.968.7)
(21.630.1)
(24.829.9)
(18.925.8)
(19.727.0)
(4.38.8)
(2.63.7)
(3.15.8)
(4.710.1)

Females, N = 9

70.5 6
30.4 6
30.2 6
25.66
27.3 6
9.6 6
4.0 6
6.2 6
8.0 6

9.5
3.6
5.4
3.3
3.6
1.6
0.6
0.8
3.4

(50.181.8)
(23.735.4)
(21.041.0)
(19.930.4)
(20.832.5)
(7.212.6)
(2.85.1)
(5.27.4)
(4.013.0)

of protection. Otherwise, its cloud-forest habitat is under constant

assault by agricultural practices, both commercial and subsistence-level.
The type-series was collected from a remnant patch (approximately 10 ha) of gallery cloud forest at 1,270 m, along the edges
of Ro Sancapech. The series of specimens collected along with
the holotype (UTA A-525915600) was collected during daylight
hours along the edges of Ro Sancapech, with the juveniles all
being found among rocks at the edge of the stream itself, the
adult female active in leaf litter near a small pool alongside the
stream, and the male holotype discovered inactive underneath
fallen wood about 10 m from the stream. The forest floor was
covered with a thick layer of leaf litter with abundant ferns and
trees in the area that were covered with abundant bromeliads
and epiphytes. The forest patch was surrounded by heavily
deforested areas and cattle pastures with abundance of rocky
outcrops and some shaded coffee plantations. Several scattered
liquidambar trees (Liquidambar styraciflua) were still present. The
discovery of early stage tadpoles of this species (described
below) in May, prior to the onset of the rainy season is indicative
of breeding during the dry season, a feature shared by other

FIG. 4. A generalized map of the highlands of Guatemala and

Chiapas, Mexico, showing the known localities for Incilius aurarius
(square symbols). The stippled pattern represents elevations 8002,000
m, and the striped pattern represents elevations >2,000 m in elevation.

478

J. R. MENDELSON ET AL.

species in the Forest toad clade (Mendelson et al., 1999). The

tadpoles were active during the day on the river bottom
consisting of gravel and sand; they were observed to be active
during the day, and none were observed at night. Other species
of amphibians and reptiles found at the type locality are
indicative of regional cloud-forest fauna (e.g., Anolis campbelli,
Ptychohyla dendrophasma, Bothriechis aurifer, and Xenosaurus
rackhami).
At the time of collection (May 1998), this was the last patch of
forest in the area, and was subject to a heavy pressure by locals
for firewood and to establish new plots for corn and coffee
crops. It is very likely that this forest patch no longer exists,
because at the time, numerous recently felled trees were
observed indicating that the patch was actively being reduced
in size. Access to this region is limited because there is a history
of social and political unrest there, including a massacre at Finca
San Francisco during the Guatemalan civil war in the 1980s that
displaced many thousands of Maya in the region. Residents of
this region frequently are hostile toward foreigners, and it is
known that several scientists have been attacked by locals in the

area (J. Monzon,

pers. com.) limiting abilities to verify the
continued existence of the species there. However, an adult
female was observed and photographed near Barillas, Huehuetenango, in April 2011 (Fig 1C, by T. Pierson; identification
verified by JRM).
Tadpoles.Description is based on four tadpoles (Gosner stages
3244; Gosner 1960; Fig. 1). Body ovoid in dorsal view, widest at
point just posterior to eyes, wider than tall, depressed in lateral
view; neuromasts not visible; snout nearly semicircular in dorsal
profile, rounded in lateral profile; eyes moderate, not part of
dorsal profile, directed dorsolaterally, separated by distance
about three times eye diameter; nostrils large, directed dorsolaterally, closer to eye than tip of snout. Spiracle sinistral, short,
opening near midbody slightly below midline, directed posteriorly, lateral wall shorter than medial wall, walls forming round
aperture. Vent tube medial, ventral, and medial margins
extended, producing the appearance of a long, dextral vent tube.
Caudal musculature highest at base, gradually tapering to a
pointed tip. Caudal fin moderately developed, extending to base
of caudal musculature, tip rounded; dorsal and ventral fins of
equivalent size, highest just posterior to midlength.
Oral disc small, anteroventral in position, width about 1.5
times distance between eyes, emarginated, with single row of
small, truncate papillae laterally; lateral marginal papillae
separated by anterior and posterior gaps; small series of
submarginal papillae at lateral tips of A-1 and A-2. Rectangular
depression present posterior to oral disc. Labial tooth row
formula 2(2)/3; A-1 longest, slightly longer than all other tooth
rows; P-1, P-2, and P-3 approximately equal in length. Upper
jaw medium, jaw sheath finely serrate, weakly convex medially,
lateral processes tapering abruptly dorsolaterally; lower jaw
narrow, width slightly less than width of upper jaw, jaw sheath
finely serrate, shallowly V-shaped.
In preservative (buffered formalin), dorsal and lateral surfaces
of body uniform reddish-brown; ventral surfaces transparent
with fine reddish-brown punctuations; gut visible ventrally, not
visible laterally. Caudal musculature uniform reddish-brown;
caudal fins transparent with fine brown reticulations on dorsal
fin. The tadpole of I. aurarius essentially is similar and
indistinguishable from that of I. macrocristatus (as described by
Korky and Webb, 1973; see also comments on that tadpole in
Mendelson, 1997, and Mendelson et al., 1999).

Remarks.Specimen UIMNH 11308 (MX: Chiapas: Palenque

ruins), a designated paratype of B. v. macrocristatus Firschein and
Smith, 1957, actually represents Incilius valliceps; this re-identification was mentioned by Mendelson (1997) but is here repeated to
avoid confusion by future scholars studying Incilius in the region.
The samples included in the phylogenetic analyses by Mendelson
et al. (2011) to represent I. macrocristatus are from near the type
locality for that taxon and so do actually represent I. macrocristatus
sensu stricto. Similarly, studies by Pramuk (2006), Pramuk et al.
(2008), and Van Bocxlaer et al. (2010) that included the taxon I.
macrocristatus were based on tissue sampled from UTA A-13014,
which we here refer to I. macrocristatus sensu stricto. The tadpoles
considered by Mendelson et al. (1999) also represent I. macrocristatus sensu stricto. Incilius aurarius is referred to as Incilius sp.
nov. throughout the text of Mendelson et al. (2011).
We propose that the conservation status of I. aurarius be listed
as Critically Endangered. We base this proposition upon
consideration of the small geographic distribution of the species,
its apparent intolerance of habitat disturbance, and the
enormous human pressures on the remnant cloud-forest habitat
along this GuatemalaMexico border region. It is a reasonable
assumption that the amphibian chytrid fungus (Batrachochytrium dendrobatidis) occurs in the region (Mendelson et al., 2004;
Cheng et al., 2011), but the susceptibility of I. aurarius to the
disease chytridiomycosis has not been evaluated.
Acknowledgments.We are grateful to all of the field
companions who ventured into the Sierra de los Cuchumatanes
at one time or another with JAC or MEA: E. N. Smith, E. D.
Brodie Jr., R. A. Nussbaum, M. V. Centeno, M. Sasa, V.
McKenzie, and I. M. Asmundsson. A. Thompson prepared the
photographs in Figure 2. P. Walker provided information
regarding I. campbelli in the Maya Mountains of Belize, and T.
Pierson allowed permission to reproduce his photograph in
Figure 1C. This paper is based on work supported by the
National Science Foundation (DEB-9705277, 0613802, 0102383)
to JAC.

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Accepted: 2 December 2011.

479

APPENDIX 1
Specimens Examined
Alda YalamIncilius aurarius.Guatemala: Huehuetenango: Nenton,
bojoch, Ro Sancapech, Finca San Francisco, 1,270 m (UTA A-52591

52560,A- 6103235 [larvae]); Nenton,

camino YalambojochYolnajab,
1,100m (UTA A-61036 [larvae]); on ridge about 2 km northwest of
Barillas, 5,900 ft [1,798 m] (MVZ 143380); city of Barillas (MVZ 143365);
Santa Cruz Barillas, 1,650 m (KU 104115 [larvae]); Municipio Barillas,
San Jose Maxbal, 1,483 m (photo voucher of uncataloged specimen at
MVZ).
Mexico: Chiapas: 9 km E Lago de Montebello, on road to Santa Elena
(CAS 13987071); Dos Lagos, 4 km east of Laguna Tziskao, in Lagos de
Montebello National Park, 4,500 ft [1,372 m] (CAS 16384445); 3 mi [4.8
km] east of Lago Tsikoa, Montebello National Park (CAS 16378289);
1012 km below Lago Tziscao, 4,000 ft [1,219 m] (CAS 16390).
Incilius macrocristatus.Mexico: Chiapas: El Mercadito, Cintalapa
(CAS 1007677); Ruins of Palenque (UIMNH 11309, 11311); ridge
between Pantepec and Tapalapa, 5,800 ft [1,768 m] (CAS 163936);

Municipio Villacorzo, between Agronimos

Mexicanos and Cerro Tres
Picos, about 4,000 ft [1,219 m] (CAS 170161); Mahosik, Tenejapa, 20 mi
[32.2 km] northeast of San Cristobal (MVZ 99521); Mahosik, Tenejapa, 18
mi [29.0 km] northeast of San Cristobal (MVZ 99522); Municipio

Tenejapa, Paraje Mahosik (CAS 16331415); 5.6 km south of Rayon

Mescalapa,
Mescalapa, 1,680 m (KU 5829499); 6.2 km south of Rayon
Mezcala (UTA A-2785759
1,690 m (KU 58300-03); 6.8 km east of Rayon
larvae); 6 mi [9.7 km] south of Soluschiapa, 1,300 ft [396 m] (UTEP 5879
5,500 ft [1,676 m] (UTEP 5885); 1.1 mi
84); 4 mi [6.4 km] south of Rayon,
[1.8 km] north of Ixtahuacan, 1,100 ft [336 m] (UTEP 9523); 3.5 mi [5.6
(UMMZ 126248); 11.3 mi [18.2 km] northwest of
km] south of Rayon
Pueblo Nuevo Solistahuacan, 5,000 ft [1,524 m] (KU 75201); 2 mi [3.2
km] west of Agua Escondida, 2,850 ft [869 m] (KU 41576); 16.1 km
northwest of Pueblo Nuevo Solistahuacan (UTA A-13014); 18.6 road km
north of Pueblo Nuevo, 1,560 m (UMMZ 123994, 4 specimens); 19.020.8
mi [30.233.5 km] north by MX Hwy 195 of Jitotol, 5,500 ft [1,676 m]
(MVZ 13893237, CAS 142614); 6.8 km north of Rayon Mezcala, 1,652 m
(UTA A-2785759 [larvae]). Mexico: Oaxaca: between La Gloria and
Cerro Azul (UIMNH 35583); mountains between La Gloria and Juchitan
(UIMNH 3358486); La Gloria (UIMNH 40995); San Isidro La Gringa,
Santa Mara Chimalapa (MZFC-EPR 37, MZFC-LCM 258); Chalchijapa,
Santa Mara Chimalapa (MZFC-EPR 67, 70, 107, MZFC-LCM 281).

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