Journal of Herpetology, Vol. 46, No. 4, 473479, 2012Copyright 2012 Society for the Study of Amphibians and Reptiles

A New Golden Toad (Bufonidae: Incilius) from Northwestern Guatemala andChiapas, Mexico

JOSEPH R. MENDELSON III,1,2,3 DANIEL G. MULCAHY,4 SARA SNELL,2,5 MANUEL E. ACEVEDO,6 AND JONATHAN A. CAMPBELL7

1Department of Herpetology, Zoo Atlanta, 800 Cherokee Avenue Southeast, Atlanta, Georgia 30315 USA2School of Biology, Georgia Institute of Technology, 301 Ferst Drive., Atlanta, Georgia 30332 USA

4National Museum of Natural History, Smithsonian Institution, MRC 162, P.O. Box 37012, Washington, DC 20013-7012 USA; E-mail: mulcahyd@si.edu6Escuela de Biologia, Universidad de San Carlos de Guatemala, Ciudad Universitaria, Zona 12, Guatemala, Guatemala; E-mail: manuelaceved@gmail.com

7Department 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 theSierra 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 Firscheinand Smith, 1957 was reviewed by Mendelson (1997) who

recognized the taxon as a full species occurring in theChimalapas region of Chiapas and Oaxaca, Mexico, the SelvaNegra region of Northern Highlands of Chiapas, Mexico, theAtlantic slope of the northwestern portion of the Sierra de lasCuchumatanes, Huehuetenango, Guatemala, and the nearbyLagos de Montebello area of Chiapas, Mexico. This is a rarely

encountered species in the Forest Toad clade recognized byMendelson et al. (2011) that is represented by few specimens,many of which are juveniles or poorly preserved. SinceMendelsons (1997) review, additional specimens and photo-graphs from Huehuetenango, Guatemala, have been collected,indicating that this population differs markedly from Inciliusmacrocristatus (sensu stricto). Based on these differences, here wedescribe the toad as a new species of Incilius.

MATERIALS AND METHODS

General terminology and measurements follow those ofMendelson (1997). Foot-webbing formulae follow the systemof Savage and Heyer (1967), as modified by Myers andDuellman (1982) and Savage and Heyer (1997). The general

format of the description and diagnosis is formatted similar tothat of Mendelson et al. (2005). Specimens examined are listed in(Appendix I). Museum abbreviations are those proposed byLeviton 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 recordsrepresenting Tropical Highlands of Chiapas, Mexico, andGuatemala); Mendelson, 1997 (in part; for specimens fromsoutheastern Chiapas, Mexico, and Sierra de los Cuchumatanes,

Guatemala).

Bufo valliceps.Flores-Villela, 1993 (in part; for recordsrepresenting Tropical Highlands of Chiapas, Mexico, andGuatemala).

Holotype.UTA A-52591 (original number MEA 1391), anadult male from Guatemala: Huehuetenango: Nenton, Aldea

Yalambojoch, Ro Sancapech, Finca San Francisco, 1,270 m

(approximately 15858007.16 00N, 91833045.90 00W; 1,270 m) collectedby Manuel E. Acevedo on 25 May 1998.

Paratypes.UTA A-52597, adult female with locality data sameas 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), havingthe following combination of characters: (1) tympanum evident,

round, smaller in diameter than the orbit; (2) canthal, supraor-

bital, supratympanic, postorbital, parietal, pretympanic, supra-

labial, 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, supraor-

bital, 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 moder-

ately 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.

3Corresponding Author. E-mail: j.mendelson@zooatlanta.org5E-mail: sara.snell@gatech.eduDOI: 10.1670/11-140

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

474 J. R. MENDELSON ET AL.

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

FIG. 2. Preserved specimens of Incilius aurarius, females top and males situated below. Top left: adult female UTA A-52597 (SVL = 79.5 mm); topright: adult female (MVZ 143380 (SVL = 67.5 mm); bottom left: adult male MVZ 143365 (SVL = 54.5 mm); bottom right: adult male holotype UTA A-52591 (SVL = 67.5 mm). Color reproduction supported by the Thomas Beauvais Fund.

NEW GOLDEN TOAD (INCILIUS) FROM GUATEMALA AND MEXICO 475

(mostly brown in I. campbelli) and by having distinctly enlargedcrests, especially in the females (low, thin crests in both sexes inI. 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, areyellowish-green but differ from I. aurarius by having reducedparietal crests and small rounded parotoid glands (vs. promi-nent parietal crests and elongate parotoid glands); additionally,females of I. luetkenii are nearly uniform dull brown, lacking thedistinctive dead leaf pattern described herein.

Males of the Golden Toad of Costa Rica (Incilius periglenes) aremore golden orange than the golden yellow coloration of I.aurarius, whereas the females of I. periglenes bear a distinctivebrilliant coloration consisting of bold yellow, black, and redmarkings that is in no way similar to a dead leaf pattern;furthermore, the cranial crests of I. periglenes are greatlyreduced, lacking the preorbital and parietal crests entirely.Color images of both Incilius luetkenii and I. periglenes werepresented by Savage (2002).

Description of the Holotype.Body robust; head wider thanlong, width 37.5% SVL, length 33% SVL; snout sharply pointed indorsal view, sharply pointed in profile, rostral keel absent;

canthal, supraorbital, parietal, pretympanic, supratympanic, andpostorbital crests present; all cranial crests well developed exceptpretympanic crest; junction of parietal postorbital, and supraor-bital crests not forming distinct knob; skin on top of head co-

ossified; nostril protuberant, directed dorsally; canthus rostralis

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

indistinct; suborbital ridge present, indistinct, extending fromangle of jaw anteriorly to level of anterior margin of orbit; notch

at symphysis of upper jaw present, distinct; eyenostril distanceabout 110% of diameter of orbit; tympanum distinct, round;tympanic annulus distinct, dorsal margin obscured by supra-

tympanic crest. Forelimbs short, moderate; hand thin, with long,thin fingers; relative length of fingers: II < IV < I < III; webbingabsent; tips of fingers not expanded, smooth dorsally; palmar,pollical, subarticular, and supernumerary tubercles present,

distinct; palmar tubercle ovoid, approximately twice size ofpollical tubercle; nuptial excrescences discrete, brown granularpatch covering most of dorsal surface of Finger I, present but less

developed on dorsal surface of Finger II. Hind limbs relativelylong, slender, tibia length 44.5% SVL; foot length 43.2% SVL;

tarsal fold absent; outer metatarsal tubercle small, ovoid, slightlysmaller than inner metatarsal tubercle; toes long, thin. Relative

length of toes: I < II < V < III < IV; webbing thin, webbingformula 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, slightlylarger, low, rounded tubercles; larger dorsal tubercles bearing

small clusters of finely keratinized points; parotoid glandsdistinct, ovoid; descending row of enlarged lateral tuberclesdistinct, comprising slightly enlarged, weakly pointed tubercles;

dorsal surface of head between cranial crests smooth; dorsal

FIG. 3. Comparison of cranial crests between adult males and females of Incilius aurarius and Incilius macrocristatus. (A) Female of I. aurarius (UTAA-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.

476 J. R. MENDELSON ET AL.

surfaces of limbs covered with densely arranged weaklypointed tubercles of various sizes; skin on throat and venteruniformly granular.

Choanae large, transversely elliptical, widely spaced; teethand odontoids absent; tongue long, posterior sixth of lengthfree; vocal slits absent.

Measurements of the Holotype (in Millimeters).SVL 67.5, headlength 24.0, head width 24.1, tibia length 28.8, foot length 31.6,orbit diameter 8.8, tympanum diameter 4.4, supratympanic crestlength 5.5.

Coloration of Holotype (Alcohol after Formalin).Uniform dullbrown, dorsal surface and limbs grey-brown; ventral surfacesdull cream with a distinct dark brown mottling posterior topectoral girdle.

Coloration in Life.Males are a uniform golden yellow, with thecolor becoming paler ventrally (Fig. 1). This color is not observedin specimens stored in alcohol. All dorsal surfaces are goldenyellow, and cranial crests are dark brown. The ventral surface iscream with indistinct grey mottling. The iris is copper with fineblack flecking.

Females are brown dorsally, becoming grey laterally. Theyexhibit a lateral row of enlarged tubercles that are grey, and thelateral surfaces below the tubercles are distinctly dark brown.The suborbital area is cream becoming dark brown over thetympanum. The dorsal surfaces of the limbs are grey-brownwith indistinct brown markings, and the tips of the digits aredistinctly yellow. The ventral surfaces are dark brown withdistinct cream mottling becoming more pronounced posteriorlyand on the ventral surfaces of the legs. There is a pale greymiddorsal stripe and black interorbital bar. The iris is darkbrown with indistinct copper flecking.

Variation.Morphometric variation is summarized in Table 1.Variation in color pattern and sexual dimorphism, especially ofthe cranial crests, are evident in Figures 13.

Etymology.The specific epithet aurarius is Latin meaninggolden and is used in reference to the distinctive coloration ofthe male of this species.

Distribution and Ecology.This species is known from a fewlocalities in adjacent regions of Huehuetenango, Guatemala, andChiapas, Mexico, on the humid Caribbean versant at elevationsbetween 1,100m and 1,798 m (Fig. 4). The few observations of thisspecies in the wild, and its phylogenetic placement among the so-called Forest Toads (Mulcahy and Mendelson, 2000; Mendelsonet al., 2011; as I. sp. nov.), suggest that this species is associatedclosely with primary cloud-forest habitat and likely intolerant ofeven minor alterations to such forest (see Mendelson et al., 1999).The species is known from within the Lagos de MontebelloNational Park, in Mexico, where its habitat receives a certain level

of protection. Otherwise, its cloud-forest habitat is under constantassault by agricultural practices, both commercial and subsis-

tence-level.

The type-series was collected from a remnant patch (approx-

imately 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 allbeing 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 andtrees 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. Thediscovery 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

TABLE 1. Morphometric variation in Incilius aurarius and Incilius macrocristatus. Mean 6 SD above, range in parentheses below; all measurements inmillimeters.

Variable

Incilius aurarius Incilius macrocristatus

Males, N = 5 Females, N = 5 Males, n =20 Females, N = 9

Snoutvent length 60.0 6 5.0 (54.567.5) 65.1 6 10.4 (53.179.5) 59.4 6 3.8 (51.968.7) 70.5 6 9.5 (50.181.8)Tibia length 26.2 6 1.8 (24.228.8) 27.3 6 4.9 (20.934.4) 25.8 6 1.8 (21.630.1) 30.4 6 3.6 (23.735.4)Foot length 27.4 6 3.0 (24.331.6) 28.2 6 4.5 (22.134.6) 26.9 6 1.5 (24.829.9) 30.2 6 5.4 (21.041.0)Head length 21.5 6 2.2 (18.424.0) 24.5 6 4.8 (20.132.0) 20.7 6 1.5 (18.925.8) 25.66 3.3 (19.930.4)Head width 22.5 6 1.9 (19.424.1) 26.5 6 5.1 (21.333.6) 21.9 6 1.7 (19.727.0) 27.3 6 3.6 (20.832.5)Orbit diameter 8.1 6 0.4 (7.78.8) 8.7 6 1.3 (7.410.8) 7.6 6 0.9 (4.38.8) 9.6 6 1.6 (7.212.6)Tympanum diameter 3.5 6 0.5 (3.14.4) 3.7 6 0.8 (2.84.9) 3.2 6 0.3 (2.63.7) 4.0 6 0.6 (2.85.1)Supratympanic crest length 4.8 6 0.7 (3.65.5) 5.8 6 1.5 (4.27.9) 4.3 6 0.6 (3.15.8) 6.2 6 0.8 (5.27.4)Parotoid gland length 8.7 6 1.4 (7.611.0) 10.5 6 1.9 (9.013.8) 8.9 6 1.4 (4.710.1) 8.0 6 3.4 (4.013.0)

FIG. 4. A generalized map of the highlands of Guatemala andChiapas, Mexico, showing the known localities for Incilius aurarius(square symbols). The stippled pattern represents elevations 8002,000m, and the striped pattern represents elevations >2,000 m in elevation.

NEW GOLDEN TOAD (INCILIUS) FROM GUATEMALA AND MEXICO 477

species in the Forest toad clade (Mendelson et al., 1999). Thetadpoles were active during the day on the river bottomconsisting of gravel and sand; they were observed to be activeduring the day, and none were observed at night. Other speciesof amphibians and reptiles found at the type locality areindicative of regional cloud-forest fauna (e.g., Anolis campbelli,Ptychohyla dendrophasma, Bothriechis aurifer, and Xenosaurusrackhami).

At the time of collection (May 1998), this was the last patch offorest in the area, and was subject to a heavy pressure by localsfor firewood and to establish new plots for corn and coffeecrops. It is very likely that this forest patch no longer exists,because at the time, numerous recently felled trees wereobserved indicating that the patch was actively being reducedin size. Access to this region is limited because there is a historyof social and political unrest there, including a massacre at FincaSan Francisco during the Guatemalan civil war in the 1980s thatdisplaced many thousands of Maya in the region. Residents ofthis region frequently are hostile toward foreigners, and it isknown that several scientists have been attacked by locals in thearea (J. Monzon, pers. com.) limiting abilities to verify thecontinued existence of the species there. However, an adultfemale was observed and photographed near Barillas, Huehue-tenango, in April 2011 (Fig 1C, by T. Pierson; identificationverified by JRM).

Tadpoles.Description is based on four tadpoles (Gosner stages3244; Gosner 1960; Fig. 1). Body ovoid in dorsal view, widest atpoint just posterior to eyes, wider than tall, depressed in lateralview; neuromasts not visible; snout nearly semicircular in dorsalprofile, rounded in lateral profile; eyes moderate, not part ofdorsal profile, directed dorsolaterally, separated by distanceabout three times eye diameter; nostrils large, directed dorsolat-erally, closer to eye than tip of snout. Spiracle sinistral, short,opening near midbody slightly below midline, directed posteri-orly, lateral wall shorter than medial wall, walls forming roundaperture. Vent tube medial, ventral, and medial marginsextended, producing the appearance of a long, dextral vent tube.Caudal musculature highest at base, gradually tapering to apointed tip. Caudal fin moderately developed, extending to baseof caudal musculature, tip rounded; dorsal and ventral fins ofequivalent size, highest just posterior to midlength.

Oral disc small, anteroventral in position, width about 1.5times distance between eyes, emarginated, with single row ofsmall, truncate papillae laterally; lateral marginal papillaeseparated by anterior and posterior gaps; small series ofsubmarginal papillae at lateral tips of A-1 and A-2. Rectangulardepression present posterior to oral disc. Labial tooth rowformula 2(2)/3; A-1 longest, slightly longer than all other toothrows; P-1, P-2, and P-3 approximately equal in length. Upperjaw medium, jaw sheath finely serrate, weakly convex medially,lateral processes tapering abruptly dorsolaterally; lower jawnarrow, width slightly less than width of upper jaw, jaw sheathfinely serrate, shallowly V-shaped.

In preservative (buffered formalin), dorsal and lateral surfacesof body uniform reddish-brown; ventral surfaces transparentwith fine reddish-brown punctuations; gut visible ventrally, notvisible laterally. Caudal musculature uniform reddish-brown;caudal fins transparent with fine brown reticulations on dorsalfin. The tadpole of I. aurarius essentially is similar andindistinguishable from that of I. macrocristatus (as described byKorky and Webb, 1973; see also comments on that tadpole inMendelson, 1997, and Mendelson et al., 1999).

Remarks.Specimen UIMNH 11308 (MX: Chiapas: Palenqueruins), a designated paratype of B. v. macrocristatus Firschein andSmith, 1957, actually represents Incilius valliceps; this re-identifi-cation was mentioned by Mendelson (1997) but is here repeated toavoid confusion by future scholars studying Incilius in the region.The samples included in the phylogenetic analyses by Mendelsonet al. (2011) to represent I. macrocristatus are from near the typelocality for that taxon and so do actually represent I. macrocristatussensu 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 tadpolesconsidered by Mendelson et al. (1999) also represent I. macro-cristatus 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 listedas Critically Endangered. We base this proposition uponconsideration of the small geographic distribution of the species,its apparent intolerance of habitat disturbance, and theenormous human pressures on the remnant cloud-forest habitatalong this GuatemalaMexico border region. It is a reasonableassumption that the amphibian chytrid fungus (Batrachochy-trium dendrobatidis) occurs in the region (Mendelson et al., 2004;Cheng et al., 2011), but the susceptibility of I. aurarius to thedisease chytridiomycosis has not been evaluated.

Acknowledgments.We are grateful to all of the fieldcompanions who ventured into the Sierra de los Cuchumatanesat 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 thephotographs in Figure 2. P. Walker provided informationregarding I. campbelli in the Maya Mountains of Belize, and T.Pierson allowed permission to reproduce his photograph inFigure 1C. This paper is based on work supported by theNational Science Foundation (DEB-9705277, 0613802, 0102383)to JAC.

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Accepted: 2 December 2011.

APPENDIX 1

Specimens Examined

Incilius aurarius.Guatemala: Huehuetenango: Nenton, Alda Yalam-bojoch, Ro Sancapech, Finca San Francisco, 1,270 m (UTA A-5259152560,A- 6103235 [larvae]); Nenton, camino YalambojochYolnajab,1,100m (UTA A-61036 [larvae]); on ridge about 2 km northwest ofBarillas, 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 atMVZ).

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 deMontebello National Park, 4,500 ft [1,372 m] (CAS 16384445); 3 mi [4.8km] 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); ridgebetween Pantepec and Tapalapa, 5,800 ft [1,768 m] (CAS 163936);Municipio Villacorzo, between Agronimos Mexicanos and Cerro TresPicos, about 4,000 ft [1,219 m] (CAS 170161); Mahosik, Tenejapa, 20 mi[32.2 km] northeast of San Cristobal (MVZ 99521); Mahosik, Tenejapa, 18mi [29.0 km] northeast of San Cristobal (MVZ 99522); MunicipioTenejapa, Paraje Mahosik (CAS 16331415); 5.6 km south of RayonMescalapa, 1,680 m (KU 5829499); 6.2 km south of Rayon Mescalapa,1,690 m (KU 58300-03); 6.8 km east of Rayon Mezcala (UTA A-2785759larvae); 6 mi [9.7 km] south of Soluschiapa, 1,300 ft [396 m] (UTEP 587984); 4 mi [6.4 km] south of Rayon, 5,500 ft [1,676 m] (UTEP 5885); 1.1 mi[1.8 km] north of Ixtahuacan, 1,100 ft [336 m] (UTEP 9523); 3.5 mi [5.6km] south of Rayon (UMMZ 126248); 11.3 mi [18.2 km] northwest ofPueblo Nuevo Solistahuacan, 5,000 ft [1,524 m] (KU 75201); 2 mi [3.2km] west of Agua Escondida, 2,850 ft [869 m] (KU 41576); 16.1 kmnorthwest of Pueblo Nuevo Solistahuacan (UTA A-13014); 18.6 road kmnorth of Pueblo Nuevo, 1,560 m (UMMZ 123994, 4 specimens); 19.020.8mi [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 andCerro 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).

NEW GOLDEN TOAD (INCILIUS) FROM GUATEMALA AND MEXICO 479

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