\\try .Em Scientific Papers Natural History Museum The University of Kansas 31 March 2005 N u mber 38: 1 -27 Systematics of the Bufo coccifer Complex (Anura: Bufonidae) of Mesoamerica By Joseph R. Mendelson IIP, Beck^ L. Williams-, CnKisTorHEK A. Sheil^ AND Daniel G. Mulcahv* Ubi ' Dcpaitiuoit of Hci-pctologi/, Zoo Atlanta, S(HI Cherokee Ave SE, Atlanta, GA 3U315 \'' •*^^-dc\TY -Departiiieut of Integrative Biologi/. University of California, Berkeley, C A 94720-iUO . .;;Vt-R^ ^Departine)it of Biology, jolni Carroll University, University Heights, OH 44118 ^Department of Biology, Utah State University, Logan^lT 84322-5305 CONTENTS ABSTRACT 2 RESUMEN 2 INTRODUCTION 2 Acknowledgments 5 MATERIALS AND METHODS 5 Morphological Data 5 Molecular Data 6 ACCOUNTS OF SPECIES 6 Biifo coccifer 6 Bufo cycladen 11 Bufo ibarrai 13 Bufo pisinnus 15 Bufo portcri 17 Bufo signifer 20 MORPHOMETRIC ANALYSES 22 MOLECULAR ANALYSES 23 LITERATURE CITED 24 APPENDIX 26 © Natural History Museum, Tlu' University of Kansas ElTlSt MfiVf Lf'TarV '^^^ ^'" "'''■*'"^*^2 Museum of Compara*ve Zoology Harvard Unlversitv' ] ds>^- Scientific Papers Natural History Museum The University of Kansas 31 Mcirch 2003 Number 38:1-27 Systematics of the Bufo coccifer Complex (Anura: Bufonidae) of Mesoamerica ubrary Joseph R. Mendelson III', Bhck^ L. Williams-, Christoi'hek A. Sheil\ ANL^ Daniel G. Mulcahv* - ^m ' Dcimiiiiieiit of Ucrpctolo^ii, Zoo Atlniitu, S(H) Cherokee Ave SE, Atlanta, GA j03]5 H '-'•^y^-rrY -Department ofhite^rative Biology, University of California, Berkeley, CA 94720-3140 V. ''c-^^^ ^Department of Biolo<;y, John Carroll Universiti/, University Hei;^hti, OH 4411S ^Department of Biology, Utah State Universiti/, Logan, UT S4322-5305 CONTENTS ABSTRACT 2 RESUMEN 2 INTRODUCTION 2 Al K\'OVVLEDGMENTS 5 MATERIALS AND METHODS 5 Morphological Da la 5 Molfxular Data 6 ACCOUNTS OF SPECIES 6 Biifo coccifer 6 Bufo cycladen 11 Bufo ibarrai 13 Bufo pisiinius 15 Bufo porteri 17 Bufo signifcr 20 MORPHOMETRIC ANALYSES 22 MOLECULAR ANALYSES 23 LITERATURE CITED 24 APPENDIX 26 © Natiiivil History Museum, The University of Kjnsds EmS^ MtiVf LF'TarV '^^^ ^'^' ' "^'■*""''*^2 Museum of Compara*ye Zoo'o^ Harvard Univ6rs[t>' 2 Scientific Papers, Naturai, Hisiorv Mi si,iiM,Tiii' Uni\i ksnv ui- Kansas ABSTRACT Many populations of toads occurrinj^ between west-central Mexico and Panama have been referred to Biifo coccifcr Cope, KSdd. While the taxonomic status of these populatitms has been c]uestioned for many decades, a tliori>iii;h ie\ it'w ot the B. coccifer complex never has been presented. Based on evidence from external morphology and a partial molecular data-set, we conclude that this complex consists minimally of six species. Herein, we recognize B. ivicifcr Cope, 1866, B. cychidcii Lynch and Smith, 1966, and B. ibanai Stuart, 1954, and describe three new species. Key Words: Bufonidae; Bufo coccifcr, Biifo cycladcn, Biifo ilnirrai, Biifo /'/s/mhhs, Bufo porlcri, Bitfo signifer; Mesoamerica; taxonom\'. KESUMEN Muchas poblaciones de sapos que se encuentian cntre el oeste-central de Mexico y Panama han sido referidas como Bufo coccifcr Cope 1866. Aunc]ue el estado taxoncimico de estas poblaciones ha sido cuestionado por muchas decadas, nunca se ha presentado una revision completa del complejo B. coccifcr. En base a c\ idcncia do mortologi'a external y una base de datos moleculares parcial, concluimos que este complejo consiste de al menos seis especies. En este trabajo, reconocemos B. coccifcr Cope, 1866, B. cycladcn Lynch and Smith, 19h6, y B. ilnirrai Stuart, 1934, y describimos tres especies nuevas. Palabraf Clave: Bufonidae; Bufo coccifcr, Bufo cycladcn, Bufo ibnrrai, Bufo pisnmus, Bufo portcri, Bufo signifer; Mesoamerica; taxonomfa. INTRODUCTION The taxon Bufo coccifcr Cope, 1866 currently is applied to toads that are distributed allopatrically in five regions of Mesoamerica (Pig. I): (1) the Tepalcatepec Valley, Michoacan, Mexico; (2) the Pacific slope of the Sierra Madre del Sur in Guerrero, Mexico; (3) the southern side of the Isthmus of Tehuantepec, Oaxaca, Mexico; (4) nearly the entire Pacific versant from northwestern Guatemala to northwestern Costa Rica; and (5) western Panama. Additional records exist from the Atlantic versants of Honduras and Nicaragua. The distribution of these populations approximately matches the distribution of low elevation tropical dry forest (Rzedowski, 1994; Campbell, 1999) in these regions. However, the Rica: Cope, 1866:130) and suggested that the holotype likely originated from somewhere on the Meseta Central of Costa Rica; this referral was supported by Savage (1974). Porter (1963) provided a range map and general diagnosis of B. coccifcr Porter (1965) discussed the distribution of the species in more detail, observing that the populations in Michoacan, Mexico, and Oaxaca, Mexico, evidently are allopatric — with a hiatus of approximately 140 km between records from Oaxaca, Mexico, and records from southeastern Guatemala. Stuart (1954a:20) referred to "a chain cif Bufo coccifcr-Wke toads" distributed through the subhumid habitats of Central America, and specifically noted undescribed species in central Guatemala, and population on the Pacific slope of the Sierra Madre del Sur another in Guerrero and the Isthmus of Tehuantepec is an exception to this generality; these toads are found at moderate elevations (ca. 1000 m) in relatively wet pine- oak forest. (See Campbell and Duellman, 2000, for habitat description.) Other records from more mesic, upland habitats include the mountains of central Honduras (McCranie and Wilson, 2002). An additional specie.s, B. ibarrai Stuart, frequently has been assigned to the Bufo coccifcr Group (e.g.. Frost, 1985). The taxonomic status and distribution of B. ibarrai was reviewed by Mendelson (2001); this species occurs primarily in upland pine-oak habitats in Guatemala and is reported here lor the first time in an adjacent region of Honduras. With reference to the broad distribution ot lUdo coccifcr, it has been suggested many times that these wirious populations likely are not conspi-citic (Stuart, 1954a, 1963; Duellman, 1960; Porter, 1963, 1965; Zweifel, 1965; McDiarmid .\nL\ I oster, 1981; Mendelson, 2001; McCranie and Wilson, 2002). Dunn and Stuart (1951) commented on region of Mexico. Stuart (1954b) subsequently described the populations in the upland pine-oak zone of Guatemala as B. ibarrai. Lynch and Fugler (1965) reported B. ibarrai from Honduras. Subsequently, the taxon has had a long period of uncertain status, typically being confused with B. coccifcr (e.g., comments by J. A. Campbell in Frost, 1985:41). Meyer and Wilson (1971) placed B. ibarrai in the synonymy of B. coccifcr. However, Mendelson (2001) recognized B. ibarrai as a ciistinct species, described the tadpole, and prcnided a new diagnosis and range map of records tiom Guatemala; also, he suggested that the species may be endemic to Guatemala. Based on examination of specimens from Honduras, McCranie and Wilson (2002) suggested that B. ibarrai does not occur in that country. Lynch and Smith (1966) referred all populations in Mexico to the new taxon Bufo cycladcn and cited differences in the ad\'ertisemenl call (tidi' Poitei; 1965) and ecology of the Mexican and Central American thestatusof the type locality for B. coccifcr (" hrnha" Costa populations. I lo\ve\'er, the morphologv of this t ixon was Toads oi- iiii- Bcro coccifir Comi'LLX K ilometers 100 Fij;, 1 , A m>ip of MfsoaniL'iiLM, sluuving thu genoriili/ed di^li ibutioiis of fo.ids ivtc-ned to Biifo locci/ci soiimi Kito, .ind of /x ibaiiai (as recognizud hi'iein). diagnosed poorly, with respect to B. ibnrrai (see Mendelson, 2001 ) cind other popiilatiotis referable to B. coccifcr (Porter, 1967). Porter (1%7) criticized Lynch and Smith (1966) for not supporting their claitns of timrphological differences atnong the populations atid also claimed that there are no real ecological differences among the populations considered; Porter proposed this ta\on to be a mniicii diibiiuii. The taxonotnic cofifiision surroiuidifig B. c\/cliu1cii is evident iti the fact that the taxon appears Ofi some recent checklists (e.g., Frost, 1985), but not oti others (e.g., Flores- Villela, 1993; Catnpbell, 1999). Se\eral authors have described the male ad \ertisefnent calls from various areas. Porter (1964, 1965) described the ad\'ertisement calls of B. cociifcr from the Tehuantepec region of Oaxaca, Mexico, in addition to those of toacHs he referred to B. coaifcr recorded in Guerrero, Mexico, and several populations in Central Atnerica (El Salvador, Honduras, Nicaragua, and Costa Rica; Porter, 1965). The various descriptions and comparisons of advertisement calls presented by Porter (e.g., 1964, 1965) are difficult to interpret because he provided no voucher nufiibers for recorded specimens, and the detail of his locality information is inconsistent. When tracking down specific localitv data provided by K. R. Porter, we found it useful to refer to recording station descriptions presented in his original, unpublished dissertation (Porter, 1962:table 1). Zweifel (1965) first reported the presence of toads he referred to B. coccifcr in Panama and demonstrated that the advertisement call of this population differed from those of both the Mexican (= Isthtnus of Tehuantepec region of Oaxaca) and other Central Americati recordings that were published by Porter (1964). McDiarmid and Foster (1981) described additional advertisement calls recorded in Costa Rica, and compared them to those reported from Mexico and Central America (Porter, 1965) and from Panama (Zwiefel, 1965); there are substantial differences in pulse SciENTiiic Papi:rs. Natlrai. Hisiorv MusiiUM.Tiiii UNi\i;Rsir-t oi Kansas labk' 1. Ciimporison ot i\ingi"s or moiins ot c\ills of the Hufo coccifcr ciimplex Ironi dlllcrcnt populations. Tlic identity ot the toads recorded from Honduras (Porter, 1965) are unknown because the original paper provides neither reference to voucher specimens nor a detailed localilv- description; thiTc ,ire at least three species in this group in ?londuras. .\o. individuals Duration Frequency Pulse Rate Species Location recorded (second) (Hertz) (per second) Source identity Mexico: Guerrero 1(6 calls) mean =5.7 mean =2800 mean =120 Porter, 1965 B. cycladcn Mexico: Oaxaca 18 1.4-6.4 2800-3350 97-115 Porter, 1964, 1965 B. coccifcr El Salvador 14 3.0-16.3 2200-2700 80-101 Porter, 1965 B. coccifcr Guatemala 7 4.2-5.8 1650-1800 48-61 Porter, 1966 B. ilmrrai Honduras 2 5.3-8.8 2200-2500 80-96 Porter, 1965 unknown Nicaragua 4 6.2-10.9 2300-2400 91-100 Porter, 1965 B. coccifcr Costa Rica 4 1.5-10.0 2300 90-95 Porter, 1965 B, coccifcr C\>sta Rica 1 1.2-5.7 2231-2539 90-105 McDiarmid & Foster, 1981 B. coccifcr Panama 1 8.0-17.0 2100-2600 66 Zweifel, 1965 B. sigtiifer rates and dominant frec|Liencies among these samples. Porter (1966) described the ad\'ertisement call of B. ibiinui. The data reported bv I^orter, Zweifel, and McDiarmid and Foster are presented in Table 1 and summarized in Figure 2. We are unaware of any recordings of populations of 6. cf. coccifcr from Guatemala or Michoacan, Mexico. The calls of male toads from the Sierra Madre del Sur of Guerrero and the Tehuantepec region of Oaxaca have higher dominant frequencies than calls of all other samples. The call of the toad in Guerrero also has a higher pulse rate than all other reported calls. Note, however, that the data from Guerrero are based on a recording of six separate calls by the same individual. These data were used, in part, by Lynch and Smith (1966) to justify recognition of all Mexican populations (including populations in Michoacan, Guerrero, and Oaxaca) as a species (B. cyclndcii) distinct Dominant Frequency Guerrero Oaxaca Central America Panama B. ibarrai 1600 2500 Hz ~1 3400 Pulse Rate Guerrero Oaxaca Central America Panama B. ibarrai 40 I 80 pulse/sec 120 Fig. 2. Cirapliical representation ol Irequencies ami pulse rate's ot advertisement calls Ironi toads ul Ihe Biito coccifcr complex Ironi dilterent regions of Mesoamerica; information shown here are a graphical representation of data presented m l.ible I IXila from "Cenlral America" here includes range of values representing samples from El Salvador, Nicaragua, and Costa Rica. Toads of thi-; Bufo cocciii-r Comi'i k.\ expressed or implied consensus of these authors is that a thorough revision of the Biifo coccifcr complex is long overdue, in tliis paper we use data from external morphology and male advertisement calls (previously published) to delimit species boundaries amting samples referable to the widespread taxon B. coccifcr. We also present a preliminary assessment of phvU>genetic relationships among the species in the group, based on mtDNA data from several of the species. Acknowledgments We are grateful to the following inclividuals for providing assistance in the field, laboratory, and library: W. Duellman, S. Cotte, K. Lips, M. Acevedo, E. Greenbaum, J. R. McCranie, J. Malone, D. Laurencio, M. Sasa-Marin, M. Ryan, M. Forstner, T. Reeder, J. Campbell, E. Smith, C. Franklin, and R. CJutberlet. Adele Cutler provided statistical acivice and assistance. Michelle Koo graciously provided access and assistance with mapping software at the California Academy of Sciences. We benefited greatly from memories, observations, and notes drawn from the considerable fieki experiences of J. Campbell, C. Myers, j. Sa\'age, W. Duellman, anci R. Zweifel. Permission and help with photographs was provided by M. Sasa-Marin, R. Zweifel, and J. Simmons. Loans of critical specimens, for extended periods of time, are gratefully acknowledged from the curators and collection managers of the following institvitions: AMNH, CAS, FMNH, TCWC, UTA, TNHC, ANSP, UIMNH, USNM, KU, UMMZ, MVZ, LACM, and LSUMZ. Some of the fieldwork for this project was supported by funds from The National Geographic Society and this research was conducted in direct association with The Research Analysis Network for Neotropical Amphibians (RANA; NSF-DEB 0130273). Comments on the manuscript were provicied by the students and faculty of the USU Herpetology Group, L. frustration in delimiting species in this complex, which Trueb, J. Pramuk, and W, Duellman. Additional help and is characterized bv advertisement calls that vary among favors were provided by J. Meik and G. Schneider, populations and subtle morphological differences. An MATERIAL AND METHODS from B. coccifcr in Central America. A call described by Zweifel (1465) from Panama differs dramatically from all others b\' having a much lower pulse rate. The relationship between the calls from Oaxaca and Central America is interesting in that they have distinctly different ranges of dominant frequencies and somewhat different (but o\'erlapping) ranges of pulse rates. However, we note that the Central America data do not include recordings from Guatemala — the closest geographic region southeast of the Oaxacan population. Recordings from Guatemala may tiiminish the apparent distinctit)n between the advertisement calls from the Tehuantepec region of Oaxaca and Central America. Porter (1966) described the call of B. ibarrni from Guatemala; this species differs from all other populations in the B. coccifcr complex by having a very low frequency and pulse rate (Table 1). In their description of Bufo ci/chhicii, Lynch and Smith (1966) placed great diagnostic importance on variation in calls between Mexican and Central American populations. We note that their summary of calls from Mexico was a composite of data from both Guerrero anti the Tehuantepec region of Oaxaca (data published by Porter, 1964, 1965; Table 1 ). VVe have concluded that these toads from Guerrero are not conspecific with those from Oaxaca (discussed below). Gergus et al. (1997) described call variation among toads of the B. Diicroscnpliufi complex; they reported wide variation in some parameters and an overall pattern of apparent plesiomorphic similarity among allopatric species. There is no reason to expect drastic differences in mate-recognition systems among allopatric members of a complex of closely related anurans with similar natural histories (Gergus et al., 1997). Nevertheless, there are apparent differences in the advertisement calls of toads of the B. coccifcr complex that suggest diagnostic differences in the mate-recognition systems of these allopatric populations. All previous workers (above citations) have expressed General terminology and measurements are those of Mendelson (1997). Adult males were identified by the presence of vocal slits and nuptial excrescences; large individuals lacking these characters were presumed to be adult females. If sex could not be determined externallv, it was verified by direct observation of the gonads. 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 descriptions and diagnoses is slightly modified from that of Mendelson (2001). Museum codes are those proposed by Leviton et al. (1985). We follow Tyler et al. (2001) in OLu- usage of the term "parotoid gland." Data from advertisement calls were taken from the literatiuv (referenced below). MORPHOMI^TRIC DaIA The following measurements were taken from adult specimens: snout-vent length (SVL); head length (HL); head width (HW); tibia length (TIB); foot length (FL); width of tympanum (TYM); length of parotoid gland (PARE); maximum width of parotoid gland (PARW); and length of supratympanic crest (SPTYM). These variables represent repeatable morphological landmarks and were measured 6 Scientific Papers, Natural History Museum. The University oi- Kansas with digital calipers and iDiinded tt) the nearest 0.1 mm. Because of the paucity of large series of females from many localities, all morphometric analyses are based only on adult males. Many of the data for Biifo ibnrnii are the same presented by Mendelson (2001), but supplemented with additional material frc)m both Guatemala and Honduras. We conducteci a Principal Components Analysis (PCA) using the covariance matrix on log-transformed morphometric measurements from 324 adult male toads; this data-set included specimens representing all species and geographic regions of the distribution of the Bufo coccifcr complex. We also performed a PCA on the residuals of the latter seven variables derived from a regression analysis (using SVL as the independent variable). This type of PCA removes the effect of size, and displays general variation in shape and proportion among the specimens (Good and Wake, 1992). We performed a Linear Discriminant Function Analysis (LDA) on the same data-set, and used residuals of variables regressed on SVL and log-transformed SVL. In this analysis, a priori groupings corresponded to the species recognized in this paper (Accounts of Species, below). Statistical analyses were performecl using SAS and Minitab software. Molecular Data We conducted molecular analyses on ail available samples, which inckided five specimens of Bufo coccifer (4 from Central America, 1 from Guerrero, Mexico) and two specimens of B. ibnrrai. Additionally, one specimen of each of the following species was included as outgroup taxa: B. conifcrus, B. valliceps, and B. mnrijius. The selection of these outgroup taxa was based on preliminary analyses of a data set containing approximately 25 species of Central American bufonids (Mendelson and Mulcahy, in prep.). Sections of tiie mitochondrial genes cytochrome-b (cyt-b) and 16S were used in the molecular analyses. Isolation and PCK amplification of the mitochondrial DNA (mtDNA) genes were performed exactly as described in Mulcahy and Mendelson (2000), which includes primer information and amplification prt)files. Products from the PCR were amplified and sec]uenced in both directions using BigDye™ Terminator Cycle Set]uencing Ready Reaction Kit (Applied Biosystems Part No. 4303152); we used the same PCR primers and standard sec]uence-reaction profile on a Perkin Elmer GeneAmp 2400 cycle sequencer. Cleaned sequences were then run on an ABI 377 automated sequencer by DGM at the Biology Department at Utah State University. Sequence comparisons and alignment were conducted with Sequencher 3.1. Phylogenetic analyses of the mtDNA sequences were conducted using PAUP* 4.0b8a (Swofford, 2002). A partition-homogeneity test of 100 replicates was implemented in PAUP (using default parsimony settings, with the exception of addition-sequence being random with 100 replicates) between cyt-b and 16S gene regions to determine whether or not the two genes yielded conflicting results. Maximum-parsimony analyses (MP) were performed on each gene separately and both combined. The program Modeltest (Posada and Crandall, 1998) was used to evaluate the best Maximum-likelihood model, using the Hierarchical Likeliht)od Ratio Tests (hLRTs) criterion. A Maximum-likelihood (ML) analysis was then conducted using the model and settings based on the hLRTs results. Because of the limited number of taxa in the phylogenetic analyses, an exhaustive search was possible, and used in the MP analysis, while the ML analysis required a heuristic search algorithm. Gaps were treated as "missing data," with characters-state optimization set at ACCTRAN. Branch support was assessed bv nonparametric bootstrap analyses using 1000 replicates of full heuristic searches, with 100 random additions at each replicate, under MP and 100 replicates of full heuristic searches, with 10 random additions under the ML criteria. Decay indices (Bremer, 1994) were also measured under the parsimony analysis using the program TreeRot (Sorenson, 1996). ACCOUNTS OF SPECIES We used data from adyertisement calls, nu)rphometry, external morphology, and DNA sequences to examine variation among samples of the Bufo coccifcr complex. These data are consistent with the "chain of Bufo coccifer-Vikc toads" distributed through the subhinnid habitats of Central America that Stuart ( 1954a) envisioned. Evaluation of our data with respect to the Evolutionary Species Concept (sensu Wiley, 1978; Frost and Hillis, 1990) supports recognition of six species in this complex. Our proposed taxonomy reflects the sentiments of the many authors (e.g., Stuart, 1954a, 1963; Duellman, 1960; Porter, 1963, 1965; Zweifel, 1965; McDiarmid and Foster, 1981; Mendelson, 2001; McCranie and Wilson, 2002) who have dealt with these toads during the last five decades. We provide species accounts and diagnoses for B. coccifer, B. ci/cladcn, and B. ibnrnii, and describe three new species from this complex; photographs of these species in life are presented in Figure 3. Bufo coccifcr Cope Figs. 3-5 Bufo coccifcr Cope, 1866:00. Hiil.it\pc: USNM 6490. Type locality: "Arriba" Costa Rica. lUifo coccifcr — Dunn and Emlen, 1932; Kellogg, 19,12; Hartweg and (.liner, 1940; Dunn and Stuart, 1951; Smith and Taylor, 1948 [in part Mertens, 19.S2; Taylor, 1932; Stuart, 1954b; Rand, 1957; Duellman, 1960 I'orter, 1963 |in part); Stuart, 1963; Porter, 1964; Porter, 1965 (in part] Toads of thh Bufo coccifer Complex Fig. 3. Species of the Bi{fo coccifer complex in life, clockwise from upper left: B. coccifer from Santa Rosa, Costa Rica (adult male: photograph hy Andrea Bernecker); B. cyclndcii from Guerrero, Mexico (adult male, UTA-JRM 4607; photograph by ]. R. Mendelson); B. ibarrni from Baja Verapaz, Guatemala (female, KU 186304; photograph by J. A. Campbell); B. pifimiiif from Michoacan, Mexico (adult male, from UMMZ series; photograph bv W. E. Duellman); B. portcri from Francisco Morazan, Honduras (subadult female, KU 103220; photograph bv W. E. Duellman); B. si\;uifcr (adult female, AMNH 69625; photograph by R. G. Zweifel). Sfii:Nrii i( P,\i'i-.ks. Nvn RAi, lIisroK^ Mii.si-;um,Thk Univhrsii'i oi- Kansas Lvnch and Fuglor, 1965; Porler and Porter, 1967; Meyer and Wilson, 1971 |in part); Villa, 1972; Savage, 1974; McDiarmid and Foster, 1981; Villa, 1983; Frost, 1985 |in part]; Savage and Villa, 1986; Villa et al., 1988; |in parti; Campbell and Vannini, 1989 [in part]; Flores-Villela, 1993 |in part]; Campbell, 1999 |in part]; Kiihler, 1999; Mendelson, 2001; Frost, 21)03 |in parti; McCranie and Wilson, 2002 |in part]; Savage, 2002. Biifo cifcliuleii — Lynch and Smith, 1966 [in part, for reference to specimens from Tehuantepec region of Oaxaca, Mexico]; Frost, 1985 ]in part]; Frost, 2003 ]in part], Biifo iharrai — Lynch and I'ligler, 1965 [in part]. Bufo vatlicepf micwli^ — Werner, 1896 (synonyiiu' b\' Mendelson, 2001). Diagnosis. — A medium to large species of Bnfo {males to 62 mm SVL; females to 82 mm SVL) having the following combination of characters: (1) tympanum e\'iclent externally, about 35-45' V' diameter of orbit in males and females; (2) cantlial, supraorbital, supratvmpanic, postorbital, preorbital, pret\'mpanic, parietal, and siipralabial crests present; (3) cranial crests well dexeloped, i(.)biist, except parietal mav be thin, low, or absent in some specimens; (4) tibia short, about 35% SVL; (5) feet short, about 35% SVL; (6) dorsal tubercles small to medium sized, elevated, rounded, scattered relatively sparsely on middorsal, dorsolateral, and lateral regions of bod\' beci)ming more denseh' arranged and distinctly spinose laterally in specimens from most regions; (7) yentral tubercles granular, smooth, or with spinose apices; (8) lateral descending row of enlarged tubercles absent; (9) skin texture not sexually dinn)rphic; (U)) \ocal slit unilateral in male; (11) in. iiitcrln/oidciii^ poorly diflerentiated from ;;;. iiitciiiiaiidibiilnris, but differentiated posteriorh' forming a large, unilobed vocal sac with heavy black pigmentation; (12) snout rounded in lateral profile, pointed in dorsal aspect; (13) parotoid glands round to oxoid and large, about Fig. 4. Dorsal and ventral aspiTts ol representalise adult spi-ciniens of Hufo coctifcr from Ketalhiileu, C.iiatcniala (male, letl: UT.'\ .A-2,5821, SVL = 57.0 mm; female, right: UTA A-29025, SVL = 75.0 mm) and B. ci/chulcn Irom t.iierrero, Mexico (male, lelt: L'MM/ 1 1 5357 ] WHO 92751, SVL = 53.8 mm; female, right: UMMZ 119270 ]WKD 1.3425], SVL = 61.4 mm). ToAUs oi- iiii-; Brio coccii i:k Comi'lilX Fig. 5. L.itei mI .ispccts ot tlif hiMd'i i.)l adult nulf >pf cimens of Biifo coccifer (upper: KU hS4(l(l) .ind B. cycln'icu (lower: KU 97434). Scale bars = 1 cm. 1.0-1.5 times size of eyelid; (14) skin between cranial crests on top of head with few to many scattered, low, rounded tubercles; (15) ventral coloration whitish cream, sometimes with some degree of diffuse dark pigmentation, stunetimes in the form of diffuse punctuations. Specimens referable to Biifo coccifer vary considerably in size, shape, coloration, skin texture, and parameters of the advertisement call across the range of the species. Biifo coccifer differs from B. ci/cladeii by: being larger (males of B. cyclndcii to 54 mm SVL, females to 62 mm); having dorsal tubercles that are relatively small and scattered about the dorsal surfaces, rounded on dorsum, becoming spinose laterally in most specimens (tubercles large, elevated, and densely arranged in B. cycliiden, rounded dorsally, becoming very spinose laterally); ventral tubercles that are smoothly granular or with small, spinose apices (ventral tubercles in /-!. c\/clih1eii with distinct, large spinose apices); relati\el\' well de\eloped parietal crests that rarely are absent (parietal crest weakly de\'eloped in B. cyclndeu, sometimes absent); and an advertisement call with a lower pulse rate (Table 1; Fig. 2). Biifo coccifer can be distinguished from B. pisiumts by: being larger (males of B. yisiuiiiifi to 51 mm SVL, females to 62 mm); and having skin tubercles that are overall larger (all tubercles minute in B. pi:^iuiiiis). Biifo coccifer may be distinguished from B. signifer by having: no, iir few, relatively indistinct dark brown markings on the venter (that of B. i^igiiifcr boldly marked with a marbled pattern); relatively thicker and higher cranial crests (all crests relatively low and thin in B. signifer); and an advertisement call with a higher frec^uency and pulse rate (Table 1; Fig. 2). Biifo coccifer differs from /->. ihnrnii by: being smaller (males of B. ibarrai to S2 mm SVL, females to 94 mm); having rounded to subovoid parotoid glands about 1.0-1.5 times size of eyelid (parotoid glands distinctly ovoid, about 1.5-2.0 times size of eyelid in B. ibarrni); and males usually with dorsal tubercles becoming spincise laterally and females with rounded tubercles middorsally (dorsal tubercles of males of B. ibarrai rounded laterally and females with spinose middorsal tubercles). Bufo coccifer differs from B. porteri by having: sharply spinose lateral tubercles in males (rounded in males of 6. porteri); and a relatively thin supratympanic crest (large, bulbous in B. porteri). Morphometric variation is summarized in Table 2. Distribution and Ecology. — Bufo coccifer occurs along the Pacific versant of Mesoamerica, from the Guatemala- Mexico border to the Guanacaste region of Costa Rica (Fig. 6). Additional records exist from interior valleys on the Atlantic versant of Honduras and from the Atlantic coastal regions of Honduras and Nicaragua. McCranie and Wilson (2002) erroneously reported this species from the Atlantic versant of Guatemala; there are no records from this region. An apparently isolated population occurs in the southwestern region of the Isthmus of Tehuantepec, Oaxaca, Mexico. This species ranges from sea level to 1435 m (Savage, 2002; record from Cartago, Costa Rica) and occurs in a variety of tropical dry forests and savanna habitats. Duellman (I960) provided photographs of the habitat and some natural history observations from the Isthmus of Tehuantepec. McDiarmid and Foster (19cSI) described the reproductive biology of a population in northwestern Costa Rica. The ecological notes provided by McCranie and Wilson (2002) for Hondiu'an populations represent mixed observations pertaining to B. coccifer, B. ibarrni, and B. porteri. Savage (2002) provided a concise summary of the ecology of this species, based on observations from Costa Rica. Tadpole. — McDiarmid and Foster (1981) and Savage (2002) described tadpoles from Costa Rica and provided illustrations. 10 Scientific Papi-ks. Nau kai History MLisi-.uM.THi-. Univi-ksha oi Kansas Table 2. Morphometric variation among males and femak's from of the Biifo ccccifcr complex; mean + SD is presented above the range (in mm). Data for B. ibarrai taken, in part, from Mendelson (21)01 ). S. cocciff) B. cydadcn B. ibarrai /' yi'-nniu-^ / ■' :: <.-:lfcr Males u = 130 n ^ 311 u ^ 79 n = 2l It =43 » = 21 SVL 51.4 ±4.8 51.1 ±2.2 62.2 ±9.1 45.9 ± 3.2 49.6 ± 4.6 51.2 ±64 41.0-64.4 47.4-54.2 42.5-82.4 38.5-51.4 40.2-58.7 42.0-64.1 Tibia length 18.3 ±1.9 18.4 ±0.9 24.1 ± 3.6 15.9 ±1.4 18,4 ±1.8 17.9 ± 2.3 13.3-22.5 16.6-20.1 15.7-31.8 13.0-18.0 15.2-21.7 15.0-23.4 Foot length 18.1 ± 1.9 18.3 ±0.7 24.6 ± 3.8 16.9 ±1.3 18.8 ±1.8 18.2 ± 2.3 13.1-23.9 16.9-19.8 16.3-33.4 14.8-19.2 15.7-22.7 13.7-23.1 Head length 17.8 ± 1.8 18.0 ±0.9 20.8 ± 2.8 15.1 ± 1.0 16,7 ±1.4 174 ±2.0 13.6-21.6 16.7-20.9 15.3-26.5 12.9-17.1 14.0-20.0 14.7-22.2 Head width 20.5 ± 2.4 20.1 ±0.9 23.9 + 3.1 17.3+1.4 18.9 ±1.6 19.8 ± 24 15.9-27.8 17.6-21.8 16.2-30.8 15.1-20.2 15.7-23.0 16.6-24.9 Tympanum 2.3 ± 0.5 2.7 ± 0.3 3.7 ± 0.7 2.3 ± 0.3 3.0 ± 0.4 3.1 ± 0.4 1.3-3.7 2.1-3.3 2.7-5.3 1.8-2.6 2.2-3.5 2.5-3.8 Supratympanic 3.2 + 0.5 2.7 ± 0.3 3.5 ± 0.5 2.6 ± 0.3 3.1 ±0.3 3.0 ± 04 crest 2.0-4.4 2.2-3.3 2.3-5.2 2.2-3.2 2.3-3.7 2.1-3.8 Parotoid length 6.3 ±1.0 6.9 + 0.5 7.8+1.5 5.4 ± 0.6 6.3 ± 0.9 6.3 ±1.2 4.3-8.7 6.0-4.8 3.2-11.0 4.1-6.5 5.0-9.0 2.8-8.8 Pan>toid width 6.6 ±1.3 6.1 ±0.6 5.7 ± 0.9 5.3 ±0.5 5.1 ±0.7 6.0 ± 0.8 3.7-9.5 5.0-7.2 3.5-7.4 4.5-6.2 3.8-6.7 4.6-7.7 Females It = 25 n =6 n = 61 » = 5 // = 14 11 = 11 SVL 67.7 ± 7.6 59.8 + 2.8 78.4 ± 7.6 59.1 ± 3.3 64.5 ± 5.8 67.2 ± 8.3 58.1-82.4 54.6-62.9 60.8-94.4 55.4-62.2 56.4-73.0 54.5-77.0 Tibia length 22.2 ± 2.4 20.0 ±1.6 28.7 ± 3.0 18.7 ±1.1 77 5 + 2.2 23.0 ± 3.4 18.8-28.1 18.3-22.3 20.2-32.8 18.0-20.6 18.7-27.2 17.7-26.5 Fool length 21.9 ±2.2 20.4 ±1,0 28.7 ± 3.7 20.3 ± 1.3 22.5 ±1.9 23.1 ±3.7 17.9-27.2 19.4-22.1 19.6-34.1 18.8-21.7 19.6-26.5 17.0-26.8 Head length ?? ? ± 2.2 20.7 ±7.6 25.2 ±2.1 18.8 ±0.9 20.9 ± 2.0 22.6 ± 2.5 19.2-26.8 18.9-22.8 19.8-28.5 17.6-19.7 18.0-25.9 18.4-25.9 Head length 25.5 ±2.5 23.0 ±1.5 29.5 ± 2.8 21.9 ±1.2 23.9 ±1.9 26.1 ±34 22.5-31.5 21.1-24.4 21.5-33.3 20.6-23.7 21.5-28.3 20.9-29.4 Tympanum 3.1 ±0.6 2.8 ± 0.5 4.1 ±0.5 3.2 ±0.1 3.5 ± 0.5 3.6 ± 0.4 2.1-4.2 2.2-3.5 3.0-5.4 3.1-3.4 2.6-4.5 3.1-4.3 Supratvnipanic 4.3 ± 0.8 3.1+0.2 4.5 ± 0.7 3.4 + 0.3 3.9 ± 0.6 4.0 ± 0.7 crest 2.9-6.2 2.8-3.4 2.9-6.0 3.1-3.8 3.0-5.2 3.1-5.2 Parotoid length 7.4 + 1.3 7.6 + 0.8 8.4 ±1.5 6.8 ±1.0 8.3 ± 08 7.8 ±1.3 5.4-9.9 6.5-8.5 3.0-11.7 5.9-8.4 6.8-9.9 5.5-9.9 Parotoid width 7.4 + 1.1 6.9 ±0.8 6.3 ± 0.9 6.6+ 1.0 6.6 + 0.4 7.7 ±1.0 ■i 7 1114 6.2-8.4 4.1-8.2 ■^ (. ~ s b 1 - 2 5.4-9.0 Remarks. — Ci)k>r photographs ot this species appear in Leenders (2()01:pl. 7), Savage (20n2:pl. 77), and Villa (1972:fig. 57). Blair's (1963) reference to Bufo coccifcr in his discussion of the evolution of North American bufonieis is unclear because he does not give the provenance of the toads he analyzed. Lynch and Smith (1966) referred specimens from the Tehuantepec region of Oaxaca, Mexico to B. ci/clndcii; however, this designation was not followed by nn)st subsequent authors (Porter, 1967). Toads from the Tehuantepec region are distinct and aiiopatric, with respect to the toads we refer to B. cyclndcii on tlu' slopes ol tiie Sierra Madre del Sur in Guerrero and western Oaxaca. There are no records of Bufo coccifcr from the Pacific Coastal Plain of Chiapas, Mexico. Despite tiie lack oi records in this region, we note that much of the region seems to be suitable subiiumid habitat. Subhumid forest and scrub habitats extend along the coastal plain here, but grade into more humici forests along the base of Volcan Tacanci (Johnson, 1989; Rzedowski, 1994). Although the lone specimen from Tapachula appears to have been found in a humid region, we note that the historical humid forests (W. E. [3uellman, pers. comm.) along the coastal margin of Volcan Tacana have been completely converted to crop and pasture lands (pers. obs.). Further fieldwork along the Pacific Coastal Plain of Chiapas is warranted to \eiif\ the allopatr\' of the population of B. coccifcr in the Islhnnis ot Ti'huanatepec, with respect to conspecifics in Ciuatemala. Johnson (19S9) implied that B. coccifcr occurs along the Pacific Coastal Plain of Chiapas, perhaps based on records from Soconiisco, Chiapas, that were Toads oi nil-; Brio C(kxii ii< C()m1'L[:.\ 11 [■ij;. li- \Lip of Central AmcriLM showing the geogr.iphie distribution of Bufo coccifcr. B. ibarnii .ind B. poi Ifi i. listed by Porter (1963); these records actually represent misidentified specimens referable to 6. imlliccpf. The single specimen from Tapachula, Chiapas (MVZ 177594), is a recently metamorphosed specirnen tiiat originally was misidentified as B. luetkeiii. This record seems to be the source of information that resulted in B. luetkenibis'mg listed incorrectly as part of the Mexican herpetofauna (Johnson, 1989; Flores-Villela, 1993)— B. luctkcni does not occur in Mexico. Johnson (1990) correctly stated that records of B. coccifcr from the Grijalva Valley of Chiapas, Mexico, were based on misidentified specimens. Biifo ci/clndcii Lynch and Smith Figs. 3-5 Bufo cydaden Lynch and Smith, 1966:19. Holotype: UIMNH 57142. Type loCiiHty; 3 mi |4.8 km] S Putio de Guerrero, Oaxaea, Mexico. Bufo Li/cliidcii — Porter, 19h7 (decKired luvucii (luluuiii\; Frost, 1985 |in part]; Frost, 2003 |in part]. Bufo aicn/I-i— Smith and Ta\ lor, I94S |in part]; Porter, 196,3, 1965 |in part]; Villa et al.. 19SS |in part): Flores-Villela, 1993 |in part); Campbell, 1999 [in part]. Diagnosis. — A small species of Bufo (males to 54 mm SVL; females to 62 mm SVL), having the following combinationofcharacters:(l)tympanumevidentexternally, about 40-50% diameter of orbit in males, about 40-45% in females; (2) canthal, supraorbital, supratvmpanic, postorbital, preorbital, pretympanic, and supralabial crests present; (3) cranial crests well developed, robust, except parietal crest thin, low, or absent; (4) tibia short, about 34% SVL; (5) feet short, about 34 % SVL; (6) dorsal tubercles medium to large-sized, elevated, rounded, densely arranged on middorsal region, becoming large 12 SciENTiiic Pai'i:rs. Naturai History Mi siim.The UNiviRsin oi Kansas and conspicuously spinosc Litciallv; (7) ventral tubercles reiati\el\' large, granular, with conspicuous spinose apices; (8) lateral descending row of enlarged tubercles absent; (9) skin texture not sexually dimorphic; (10) vocal slit unilateral in male; (11) iii. intcrln/oidciifi poorly differentiated from )/;. intcniiaiidibuhnis, but differentiated posteric^rly forming a large, unilobed vocal sac with heavy black pigmentation; (12) snout shape rounded in lateral view, pointed in dorsal view; (13) parotoid glands round to ovoid and large, about 1.5-2 times size of eyelid; (14) skin between cranial crests on top of head with few to many scattered, low, rounded tubercles; (15) ventral cciloration whitish cream, with scattereci dark brown, indistinct to distinct, punctuations in some individuals. Biifo ci/tindcii may be distinguished from all other members of the Bnfo coccifcr Group by the combination of: its relatively small size; relatively large, scattered, rounded tubercles on the middorsum; large, spinose tubercles on the dorsolateral and lateral surfaces; relatively large, spinose tubercles on the venter; large parotoid glands; weakly developed (or absent) parietal crest; and presence of scattered, dark brown punctations on the venter (but this last character is variable amtmg specimens). Biifo ci/cladcn differs from B. coccifer by being smaller (males of B. coccifcr to 62 mm SVL, females to 82 mm), and possessing the following characters: large, elevated, densely arranged middorsal tubercles (middorsal tuber- cles relatively small, scattered in B. coccifcr); large, strongly spinose dorsolateral and lateral tubercles (dorsolateral and lateral tubercles smaller in B. coccifcr, variably roundeti or moderately spinose); relatively large, strongly spinose ventral tubercles (ventral tubercles small, rounded, or \sith tiny spinose apices in B. coccifcr); weakly de\'eloped, or no, parietal crests (parietal crest relativeK' well developed in most specimens of B. coccifcr); and an adverti.sement call with a higher pulse rate (Table 1, Fig. 2). Biifo cycladcn differs from B. pisinnus by having: much larger, distinctly spinose dt)rsolateral tubercles (smallei-, more rounded in B. pisinnus) and relatively larger parotoid glands (1-1.5 times si/e of eyelid in B. pisinnus). Superficially, B. cyclndcn resembles B. signifcr, but it is smaller (males of B. si^nifcr to 64 mm SVL, females to 77 mm) and has the following features: a whitish-cream venter, with or vvitht)ut scattered dark brown punctations (venter with bold brown-black marbling in /'. sis^nifcr); and smaller and less spinose tubercles overall (but this latter character is somewhat variable and subjective). Bufo cyclndoi differs from B. ibarrai bv: being smaller (males of B. ibarrai to 82 mm SVL, females to 94 mm); having males with conspicuously spinose dorsolateral and lateral tubercles; having relali\el\' larger, more densely arranged middorsal tubercles (all such tubi'rcles relatixely small, scattered, rounded in males of B. ibarrai); and having rounded middorsal tubercles in females (tubercles spinose in females of B. ibarrai). Bufo ci/cladoi differs from B. portcri by: having sharply spinose lateral tubercles in males (rounded in males of B. portcri); and relatively thin supratympanic crests (large, bulbous in B. portcri). Morphometric \ariation is summarized in Table Distribution and Ecology. — Bufo cycladcn occurs in a relati\eh' narrow ele\ational band along the Pacific slope of the Sierra Madre del Sur in Cjuerrero and Oaxaca, Mexico (Fig. 7). Most specimens ha\e been collected at, or near, Agua del Obispo, Guerrero. Althc)ugh ctimmcinly cited in the literature, Agua del Obispo does not appear on most maps because it is a private hacienda; it is located near the 1000-m contour (17°18' N, 99" 28' W) along the o\d high\s'a\' between the towns of Tierra Colorada and Chilpancingo, Guerrero. One other series (including the holotype, UIMNH 57142) is known from Putia de Guerrero, Oaxaca; this locality lies at an elevation similar to that of Agua del Obispo, Guerrero, about 150 km (airline) eastward on the same slope of the Sierra Madre del Sur. 100° 1 V k. 20° ▲ B.cycladen ^~-~~.^_^^*- • B. pisinnus ^"^^-—^ 1 < 300 m 1 300-900 m 1 1 900-2100 m ^^° WM =• 2100 m 0 100 15° Kilometers 100° I ij;. 7. Map ol LiMitiMl Mexico with dot localities tor Bufo cudadcii iTid K. pii^iiiiiu!^ (see species account lx>lou) indicating specimens ■x.iTiiined, which reprc-enls all known Imalities. Toads oi- thk Blfo coccii i:k Complex 13 Presumnblv, the distribution of this species is continuous along the middle-elevations of this slope, between these two localities. Campbell and Duellman (2000:19) described the habitat at Agua del Obispo as "...an area of scattered, small pines and brush that appears to be ecotonal between tropical deciduous forest, which is found slightly below, and pine-oak forest, which is present in the mountains above." Field work in this region by JRM and associates in 2000 and 2002 reinforced that B. ci/chulcii seems to be restricted to this narrow, ecotonal habitat. In this area, we encountered: B. Jiuiriinn^ and B. iiuiniioivtis at the lowest elevations between Acapulco and near Las Cruces; only B. manuorcits in the vicinity of Tierra Colorada (±M00 m); B. iimriinif and B. cycladcn near Agua del Obispo; only B. l>crplcxii> in tine Zumpango del Rio/Chichihualco area; and only B. occidciitalis at high elevations (over 2000 m) near Carrizal de Bravo. Da\'is and Dixon (mtiS) reported some additional records of Bufo from along this transect, but we have not verified the identifications of those specimens. The sporadic appearance of B. iiuirinii^ along this transect is noteworthv; we also note that Davis and Dixon (1965) did not report this species from anywhere in the area. Tadpole. — The tadpole of B. ci/chnicii is unknown, as are aspects of its reproductive biology. Davis and Dixon (1965) reported females collected on 22 June " . . .contained numerous small eggs." Remarks. — The type locality for this taxon is near Putla, Oaxaca, Mexico, which is located at about 750 m elevation on the Pacific slope of the Sierra Madre del Sur; specimens designated as paratypes were collected at the type locality and from around Agua del Obispt^, Guerrero, Mexico (Lynch and Smith, 1966). However, these authors referred all Mexican populations of B. coccifcr to this new taxon (thereby including populations of three different species of toads, which we have identified as B. coccifci; B. cycladcu, and B. yifiiniiis). Furthermore, it is evident from their map (Lyncli and Smith, 1966:fig. 2) that their comparative samples of "B. coccifcr" from Guatemala, Honduras, and Nicaragua may have included specimens that we recognize as B. coccifcr, B. iharrai, and B. portcri. Owing to this confusion, the diagnosis of B. cycladcn presented in the original description has not been widely accepted (Porter, 1967; McDiarmid and Foster, 1981; Frost, 1985, 2003). We apply the taxon 6. c\/clndeii only to those populations on the Pacific slope of the Sierra Madre del Sur of Guerrero and Oaxaca, Mexico. It seems likely that the confusion associated with 6. cyclnden is the result of authors' having inadvertently compared a variety of different species in their efforts to distinguish B. coccifcr and B. ciicladcii. Mendelson (2001) provided additional comments on this problem. Bufo iharrai Stuart ' Figs. 3, 8, 9 Hufo iharrai Stuart: I S9. Holotypc: UMM7. 108000. Type kicality: .AsiTiMdero San Lorenzo |about 12 airline km sli^hth' East of North of LiLipa], Jalapa, GuatemaLi, 1725 ni. Bufo coccifcr — Cope, 1887 |in part]; Meyer and Wilson, 1971 |in part]; Campbell and Vannini, 1989 |in part]; Campbell, 1999 |in part]; McCranie and Wilson, 2002 [in part]. Hufo iharrai — Lyndn and Fugler, 1965 |in part]: Porter, 1966; I'rost, 19S5; Camplx'll, 1999; Mendelson, 2001; Frost, 200.1. Bufo luicrotif — Sdimidt and Stuart, 1941 |in part, tor reference to UMMZ840H3]. Diagnosis. — A large species of Bufo (males to 82.4 mm SVL; females to 94.4 mm SVL), having the following combination of characters: (1) tympanum e\'ident externally, about 45'7- diameter of orbit in males, about 40' r, in females; (2) canthal, supraorbital, supratympanic, postorbital, preorbital, pret\-mpanic. Fig. 8. Lateral aspects ot the heads ot adult male specimens of Bufo iharrai (upper: KU 58413) and B. pi^iiiuiis (lower: UMMZ WED 233723; holot\ pe). Scale bar = 1 cm. 14 Scientific Papers, Natural History Museum, The University of Kansas Fig. y. Dorsjl and vi-ntrdl dspects of representative adult specimens of Bufo ibiinni from Departamento Guatemala, Guatemala (male, left: UTA A-25825, SVL = 69.S mm; female, right: UTA A-47572, SVL = 93.3 mm) and B. pi^imnis from Michoaain, Mexico (male, left: UMM2 115353 [WED 10971], SVL = 43.5 mm; female, right: UMMZ 121578, SVL = 60.4 mm). parietal, and supralabial crests present; (3) cranial crests well developed, parietal crests low, thin, sharply angled medially; supratympanic crest large, bulbous; (4) tibia short, about 40% SVL; (5) feet short, abi>ut 40';, SVL; (6) middorsal tubercles sparse, roundecl, becoming spinose lateralh' in females, all dorsal tubercles in males rounded, usually indistinct or absent lateralh'; (7) wntral tubei-cles areolate, non-spinose in males and finelv spinose in females; (8) lateral descending row of enlarged tubercles indistinct or absent; (9) skin textinv sexualK- dimoiphic; (10) vocal slit unilateral in male; (11) ni. iiilcr!n/oulru> poorly differentiated from ;;;. iiilciiiuiiKlibitliiri^. but differentiated posteriorly, forming a large, unilobed vocal sac with variable amounts of pigmentation; (12) snout shape rounded in lateral \'ievv, weakK' pointed in dorsal view; (13) parotoid glands large, ovoitf, length about 2 times size of eyelid; (14) skin between cranial crests usually smooth, lacking tubercles; (15) \'entral coloration dull cream with some diffuse gra\' mottling in some inctividuals. Biifo ibarrai is the largest species in the B. coccifer Croup and differs from all species except B. portcri bv: ha\'ing i"elati\el\' massi\e cranial crests, with the supratympanic crest being distinctb' bulbous; and ha\ing sexually dimorphic skin texture with distincth' roimded lateral tubercles in males and spinose lateral tubercles in females. Bufo ihiiiiiii closely resembles B. porlcvi but differs b\: being larger (males to 82.4 mm SVL and females to <-)4.4 mm SVL \s. S^.M mm in males and 76.2 mm in females); and ha\ing the caudal muscuLiture ot the tadpole uniloini pale blown (cream and hea\il\' punctated with brown in B. portcri). Morphometric \ariation is summarized in Table 2. Distribution and ecology. — Bufo ihnrrni occurs at moderate ele\ations (1360-|9S() m) in the highlands of Toads of the Blfo coccii kk Compi.bx 15 Fig. 10. Map of Guiiteniiila and western Honduras with dot localities for Bufo ibnrnii and B. poilcn (see species account below) indicating specimens examined. central and southern duatemala (Mendelson, 2001), and new information presented here extends that range into contiguous regions of Honduras (Fig. 10). The Honduran localities range up to 2020m elevation, and all represent Premontane Moist Forest (e.g., McCranie and Wilson, 2001: pl.2E) or Lower Montane Moist Forest (e.g., McCranie and Wilson, 2001:pl.4E) and lie in the western ranges of the Southern Cordillera Region (sensu McCranie and Wilson, 2001), in the departments of Intibuca, Lempira, and Ocotepeque (Fig. 10). These discoveries refute Mendelson's (2001 ) premature speculation that B. ibnvrni ma\' be endemic to Guatemala and refute McCranie and Wilson's (2001) conclusion that B. ibanai does not occur in Honduras. Tadpoles. — The tadpole was described bv Mendelson (2001 ). Remarks. — Altliough Mendelson (2001) discussed diagnostic features that distinguish B. coccifcr and B. iharrai, he did not present multi\ariate analyses of morphometrv. In addition, Mendelson's (2001) research was based only on specimens from Guatemala. During the course of this stud\', Eric N. Smith kindly sent to us specimens of B. iharrai he collected in Ocotepeque, Honduras. These toads are consistent with the diagnosis presented b\ Mendelson (200 1 ) and have a similar (0.67' <'. di\-ergent) mitochondrial- DNA sequence to B. iharrai collected at the opposite end of the range of the species, in El Quiche, Guatemala. We also discovered additional specimens from western Honduras among museum collections (Appendix II) that match tiie diagnosis provided b\' Mendelson (2001 ); these specimens Bufo pisinuus new species Figs. 3, 8, 9 Bufo coccifcr Porter, 1%3 [in part]; Frost, 1985 |in part]; Villa et al., mSS |in part]: Flores-Villela, IW.I |in part]: Campbell, 1999 |in part]; I i-ost, 2003 |in part], Bufo ci/clihicn Lvnch and Smith, l"-l(i(i |in part, tor reference to specimens from Michoacan, Mexico]. Holotype.— UMMZ 233723 (WED 10973), an adult male from 6.2 mi [10 km] E Apatzingan, 1100 feet [335 m] elevation, obtained b\' W. E. Duellman and R. E. Etheridge on 2 August 1936. Paratypes. — All from Michoacan, Mexico: 6.2 mi [10 km] E Apatzingan, 1100 ft [333 m] (UMMZ 115353 [15 specimens]); 7 mi [11.2 km[ E Apatzingan, 1100 ft [335 m] (UMMZ 112794 [6 specimens), 115335); 1 mi [1.6 km[ W Apatzingan, 1100 ft [335 m] (UMMZ 115354); 3 mi [4.8 km] S Lombardia (UMMZ 121578). Diagnosis. — A small species of Bufo (males to 51 mm SVL; females to 62 mm SVL), having the following combination of characters: (1) tympanum evident , about 35^0 '}o diameter of orbit in males, about 40-50% in females; (2) canthal, supraorbital, supratympanic, postorbital, preorbital, pretympanic, supralabial crests present, parietal crest a thin ridge, or absent; (3) cranial crests weakly developed, thin, low; (4) tibia short, about 34';;, SVL; (5) feet short, about 36% SVL; (6) dorsal tubercles small to medium sized, low, rounded, relatively densely arranged middorsally, becoming smaller and spinose laterally; (7) ventral tubercles tiny, evenly granular, appearing smooth, especially in males, but tiny, spinose apices apparent under microscope; (8) lateral descending row of enlarged tubercles absent; (9) skin texture not sexually dimorphic; (10) vocal slit unilateral in male; (11) ;;;. iiitcrln/oideus poorly differentiated from ;;/. iutcniiaiidihiilaris, but differentiated posteriorly forming a large, unilobed vocal sac with heavy black pigmentation; (12) snout shape acutely rounded in lateral view, sharply pointed in dorsal view, snout shape in females similar but more genth' rounded in lateral view; (13) parotoid glands ovoid, about 1.0-1.5 times size of eyelid; (14) skin between cranial crests on top of head usually with many scattered, low, rounded tubercles; (15) ventral coloration usually 16 Scii-Miiic Paplrs, Nailkal History Museum, Thh Umvl;rsha oi Kansas immaculate whitish cream, some indixiduals with tinv, scattered black flecks. Biifo pnsiiiiitif can be distinguished from all other members of the B. coccifcr Group b\' its small si/e, and possession of: relatively weakly developed cranial crests, the parietal crest appearing only as a thin ridge among surrounding tubercles and frequentlv is absent; smaller, more densely arranged dorsal tubercles; and smaller, less spinose ventral tubercles. Bitfo pisinniis differs from B. coccifcr by having: a relatively shorter snout and an advertisement call with a higher pulse rate (120 pulses per sec vs. maximum of 115 pulses; Table 1, Fig. 2). Biifo pisi)uni^ differs from B. cycladcn by having: much smaller, and less spinose dorsolateral tubercles (large and conspicuously spinose in B. cycladcn); and smaller parotoid glands (about 1.5-2.0 times size of eyelid in B. cyclndcn). Biifo p'ifiiiniis differs from B. sigiiifcr by having: a whitish-cream venter, with or without scattered black flecks (venter with bold brown-black marbling in B. sigiiifcr); smaller parotoid glands (about twice size of eyelid in B. sigiiifer); and an advertisement call with a higher frequency and faster pulse rate (Table I, Fig. 2). Bufo pisiimiis differs from B. ibarrai by having: smaller, more rounded parotoid glands (glands in B. ibnrnii ovoid, much higher, and about twice size of the eyelid); and rounded mid-dorsal tubercles in both sexes (tubercles in B. ibarrai rounded in males, spinose in females). Bufo pisiunus differs from B. portcri by having: sharply spinose lateral tubercles in males (rounded in males of B. portcri); and a relatively thin supratympanic crest (large, bulbous in 6. portcri). Description of holotype. — Body robust; head wider than long, width 3M.1';, SVL, length 35.3";- SVL; snout sharply pointed in dorsal view, rounded in profile, rostral keel distinct; canthal, preorbital, supraorbital, pretympanic, supratvmpanic, and postorbital crests present, distinct; parietal crests reduced, barely distinct; skin on top of head co-ossified; nostril not protuberant, directed dorsallv; can thus rostralis forming distinct, raised, canthal crest; loival region cinna\'i-; lip distinct, rounded; suborbital crest present, distinct, extending from angle of the jaw anteriorlv to level of anterior margin ot orbit; notch at symphysis of upper jaw pivsenl, distinct; e\'e-nostril distance 5. 7'.r, diameterof orbit; t\-mpaniim distinct, nearh' round; tympanic annulus distinct onl\' along anterior and ventral margins, upper margin contacting supratympanic crest, posterit)r margin obsciu-ed by overlying llesh. Forelimb short, robust; hand broad, with short, slender fingers; relative length ot fingers: II • I < IV ■ III, webbing and lateral fringe on fingers absent; tips ot lingers not enlarged, smooth dorsallv, demarcated proximalK' b\- distinct dermal told; palmar tubercle elistincl, large, ovoid; pollical tubercle smaller than palmar tubercle. ovoid; subarticular tubercles distinct, elevated, triangular in profile, single except distal tubercle on Finger 111 bilid; supernumerary tubercles of unequal size, small, distinct, scattered evenly over palm and ventral surfaces of fingers; nuptial excrescences present as brown granular patches on medial surfaces of Fingers I and II. I iind limbs short, slender, tibia length 35.8',' SVL; foot length 38.0% SVL; tarsal fold absent; outer metatarsal tubercle minute, elevated, ovoid; inner metatarsal tubercle slightly larger than outer metatarsal tubercle, distinctly elevated, ovoid; toes long, slender, relative lengths of toes: I < 11 < V < III < IV; lateral fringe present on Toes II, 111, and \', absent on Toes I and IV; webbing thin, webbing formula 11 — 3II2— 3III2— 4IV4— 2V; tips of toes not enlarged, smooth dorsally, demarcated proximally by distinct dermal fold; subarticular tubercles distinct, elevated, triangular in profile, bifid; supernumerar\' tubercles unequal in size, distinct, distributed evenlv over \-entral surfaces of foot and toes. Skin on dorsum of bodv rugose with e\enl\' distributed, small, rounded tubercles of relatively equal size, becoming sharpiv pointed laterally; parotoid glands about same size as evelids, o\oid, oriented perpendicular to midline of bodv; lateral rov\' of enlarged tubercles barely evident; dorsal surface of head smooth with few, small, rounded tubercles scattered in interspaces between cranial crests; dorsal surfaces of limbs covered with small, weakly pointed tubercles; skin on throat and other ventral surfaces granular, covered with tinv flattened and weakly pointed tubercles. Choanae small, rounded, widely spaced; teeth and odontoids absent; tongue long, ovoid, about four times as long as wide, free posteriorly for about one-fourth its length; vocal slit unilateral, sinistral. Coloration of holotype. — In preser\ati\e (ethanol), dorsum ot bod\' and limbs mottled evenly with pale brown and dark brown markings; irregular cream blotches present posterior to each parotoid gland; top ot head uniform dark brown with distinct cream interorbital bar; distinct cream middorsal stripe extending from snout to posterior end o\ urostvie, irregularlv widened at se\'eral points along its length; lateral sin-faces pale brown with dark brown flecking. Ventral surfaces nearh' immaculate cream: dark \'ocal sac visible through gular skin. Measurements of the holotype (in mm). — SVL 45.0, HI. IS», IIW 17.(1, IL Ih.l. 14 17.1, orbit diameter 5.4, tympanum diameter 23, suprahmpanic crest length 2.7, parotoid gland length ^.4, pai'otoid gland width 4.8. Coloration in life. — Duellman (l'-)6l:2I) described coloration "...yellowish tan ground color with dark brow n spots; middorsal stripe is deep ncIIow or ci-eam color. 1 he \ enter is dust\- cream color, anil the iiis is pale gold." Toads of the Bcro cocciier Complex 17 Variation. — Morphometric Vciriation among specimens examined is summarized in Table 2. The parietal crest is poorly developed in most specimens, and essentially absent in a few individuals (e.g., UMMZ 121578). Some individuals (e.g., UMMZ 112794) have a dark-brown ground color on the dorsum, which effectively obscures the dark brown dorsal blotches; most specimens have a pale brown to grayish ground color on the dorsum, and the overlying brcnvn blotches are distinct. The brown blotches on the dorsum may be relativelv small, incorporating one to three dorsal ttibercles (e.g., UMMZ 115353 [WED 10972]), or the blotches may be larger, incorporating more than 12 dorsal tubercles (e.g., UMMZ 115353 [WED 10974]). The narrow middorsal cream stripe invariably is present in all individuals examined. Scattered black flecks on the ventral surface may be absent {c.;^., UMMZ 121578), restricted to the peripheral ventral surfaces (e.g., UMMZ 1 15353 [WED 10970]), or scattered relatively evenly across the venter (e.g., UMMZ 115354 [WED 10976]). Etymology. — The name pisiiiiiiis is the Latin word pi>iuiiii<, meaning small, in reference to the diminuti\'e size of this species. Distribution and ecology. — Bufo pisiiuiii^ is known only from the Tepalcatepec Valley, which represents the western extension of the Balsas Basin (Fig. 7). Duellman (Field Notes, 16 July 1960) described the area arciund Lombardia, Michoacan, as having grassy areas interrupted by areas of mescjuite with many reddish rocks and barren areas. Duellman (1961) suggested that this species is likely widespread in region, but noted that it does not occur along the coast of Michoacan. Duellman ( 1 961 ) reported breeding choruses in muddy ditches and flooded grassy fields after heavy rains in June and August. Tadpoles. — The tadpole of B. pifiiniiiis is unknown. Bufo porteri new species Figs. 3, n, 12 Bufocoi'iifcr — MevtTiind Wilson, l'-)7I |in p>irt|; iTost, 198S [in portl; McCranie dnd WiKon, 2002 |in part): Campbull, Iwm |in part]: Frcist, 20(1^ |in part|. Bufo ibarnii — l.vnch iind Fiigler, I'-Ui? |in part]. Holotype. — KU 97519, and adult male from 6 mi [9.6 km] NE Escuela Panamericana, Cerro Uvuca, 520(1 ft [1584 m], Francisco Morazan, Honduras, obtained by K. R. Porter on 30 June 1964. Paratypes. — All from Franciso Morazan, Hondinas: 6 mi ]9.h km] NE Escuela Panamericana, Cerro Uvuca, 5200 ft [1584 m] (KU 97520-26); 6 mi 19.6 km] NE Escuela Panamericana, Cerro Uvuca, 6000 ft 11828 m] (KU 97514); W slope Cerro Uyuca, 1650 m (KU 103221); Parque Nacional LaTigra aLxiveSan Juancito, 2100 m (KU 192294); Cerro La Tigra, 1840-1890 m (KU 194216-19, 209249). Fig. 11. Lateral aspects of the heads of adult male specimens of Bufo portcn (upper; KU 97520) and B. ngnifcr (lower: AMNH 69626). Scale bar = 1 cm. Diagnosis. — A small to medium-sized species of Blip (males to 59.9 mm SVL; females to 76.2 mm SVL), having the following combination of characters: (1) tympanum evident externallv, about 40',, diameter of orbit in both males and females; (2) canthal, supraorbital, suprat\'mpanic, postorbital, preorbital, pret\'mpanic, parietal, and supralabial crests present; (3) cranial crests well developed, parietal crests low, thin, supratympanic crest large, bulbous; (4) tibia short, about 37' ,'. SVL; 5) feet short, about 37"^. SVL; (6) middorsal tubercles sparsely arranged, rounded, becoming spinose laterallv in females, all dorsal tubercles in males rounded, usually bect)ming indistinct or absent laterallv; (7) ventral tubercles areolate, non-spinose in males, fineh' spinose in females; (8) lateral descending row of enlarged tubercles indistinct or absent; (9) skin textu re sexuall\'dimorphic;( 10 )\ocal slit unilateral in male; (11) iii. iiitciin/ouicu> poorly differentiated from III. iiitcniiiiiidibiilnrifi differentiated posteriorly, forming a Scientific Paphrs, Naturai. Hisiory Museum. The University of Kansas Fig. 12. norsjj iind ventral aspects ot representative adult specimens of Biifo portcrl from Francisco Morazan, Honduras (male, left: KL' ^7519, SVL = 51.6 mm; female, right: KU y7.S14, SVL = 60.7 mm) and B. signifcr Panama (adult male, left: TNHC 31341, SVl, = 49.7 mm; subadult female, right: KU 1 1 3359, SVI . = 54. 1 mm ). Itirgc, unilobed vocal sac hcavih' pii^LMiiented in black; (12) snout shape roiindccl in lateral \ic\v, wcakh' poifited in dorsal xiew; (13) pafotoid glatids moderate, rounci to ovoid, length about 1.00-1.23 titnes size of eyelid; (14)skifi between cranial crests usually smooth, lacking tubercles; (15) ventral coloration is dull cream with some diffuse gray mottling in some individuals. Biifo poiicii ma\' be distingiushed trotn all other tnember of file B. coaifcr Group, except B. ihninu, the sexual dimorphism in the texture of the dorsal skin and the relatively robust cratiial crests, especialh' the supratympanic crest. Biifo poiicri closely resetnbles />'. ibanni but differs by being smaller (tnales to 54.9 mm SVL and females to 76.2 mm SVL \'s. males to >S2.4 mm SVL and females to 94.4 mtii SVL); adult male B. ]wrtcn as small as 43.9 mm SVL have been observed (e.g., LSUMZ 46431 ). The caudal musculature of the tadpole of B. poiicri is creafTi w ith hea\ \ brown punctations (See description b\' McCranie and Wilson, 2002:173; caudal musculature if! B. ibarrai is unifortii pale browti.) Description o( holotype. — Bod\ robust; head \\ ider than long, width 40.0% SVL, length 36.4% SVL; snout sharply pointed in dorsal view, pointed in profile, rostral keel distinct; canthal, preorbital, supraorbital, pretytnpanic, supratympatiic, and postorbital crests present, distinct; supraorbital atid suprat\'mpanic crests distintly thickened; parietal crests present, not reduced; skin on top of head co-ossified; nostril protuberant, directed dorsally; canthus rostralis forming distinct, raised, canthal crest; loreal region concaM'; lip distinct, rounded; suborbital crest present, distinct, extending from angle of the jaw anteriorly to le\'el of afiterior margin of orbit; notch at s\'mphvsis of upper Toads of ihh Buio coven ik Complex 19 jaw present, distinct; e\'e-n(istril distance 37.9",- diameter of orbit; tympanum distinct, nearly round; tympanic annulus distinct only along anterior, xentral, and posteroventral margins; upper margin of tympanic annulus contacting and obscured by supratvmpanic crest. Forelimb short, robust; hand broad with short, slender fingers; relative length of fingers: II < I < IV < III, webbing and lateral fringe on fingers absent; tips of fingers not enlarged, smooth dorsallv, demarcated proximally by distinct dermal fold; palmar tubercle distinct, large, rounded; pollical tubercle smaller than palmar tubercle, rounded; subarticular tubercles distinct, ele\ated, triangular in profile, all single except distal tubercle on Finger 111 bifid; supernumerary tubercles of unequal size, small, distinct, scattered evenly over palm and ventral surfaces of fingers; nuptial excrescences present as brown granular patches on medial surfaces of Fingers l-IIl. Hind limbs relatively long, slender, tibia length 41.7",. SVL; foot length 40.5",'. SVL; tarsal fold absent; outer metatarsal tubercle very small, rouncied, indistinct; inner metatarsal tubercle much larger than outer metatarsal tubercle, distinctly elevated, ovoid; toes long, slender, relative lengths of toes: I < II < V < III < IV; lateral fringe barely evident on Toes III and IV, absent on other toes; webbing thin, webbing formula II — 3II2 — 3III2 — 4IV4 — 2V;'tips of toes not'enlarged, smooth dorsally, demarcated proximally bv distinct dermal fold; subarticular tubercles distinct, elevated, triangular in profile, bifid on Toes III and IV; supernumerary tubercles unec^ual in size, distinct, distributed evenly over ventral surfaces of foot and toes. Skin on dorsum of body relatively smooth with scattered, small, rounded tubercles of unequal size, many bearing tiny, indistinct single keratinized apices; tubercles on lateral surfaces indistinct, roimded or ovoid, ncit pciinted; parotoid glancis about same size as eyelids, ovoid, oriented slightly divergent tci midline of body; distinct lateral row of enlarged tubercles absent; dorsal surface of head smooth \yitlT few, small, rounded tubercles scattered in interspaces between cranial crests; dorsal surfaces of limbs covered with small weakly pointed tubercles; skin on throat and other ventral surfaces smooth!)' granular. Choanae moderate in size, rounded, widely spaced; teeth and odontoids absent; tongue long, ovoid, about four times as long as wide, free posteriorly for about one-foiuih its length; vocal slit unilateral, dextral. Coloration of holotype. — In preservative (ethanol), bod\' pale brown with indistinct, medium-brown marbled markings diffused across middorsal and lateral areas; distinct, thin cream middorsal stripe present; two oblong dull cream patches on dorsolateral areas; distinct dark brown bar extending across area between parietal crests and another covering area between canthal crests; all limbs with indistinct medium brown crossbars; venter immaculate dull cream; dark vocal sac clearly \'isible through gular skin. Measurements of the holotype (in mm). — SVL 51.3, HL 18.7, HW 20.5, TL 21.4, FL 20.8, orbit diameter 7.6, tympanum diameter 3.0, supratympanic crest length 3.1, parotoid gland length 6.9, parotoid gland width 4.8. Coloration in life. — McCranie and Wilson (2002:pl '■'D) provided a color photograph of this speceis; see also Figure 3. Variation. — Morphometric variation among specimens examined is summarized in Table 2. The parietal crests in females generally are well developed and distinct; however, the condition of the crest varies among males and may be relatively robust (e.g., KU 209249) or reduced to an indistinct, thin sliver of raised bone (e.g., KU 97523). The brown dorsal blotches may be very dark brown, and therefore quite distinct (e.g., LSUMZ 46398), or they may be only slightly darker than the brown ground cc^lor of the dorsum (e.g., LSUMZ 46445). The ventral coloration is either uniform dull cream (e.g., KU 103221) or bearing diffuse, grayish-brown marbling that ranges from moderate (e.g., KU 97522) to extensive (e.g., KU 194216). The middorsal cream stripe invariably is present on all individuals examined; this stripe is quite indistinct in a small nimiber of individuals (e.g., KU 97520) and may appear irregular (i.e., not ft)rming a straight line; e.g., LSUMZ 45441). Etymology. — The specific epithet is a patronym that honors Kenneth R. Porter and his series of papers (e.g.. Porter, 1963, 1964, 1965)onthesystematicsof Mesoamerican Biifo, and also recognizes his numerous field efforts that resulted in many of specimens referred to herein. Distribution and ecology. — Bitfo povtcvi is known from the Hondiu-an departments of Comavagua, Francisco Morazan, and La Paz (Fig. 10). The known localities for this species generally represent Lower Montane Moist Forest habitats (e.g., McCranie and Wilson, 2001 :pl. 4C) in the Montanas de Comayagua region. These localities differ markedly from the lower elevation, Dr\' and Arid Forest habitats (e.g., McCranie and Wilson, 2001 :pl.lC) occupied b\' B. coccifer. Tadpoles. — Tadpoles leferrable to B. yoiicri (based on geography) were described bv McCranie and Wilson (2002:173). The tadpole of this species resembles that of B. coccifer (McDiarmid and Foster, 1981; Sa\age, 2002), from which it is distinguished by having submarginal papillae on the oral disc (absent in 6. coccifer) and cream-colored caudal musculature with brown punctations (boldly marked with brown saddles in B. coccifer). Remarks. — We have allocated many Honduran specimens that previously were referred to B. coccifer 20 Scientific Paphrs, NATiiRAi. Hisiok^ Mu.shum,Thh UNivi-.Rsrn oi Kansas (e.g., McCranie and Wilson, 2001) to the new taxon B. porteri, or to B. ibarrai (discussed above, and Appendix II). The difficulty of identifying specimens from this country is exacerbated bv the resemblance of B. porteri and B. ibarrai. The wide range of variation among specimens of "B. coLxifcr" from Honduras described by McCranie and Wilson (2001 ) seems to be attributable to the fact that three relative!)' similar species occur in close proximit)' in that country. At a general le\el, this species appears to be parapatric with respect to the distribution of Bnfo coccifcr. Thus, the distribution and habitat associations of this species, with respect to those of B. coccifer, resemble the relationship between B. coccifcr and B. ibarrai in Guatemala as described by Mendelson (2001 ). Inasmuch as species of Bnfo frec]uentl\- are interfertile (Blair, 1963; Masta et al., 2002), it is possible that B. porteri may hybridize with B. coccifer if the species co-occur on the lower slopes of the Pacific Versant of Honduras. Similarly, hvbrids between B. porteri and 6. ibarrai eventually may be found. Bnfo sigtiifer new species Figs. 3, 11, 12 Bufo coccifer Dunn, 1933; Zweifel, 1965; Frost, 1985 [in part, tor reference to specimens from Panama]; Villa et al., 1988 |in part, for reference to records from Panama]; Campbell, 1999 |in part]. Holotype.— AMNH 69626, an adult male from 7 mi N [11.2 km] and 2 mi [3.2 km] W of David, Chiriqui', Panama, obtained bv R. G. Zweifel on 25 June 1962. Paratypes. — All from Panama. Chiriquf: 7 mi [ 1 1 .2 km[ E ConcepcicSn (AMNH 69627); 2.5 mi [4.0 km| NE David (TNHC 3134-43); 23 km NNE San Felix, 900 m (USNM 297511-21). Code: El Valle de Anton (AMNH 59634, 59637); 16 km S, 9 km W Penonome, 30 m (KU 11 5359-6 1 ); 3.2 km W Agua Dulce, 15 m (KU 115362). Herrera: Jacinto, 2250 ft [686 m] (ANSP 22341-44); 3 mi [4.8 km] SW Pan American Hwy, on road past Potuga (UMMZ 167373). Veraguas: 14 km NE Sona, 75 m (KU 95432). Diagnosis. — A medium-sized species of Bufo (males to 64 mm SVL; females to 77 mm SVL), ha\ ing the following combination of characters: (1) tympanum evident externally, about 40-45'('. diameter of orbit in males, about 40-50% in females; (2) canthal, supraorbital, supratympanic, postorbital, preorbital, pret\nipanic, parietal and supralabial crests present; (3) most cranial crests distinct and thick, except parietal crests low, thin, sometimes intermittent; (4) tibia short, about 35'f' SVL; (5) feet short, about 36' V, SVF; (6) dorsal tubercles medium- sized, prominent, roimded, rel,ili\el\' denseh' arranged middorsally, becoming smaller, more concentrated, antl spinose laterally; (7) ventral tubercles granular, willi small, distinct spinose apices; (8) lateral desci'nding row of enlargeti tubercles absent; (9) skin texture not sexualh' dimorphic; (10) vocal slit unilateral in male; (11) iii. iiiterh\/oiiieiis poorly differentiated from iii. iiiteriiniiulibulari^, but differentiated posteriori)' forming a large, unilobed vocal sac with heavy black pigmentation; (12) snout shape rounded in lateral view, pointecf in dorsal \'iew; (13) parotoid glands round to subovoid, about twice size of e\'elid; (14) skin between cranial crests on top of head usually with few, scattered, low, rouncfed tubercles; (15) ventral coloration cream with distinct, bold, brown-black marbled pattern, becoming indistinct o\er pehic patch. Bnfo ^i^'^iiifer may be distinguished from all other members of the Bufo coccifer Group bv ha\'ing a cream venter o\'erlain with a distinct, marbled pattern of brown- black markings. All t)ther species in the group have immaculate, ornearh' immaculate, cream ventral surfaces with the exception that some indi\iduals of B. cucladeu may ha\'e some dark brown mottling, and some indix'iduals of B. pisinniis mav ha\'e some tiny, black flecks. Bufo si^i^uifer differs from B. coccifer by having: relatixel)' thinner and lower parietal crests (typically higher and thicker in B. coccifer); relatively smaller tympana; and an advertisement call with a lower frequencv and pulse rate (Table 1, Fig. 2). Bufo :^igiiifer is superficially quite similar to B. ci/cladeu but differs by: being larger (males of B. cycladen to 54 mm SVL, females to 62 mm); having a distinctly marbleci \'entral pattern (variably present, but always less developed in B. cycladen); and having generallv smaller, less spinose tubercles overall (but this latter character is someu'hat \'ariable and subjective). Bnfo signifer differs from B. pisinnub bybeing larger (males of B. pi^inuns to 51 mm SVL, females to 62 mm) and by having: larger middorsal tubercles (small in B. pisinnns); larger parotoid glands (glands about 1.0-1.5 times size of eyelid in B. pisiniius); spinose tubercles on the venter (granular, non-spinose in B. pisi)i}uis); an advertisement call with a lower frequency and pulse rate (Tablel, Fig. 2); and better-developed cranial crests, especially with respect to the parietal crest (crests weaklv developed, and parietal crest very reduced or absent in B. pissiiin^). Bnfo signifer differs from B. ibarrai by being smaller (males of B. ibarrai to 82 mm SVL, females to '^'4 mm) and b\' having: smaller, more rounded parotoid glands (glands in B. ibarrai ovoid, about twice the size of the e\'elid); and rounded mid-dorsal tubercles in both sexes (tubercles in B. ibarrai roimded in males, spinose in females). Bnfo fiignifer differs trom B. porteri by having: sharplv spinose lateral tubercles in males (roimded in males of B. porteri); and a relatively thin supratympanic crest (large, bulbous in B. porteri). Description of holotype. — Body robust; head wider than long, width 39.9', SVL, length 34.7';;, SVL; snout sliarph' pointed in dorsal \'iew, rounded in profile, rostral keel distinct; canthal, pivorbital, supraorbital, pretympanic, siiprat\mpanic, and postorbital crests present, distinct; Toads of the Bufo coccih.r Complex 21 parietal crests present, relati\el\' indistinct; skin on top of head coossified; nostril protuberant, directed dorsally; canthus rostralis forming distinct, raised, canthal crest; loreal region concave; lip distinct, rounded; suborbital crest present, distinct, extending from angle of the jaw anteriorly to level midway between orbit and nostril; notch at symphysis of upper jaw present, shallow, indistinct; eye-nostril ciistance 47.6' r. diameter of orbit; t\mpanum distinct, nearlv round; t\mpanic annulus distinct only along anteroventral margin, upper margin contacting supratympanic crest, posterior margin obscureci by overlying flesh. Forelimb short, robust; hand broad, with short, slender fingers; relati\'e length of fingers: II < I < IV < III, webbing and lateral fringe on fingers absent; tips oi fingers not enlarged, smooth dorsallv, demarcated proximalh' by distinct dermal fold; palmar tubercle distinct, large, round; pollical tubercle smaller than palmar tubercle, ovoid; subarticular tubercles distinct, elevated, triangular in profile, single except ciistal tubercle on Finger III bifid; supernumerary tubercles of unequal size, large, distinct, scattered evenly over palm and \entral surfaces of fingers; nuptial excrescences present as bro\N'n granular areas on entire dcirsal surfaces of Fingers I and II, including lateral surfaces of tips of fingers, lateral surfaces of distal phalange of Finger III, and on medial surface of pollical tubercle. Hind limbs short, slender, tibia length 34.5' «'. SVL; foot length 35.1"o SVL; tarsal fold absent; outer metatarsal tubercle small, elevated, ovoid; inner metatarsal tubercle slightly larger than outer metatarsal tubercle, distincth elevated, ovoid; toes long, slender, relative lengths of toes: I < II < V < III < IV; lateral fringe on toes barely evident on Toes II and III, absent on other toes; webbing thin, webbing formula II — 27,111'/, — 3III2 — 4IV4 — 2V; tips of toes not enlarged, smooth dorsally, demarcated proximally by distinct dermal fold; subarticular tubercles distinct, elevated, triangular in profile, single except two most distaltubercles on Finger IV and most distal tubercle on Finger V bifid; supernumerary tubercles unequal in size, distinct, distributed evenly over ventral surfaces of foot and toes. Skin on dorsum of body rugose with scattered, rounded tubercles of unequal size, becoming sharply pointed laterally; parotoid glands about same size as eyelids, slightly ovoid, oriented parallel to midline of body; lateral row of enlarged tubercles barely evident; dorsal surface of head smooth with few, small, rounded tubercles scattereti in interspaces between cranial crests; dorsal surfaces of limbs covered with small to large, mostly pointed tubercles; all ventral surfaces rough, co\'ered with small, conical tubercles. Choanae small, ovoici, widely spaced; teeth and odontoids absent, but ventral surface of neopalatine appears serrate; tongue long, ovoid, about four times as long as wide, free posteriorly for about one-fourth its length; \'ocal slit unilateral, dextral. Coloration of holotype. — In preservative (ethanol), dorsal areas of body and limbs mottled evenly with pale brown and dark brown markings, with a bilateral series of small, oblong cream markings dorsolaterally; top of head uniform dark brown with distinct cream interorbital bar; distinct cream middorsal stripe extending from snout to posterior end of urostyle; lateral surfaces dull brown with some dull cream supralabial spots. Ventral surfaces dull cream with indistinct pale gray markings, forming loosely reticulate pattern; dark vocal sac barely visible through gular skin. Measurements of the holotype (in mm). — SVL 57.1, HL 20.5, HVV 23.0, TL l'-).9, FL 20.0, orbit diameter 8.2, tympanum diameter 4.4, supratympanic crest length 3.4, parotoid gland length 6.8, parotoid gland width 5.9. Coloration in life. — Original field notes by R. G. Zweifel { 1 2 June 1 962) describe AMNH 69625 as " . . .a rather black and white looking toad, the whiteness coming from 80° 10° B. agnifer < 300 m 300-900 m 900-2100 m I ""H > 2100 m 0 100 Kilometers 80= Fig. 13. Map of western Panama with dot localities for Biifo >iguiffr indicating specimens examined, which represent all known localities. 97 SciHNTiHC Paphrs. Natural History Museum, The University of Kansas the lateral stripe which begins at tlie parotid and runs back to the groin and by the central vertebral line. The blackness is supplied by the dorsal blotches which are almost jet- black. The ground color between the blotches is dark gray. The side below the lateral line is a mixtiuv ot black and gray. The eyelids are a stripe with grayish white and black. The interorbital area is black with a grayish white band. The \'entral surfaces are mottleci gray and white; there is a dark gray spot in the middle posterior gular region. The hind legs are barred very with dark gray and grayish white as are the limbs right out to the tips of the toes." Variation. — Morphometric variation among specimens examined is summarized in Table 2. The parietal crests in males may be relatively distinct and well formed (e.g., TNHC 31343, or they may be essentially absent (e.g., TNHC 31342); this crest is always relatively distinct in females. The dorsal pattern usually consists of a contrasting array of dark brown or black blotches over a pale brown ground color, but some specimens (e.g., TNHC 46261 ) have a nearly uniform, dull brown dorsimi. Nearly all specimens of this species bear a distinctive, highly contrasting marbled pattern on the venter; this pattern is present, but less distinct in a few specimens (e.g., TNHC 46261, USNM 297515, AMNH 69626). The middorsal cream stripe is present in all individuals, but may be incomplete (e.g., AMNH 69627). Etymology. — The name sij^iiifcr Latin, meaning bearing marks, refers to the ventral coloration of this species. Reproductive biology. — The tadpole of B. signifcr is unknown. Zweifel (personal communication) found calling males in a "weedy, shallow, muddy roadside pool" cin 25 June 1962; also found in this pool were ElnchiitocU'is sp. and Lcptodncti/liib in^ulnniiii. Distribution and ecology. — This species is known from each of the disjunct areas of Tropical Dry Forest (Campbell and Lamar, 2004:47, color map 6) along the Pacific Coast of Panama (Fig. 13). These regions are located in the vicinity of city of David, and the Province of Veraguas eastward toward the Canal Zone. MORPHOMETRIC ANALYSES Overall morphometric variation fc)r all species is summarized in Table 2. Two Principal Components Analyses were conducted: one with log-transformed data, and one using residuals of each variable regressed on SVL. These analyses produced similar results (not presented here) wherein the first PC had the greatest eigenvalue (and accounted for the majority of the variation explained ) and represented an overall size axis. Despite reasonably high loadings for variables such as TYMP and PARW on the second PC, plots individual scores for each specimen did not clearly distinguish among the species. As could be expected, Biifo coccifer showed the most overall variation, while the smallest species (6. pisiiniiifi) was somewhat distinct from the largest species (B. ihnrrai) along the size axis (PCI). Considered together, the results of these analyses indicate that, with the exception of overall size, there has been relatively little morphometric differentiation among the species in this group. Variable selection procedures (forward, backward, and stepwise) for LDA suggested retention of all nine variables. i.e., residuals of eight variables regressed on SVL and log-transformed SVL. The cross-validated classification- matrix from the LDA is shown in Table 3. The majority of individuals of all species were correctly classified. Individuals of Biifo sigiiifer and B. porteri were misclassified inconsistenth' among all other species. Biifo pisimiiis and B. cycladcn had the greatest percentage (86%, 80'v,) of correctly classified individuals, respectively. The small size of 6. pisiiuuifi likely is responsible for this high degree of correct classification, but we note the 11 individuals of the quite variable species B. coccifer were misclassified as B. pifiinius; this result suggests that small B. coccifer are morphometrically similar tti B. pisiiniiis. Similarly, the relatively large size of B. ibarrai likely is responsible for the relatively high percentage (79' ,',) of correctly classified individuals of that species (and the very few individuals of other species that were identified as B. ibarrai). In general, these results are consistent with the results of the PCA. Despite its wide range of overall variation (Table 2), 77% of indi\iduals of B. coccifer were classified correctly. Table 3. Cross-validated classification-matrix and overall correct classification percentages (rounded to nearest integer) from Linear Di.scriminant Analysis based on residuals of eight morphometric variables regressed on SVL and log transformed SVL from all species in the Bufo coccifer Group; analysis includes only adult males. Values in boldface indicate number of individuals correctly classified. Per cent correct N B. coccifer B. cycladcii B. ibarrai B. pi: isiiuif B. sif;nifcr B. porteri cla ssification B. coccifer 106 82 4 2 11 4 3 77 B. cyctaden 30 1 24 0 2 3 0 80 B. ibarrai 79 0 2 62 0 2 3 79 B. pi'^aiiiu^ 21 1 1 0 18 1 0 86 B. aignifcr 21 0 2 1 1 16 1 76 B. porleri 43 3 1 1 5 1 34 79 Toads of the Bufo coccifer Complex 23 Table 4. Voucher numbers and locality int'omiation for specimens o( Bii/o used in the molecular analyses. GenBank accession numbers are given for I6S and cyt-b. respectively. Taxon Voucher No Local itv GenBank No. B. ihcinai I B. iharrai 2 B. coccifer HS B. coccifer Nl B. coccifer HO B. coccifer CR B. cycliulen B. conifenis B. valHceps B. mcirimis UT.^-ENS 10270 UTA-.IAC 19612 KU 290030 SDSNH-ARH0I3 USNM 547980 TC'WC 8399S UTA-JRM 4607 MVZ 203775 MZFC-JRM 3868 UTA A-50864 Honduras: Ocotepeque (iuatemala: Quiche HI Salvador: Morazan Nicaragua: Ometepe Is. Honduras: Gracias A Dios Costa Rica: San Jose Mexico: Guerrero Costa Rica: Cartago Mexico; Veracruz Guatemala: Zacapa .A'i'927854,.'\Y927S6l .'\Y927S55, AY927862 .•\Y927S56, AY927S63 .•\Y927857. A>'927864 .•\'i'92930I.AY929303 .^929302. AY929304 AY927858. AY927865 U52800. AY008255 AYO0821I.AYOO82I2 AY927860. AY927867 MOLECULAR ANALYSES Sequences of 547 and 410 base pairs were obtained for 16S and cyt-b mtDNA genes, respectively; specimen information and GenBank Accession numbers are listed in Table 4. These sections correspond to the 4004—1556 and 16422-16818 positions for 16S and cyt b, respectively, on the mtDNA genome of Xeuopus [Pipidae] (Roe et al., 1985). Of these 956 base pairs, 765 were constant, 191 were variable, and 85 were considered parsimony-informative characters. The partition-homogeneity test results indicated that trees from the separate genes were not significantly different from one another (P = 1.0). Parsimony analysis of the 16S region produced four trees. Parsimony analysis of the cyt-b region produced one tree that was identical the combined analysis shown in Figure 14. The differences among the 16S trees were the monophyly/paraphyly of Bufo iharrai, and the relationships among B. coccifer (sensu stricto) samples. Parsimony analysis of all nucleotide data combined evaluated a total of 2,027,025 trees, with the best tree score of 278 steps (Fig. 1 5) and the worst tree score of 373 steps. The frequency distribution of trees scores had a mean of 351.60 steps (sd = 14.8; g, = 1.20; g, = 0.94). The hLRTs selected the TrN + I + G model (Tamura and Nei, 1993) as the most significant (p < 0.01; -InL = - 2612.9934) with base frequencies of A = 0.3071, C = 0.2448, G = 0.1620, T = 0.2860. The substitution rate matrix was: A-G = 6.4270, C-T = 12.1788, all others were equal to 1; the proportion 6. valliceps B. marinus B. coniferui B. cycladen B. ibarrai Guatemala 5. ibarrai Honduras B. coccifer Nicaragua 6. coccifer Honduras S. coccifer El Salvador B. coccifer Costa Rica Fig. 14. Phylogenetic relationships among samples used in molecular analyses; see Table 4 for specimen information. Topology is shown from the maximum-parsimony exhaustive search. Bootstrap values greater than 50' ». (MP /ML) are shown for each analysis above branches and decav indices are shown below. 98/99 100/82 91 89 10 11 2 97/74 2 61 7 1 51 1 Table 5. Pair-vvisc sequence divergence of 1 6S and cyt b combmed for specimens ofBii/o used in the molecular analyses. Absolule distances are shown above the diagonal, and Tamura-Nei corrected distances are below the diagonal. Taxon 1 "> 3 4 5 6 7 8 9 10 1 . B- il''ork. McDiarnud, R. VV., and M. S. Foster 198 1 . Breeding habits of the load Bufo cihi ifcr in Costa Rica, with a description of the tadpole. Southwestern Naturalist 26:.35.3-363. Toads oi nil; Bi i o coccii i.k C()mi'i.i;x MendeKon, J. R., HI. 1997. A new species of Bufo ( Anura: Butonidac) from the Pacific Highlands of Guatemala and southern Mexico, with comments on the status of Bufo vnUkiyi iiuuwcristntii>. Hcrpetologico 53:14-30. Mendelson, J. R., HI. 2(101. A re\'ievv of the Guatemalan toad Bufo ihnrrai (Anura: Bufonidae), with distributional and taxonomic comments of Bufo vnlhccps and Bufo coccifcr. Pp. 10-iy iii J. D. Johnson and R. G. Webb, and O. Flores-Villela (eds.), Mcfoaiitericnu Hcrpctology: Si/slciimtics, Niiluml Hilton/, nud Coii>cn'nliou. Tiie L'ni\'ersit\' of Texas at El Paso, El Paso, Texas. Mertens, R. 1932. Die .Amphibien und Reptilien von El Salvador, auf Grund der Reisen von R. Mertens und A. Zilch. Abhandlen Senckenbergischen Naturforschenden Gesellschatt 487:1-83. Meyer, J. R., and L. D. Wilson. 1971. A distributional checklist of the amphibians of Honduras. Contributions Science Natinal Histor\- Museum Los Angeles County 218:1-47. Myers, C. W., and W. E. Duellman. 1982. A new species of Hi/ln from Cerro Colorado, and other tree frog records and geographical notes from western Panama. American Museum Noxitates 2752:1-23. Mulcahy, D. G., and Mendelson, J. R., 111. 2000. Phylogeography and speciation of the morphologicallv variable, widespread species Bufo vnlliccpf. based on molecular evidence from nitD\A. VIolecular Phylogenefics and Evolution 17:173-189. Porter, K. R. 1962. Evolutionarv Relationships of the Bufo ;v7//;a7's Group in Mexico. Ph. D. Dissertation, University of Texas, Austin, Texas. Porter, K. R. 1963. Distribution and taxonomic status of seven species of mexican Bufo. Herpetologica 19:229-247. Porter, K. R. 1964. Morphological and mating call comparisons in the Buto valUccps complex. The American Midland Naturalist 71:232-245. Porter, K. R. 1965. Interspecific variation in mating call of Bufo coccifcr Cope. The American Midland Naturalist 74:350-356, Porter, K. R. 1966. Mating calls of six Mexican and Central American toads {genus Bufo). Herpetologica 22:60-67. Porter, K. R. 1967. Bufo cyclndcn (Bufonidae): a case of uoiiicii dubunn. Southwestern Naturalist 12:200-201. Porter, K. R., and W. F. Porter. 1967. Venom comparisons and relationships of twenty species of New World toads (genus Bufo). Copeia 1967:298-307. Posada, D., and K. A. Crandall. 1998. Modeltest: testing the model of DNA substitution. Bioinforniatics 14: 817-818. Rand, A. S. 1957. Notes on ampliibians and reptiles from El Salvador. Fieldiana Zoology 34:505-534. Roe, B. A., D. P Ma, R.' K. Wilson, and J. F. H. Wong. 1985. The complete sequence of the Xcnopus hicoh mitochondrial genome. Journal Biological Chemistry 260:9759-9774. Rzedowski, J. 1994. Vegetaciiin de Mexico. Limusa Noriega Editores, Mexico City, Mexico. SAS software. Version 8 of the SAS S\stem for Windows. 1999-2000. SAS Institute Inc. SAS and all other SAS Institute Inc. product or service names are registered trademarks or trademarks of S.AS Institute Inc., Cary, North Carolina. Savage, J. M. 1974. Type localities for species of amphibians and reptiles described from Costa Rica. Revista de Biologi'a Tropical 22:71-122. Savage, J. M. 2002. Ihe Amphibians and Reptdes of Costa Rica: A Herpetofauna between Two Continents, between Two Seas. University of Chicago Press, Chicago, Illinois. Saxage, J. M., and W. R. Heyer. 1967. Variation and distribution in the frog genus PhyUoincduf:! in Costa Rica, Central America. Beitrage Zur Neotropischen Fauna 5:111-131. Savage, J. M. and W. R. Heyer 1997. Digital webbing formulae for anurans: a refinement. Herpetologica! Review 28:131. Savage, J. M., and J. D. Villa. 1986. Introduction to the herpetofauna of Costa Rica. Contributions to Herpetology 5, Society for the Study of Amphibians and Reptiles, Oxford, Ohio. Schmidt, K. I'., and L. C. Stuart. 1941. The herpetological fauna of the Salama Basin, Baja Verapaz, Guatemala. Zoological Series of the Field Museum of Natural History 24:233-247. Smith, H. M., and E. H. Taylor. 1948. An annotated checklist and key to the amphibia of Mexico. Bulletin United States National Museum 194:1-118. Soren.son. M.D. 1996. TrccRol. University of Michigan. .Ann Arbor. Stuart, L. C. 1954a. A description of a subhumid corridor across northern Central America, with comments on its herpetofaunal indicators. Contributions Laboratory Vertebrate Zoology, University of Michigan 63:1-26. Stuart, L. C. 1954b. Descriptions of some new amphibians and reptiles from Guatemala. Proceedings of the Biological Societ\- Washington 67:159-178. Stuart, L. C. 1963. A checklist of the herpetofauna of Guatemala. Miscellaneous Publications, Museum of Zoology, Unix'ersitv of Michigan 122:1-150. Swofford, D. L. 2002. PAUP". Phvlogenetic .AnaKsis Using Parsimonx' (*and other methods). Version 4.0b 10. Sinauer .Associates, Sunderland, Massachusetts. Tamura K., and M. Nei. 1993. Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA and chimpanzees. Molecular Biology and Evolution 10:512-526. Taxlor, E. H. 1952. A reviexv of the frogs and toads of Costa Rica. Universits- of Kansas Science Bulletin XX,\V:577-942. Tyler, M. J., T. Burton, and A. M. Bauer. 2001. Parotid or parotoid: on the nomenclature of an amphibian skin gland. Herpetological Review 32:79-81. Villa, J. 1972. Anfibios de Nicaragua. Institito Geografica Nacional and Banco Central de Nicaragua. Villa, J. 1983. Nicaraguan fishes, amphibians, and reptiles: a checklist and bibliographx'. Escuela de Ecologia y Reairsos Naturales, Unix'ersidad Centroamericana, Managua, Nicaragua. Villa, ].. L. D. Wilson, and J. D. Johnson. 1988. Middle American Herpetology: A Bibliographic Checklist. University of Missouri Press, Columbia, Missouri. Werner, F. 1896. Beitrage zur kenntniss der reptilien und batrachier von Centralamenka und Chile. Verhandl. k.k. Zool.-Bot. Gesell. Wien 46:1-24. VVilex', E. O. 1978. The exolutionarv species concept reconsidered. Sx'stematic Zoology 27:17-26. Zxveifel, R. G. 1965. Distribution and mating calls of the Panamanian toads, Bufo coccifcr and B. ■^'rudii/iisiis. Copeia 1965:108-110. 26 SciENTiiic Papkrs. Natlral Histor'> ML'siaM.Tiiii Universh^ oi Kansas APPENDIX Specimens examined Note that data presented here has been taken verbatim fnmi museum catalogs and tags. We have not attempted to correct various spellings, nor to alter original estimates of distance and elevation. We do not consider it reasonable to re-estimate primarv locality data. Biifo coccifer: Costa Rica: Ai .muei a: Rio CIrande (LACM 594.'$2). PuNTARtNAS: 1.5 km W Barranca, 20 m (KU fi.S.'iSl); 4 km WNW Esparta, 45 m (KU 65397-90, 65393, 65397-98, 65400, 6.5404-05, 65414). San Jost: Guadalupe, 1190 m (KU 65379); Esca/ii, 1100 m (KU 65380). Guatemala: Cimqiumii a: above headijuarters of Finca San Jose, ca. 6.0 km SE Concepcion l.as Minas (UTA A-38119); Finca San Jose, ca. 6.0 km SE Concepcion Las Minas on rd to Las Presas (UTA A-381 23-26); Concepcion Las Minas, Valle Arriba (UTA A-47566); 2.6 mi N intersection of CA-12 and CA-10 (UTA A-38120-21); Jalaca: Volcan Jumay (UTA A-47591). RtTALHULEU: 3.2 km N Champerico (UTA A-25814-'23); 3.7 km N Champerico (UTA A-29021-25). Santa Rosa: El Oratorio, 3.0 km E CA-8 (UTA A-38143). Hi Salvador: Chalatengo: 16.5 km WNW Chalatengo (KU 184659-62); 10 km NE La Palma, Canton Las Pilas (KU 184663-69), Cuscatlan: Tcnancingo, Rio Quezalapa (KU 184572 i73); 7 km W Cojutepeque, 2900 ft (KU 97495); 8 km W Cojutepeque, 2850 ft (KU 97499-504); Cojutepeque, 3220 ft (KU 97505-09); 0.5 km SE Cojutepeque, 2520 ft (KU 97510-11); 2 km E Cojutepeque, 3250 ft (KU 97496). l.A Libtrtad: 10 mi NW Santa Tecla (KU 41411-29); 5 mi W Colon (KU 97473-86); 5.4 mi W La Libertad, 680 ft (KU 97487-92); San Bartolo, 11 km E San Salvador, 2595 ft (KU 97493). La Union: Laguna Olomega (KU 184574). M(1ra/an: 12 km NE Perquin, Canton, El Zancudo (KU 184578-609); grounds of Hotel Perquin Lenca, Perquin, 1150 m (KU 290030). San Salvador: 1 mi NW San Salvador (KU 42613-24); Instituto Tropical (KU 61871-74); 16 km E San Salvador, 2880 ft (KU 97494); 3 km SE llopango. Canton A.sino (KU 184575-77, 184670-74); San Salvador, Cuidad Universitaria (KU 184612-58, 184715-16). Santa Ana: Rancho San Jose, 800 m (KU 65372). San Vicenit;: 1 km E El Carmen (12 km E Cojutepeque), 2620 ft (KU 97497); 6.5 km E El Carmen (17.5 km E Cojutepeque), 2700 ft (KU 97498). Honduras: Ciiolltra: 6.8 mi S Prespire, 380 ft (TNHC 31461); Choluteca (USNM 167195); 1.5 km NW Choluteca, 170 m (KU 65375-76); San Francisco de Colon, 1 130 m (KU 65374). El Paraiso: 4.4 mi SW Santa Maria, 1960 ft (KU 97512-13). Francisco Mor,-\/an: 10.3 mi S La Vente, 530 ft (TNHC 31454); vie. of Tegucigalpa (LSUMZ 24133-34). Ckacias A Dios: Rus Rus, 60 m (USNM 547977, 547980); Mocoron (UTA-CJF 1883-84). Olancho: 1 km NW Catacamas (LACM 47973-74, 21584-89); Catacamas, 460 m (KU 194214); ca. Dulce Nombre de Culmi, 600 m (KU 194213). Valle: 1 mi E Goascoran (L.A.CM 47975-77); 3 mi E Goascoran (LACM 47978-82); 5.2 mi SE Jicaro Galan, 250 ft (TNHC 31456, 31459). Mtxico: Chiapas: road from Tapachula to Puerto Madero. Oaxaca: O.I mi S jet 185 & 190, on 185 (LACM 38160-61 ); Juchitan (USNM 51 175); 4 mi S Juchitcin, 120 ft (KU 974.34-42,); 1.5 mi N Juchitan, 120 ft (KU 97443); 3 mi N Juchitan, 120 ft (KU 97444-17); 0.5 mi S Juchitan, 140 tt (KU 97448-50); 5 mi N Juchitan, 110 ft (KU 97451-62); Juchitan, 9 mi 1-; jet hvvys 190 and 1S5 (TNHC 30968); Hwy 18,5, 14.2 mi N jet Hvvy 190 ( INHC 5,3682, 53684);) mi NW Zanatepec (TNI IC 31 .338); 20 mi W Zanatepec (TNI IC 27161 ); 3 mi W Zanatepec (TNHC 271 65); 10 mi W Zanatepec (TNI IC 27292); Zanatepec (TNHC 27300); 17 mi E Tehuantepec (TNHC 27282, 27284, 27286, 27288, 27290, 27305); 40 mi E Juchitan (TNHC 27291 ); edge of Tepanfepec (TNHC 2729,3,27295,29297), Nicarac;ua: Granada: shoreofl.agode Nicaragua, ca 2 mi from Granada (LACM 67585). Man.-u.la: Managua, S shores of Lake Managua (LACM 28165-69); Los Kobles (LACM 37957-,5S, KU 17.3951-53, 173955-.56); 2-5 km STipitapa(KU 173958), Rivas: Nandaime, 400 ft (KU 97547). Zi i aya Noktl: Leicus Creek at La Tronquera (NIPCX) Lumber Plant), .56 mi NW Puerto Cabeza (l,At:M 14580S); Slilma Sia, 16 km SE Waspam, Comarca de El Cabo (LACM 145813). Bufo cyclailen: Mfxico: Ck trrtro: near Palo Blanco (FMNH 99682, 99686, UIMNT I 248,34-,38); Xaltianguis (UIMNH 24833); Agua del Obispo, KM 350-351 (UlMNl I 248,39); Agua del Obispo, KM 357 (UIMNH 24840); 4 km bevond Agua del Obispo (UIMNH 24841^5); near Agua del Obispo (FMNH 99684-85, 99687-90, 105394, 107984-91, USNM 11548283); Agua del Obispo (UIMNH 24846, 24848-50, 24875-76, 57143); Agua del Obispo, 980 m (KU 86672-73); Agua del Obispo, 2900 ft (UMMZ 115357 1 6 specimens]), Oax.aca: 3 mi S Putla (UIMNH 57144-51), Bufo ibnrrni: Guatemala: Baia Vi hai'az: ChiIasco(UTAA-47567-69); circa 5 km S Chilasco, 1800 m (MVZ 143379); 8 km ESE Chilasco, Finca Miranda, 6500 ft |198] m| (MVZ 150931); 50,2 km NW El Rancho (UTA A-=.016); CA-14, 29,0 mi |46,7 km] NW El Rancho (UTA A-.5049); CA- 14, 50,2 km NW El Rancho (UTA A-5015): 4,8 mi (7,7 km] SSE Puruiha, Plantacion Santa Teresa (UTA A-74 1 7); 9, 1 mi (14,6 km] W Salama (by road to Puruiha) (UTA A-7432); 2,4 mi ]3,9 km] W Puruiha (UTA A-8502-<')7); 3.5 mi ]5.6 km] W Puruiha (UTA A-30495 larvae); 3.2 km WNW Puruiha (UTA A-17117-18); 3.5 km W western Puruiha turnoff (UTA A- 17242- 1 7244); 2.7 km W western Puruiha turnoff (UTA A-I7245); 3.8 km W Puruiha, 1536 m (KU 186288-303); 7,7 km SE Puruiha, 1615 m (KU 186304); 3,8 km W Puruiha, 1524 in (KU 190067); 4.2 km W Puruiha, 1524 m (KU 190068-70); 3,4 km W Puruiha, 1524 m (KU 190071); 2,0 km W Puruiha (UTA A-38145^9); Hwy CA-17 between El Rancho and Coban, km 126 (UTA A-43977-78); 1 km S San Geronimo (UMMZ 84083), El Quich6: Jovabaj (KU 186305); La Primavera, between Sacapulas and Santa Cruz Quiche, 6600 ft ]2012 m] (UMMZ 126307), Gi .ufmai a: Amatitlan (UTA A-.38144); 1 1,2 km SW Guatemala City 4600 ft [1402 m] (KU 97,595-609); 21 km SW Guatemala City, 4480 ft 11366 m] (KU 97610-19]; Guatemala Citv, /one 10, 4820 ft ]1469 m] (TNHC 31384, 31387, 31390, 31392, 31395, 31399, 31401-02, 31405, 31408, 31416-20, 31422, 31426, 31430-33); Guatemala City, between zone 5 and zone 15, km 2,5 (UTA A-25824): E side Guatemala City, zone 16, 1 km N Vista Hermosa III on road to Santa Rosita (UTA A-25825-32); Santa Catarina Pinula, San Miguel Buena Vista, 1700 m (UTA A-43951, UTA A-47570-74); Guatemala City zone 15, Vista Hermosa IIL 1510 m (UTA A-28959-60);Parque San Jorge Muxbal 1850 m (UTA A-32993). HuEHLErtNANt.o: Aguacatan (UMMZ 120046); 2 km NE Aguacatan, 1640 m (UMMZ 120047-48); 2,8 km E Aguacatan, 1600 m (KU ^8412-13); Huehuetenango, patio of Casa Maryknoll (UMMZ 124382); 22 km SSW Huehuetenango (KU 116959); 3 km W I luehuetenango, 6100 tt ]1859 m] (TNHC 29452-57); at La Libertad, 1700 m (MVZ 143343-57); San Pedro Necta (UMMZ 130059 larvae); circa 1 km F San Pedro \ecta, 1615 m (UMMZ 119352). Jaiara: Jalapa (TNHC 33666-72); 8.5 km NW Jalapa (INHC 31442); 7.5 km WSW Jalapa, on road to Miramundo (UTA A-,391I4 larvae); Jalapa-Miramundo rd, at km 101 (UTA A-,381 18); Falda Oeste Volcan Jumav (UTA A-47,565); 1,6 mi [2,5 km] NE El Mojon (UTA A-38127-40); 0,7 nii'l 1 . 1 km] NE EI Mojon (UTA A-38141 ); Aserradero San Lorenzo (circa 12 air km NNE Jalapa), 1725 m (UMMZ 108000, 106806 ] 10 specimens], 106807 ]3 specimens], Proc.reso: Finca Bucaral (UMMZ 10(iS08, 1395 16 larvae), S.u aiii-lqui/: 3 km W Dueiias (TNHC 31378); 1,3 mi ]4,4 km] W Finca San Rafael Urias at Dueiias (TNHC 31344, 31379-80); San Antonio (CAS 70826-27); Volcan Agua (CAS 70719-825), Honduras: Kiiiiii. a: water supply area for La Fsperanza (LACM 45247—18) 1,5 mi NE La Esperanza (LACM 47992-96); I mi NE La Esperanza (LACM 47998); La Esperanza (LACM 47998-48004); 25, 7 km NW La Esperanza, 1340 m (USNM 523689-93); 18,1 km NW La Esperanza, 1740 m (USNM 523694-96); 8,7 km NW La Esperan/a, 1540 m (USNM 523697); Zacate Blanco, 2020 m (KU 194220-21 ); ca, Miguelito, 60,3 km SF Gracias (Depto. Lempira), 1310 m (KU 209250, 2092,53), Llmrira: Frandique (LSUMZ 46432, 497.38); above Villa Verde, 1280 m (KU 209240), OtxiTEPEOiiL: 12,5 mi E Nueva Ocot..peque (LACM 47983-85); 6.5 mi E Nueva Ocotepeque (LACM 47986-91 ); ■•14'29.48'N, 88-46.83'W" (UTA A-52960, ,53662); Belen Gualcho, 1470 m (KU 194208); El Chaguiton, 1870 m (KU 194209-12; USNM .523712-13); Fl Volcan, 1760 m (USNM 523714-18), Bufo /lisiunus: Mexico: Miciioaian: 9 mi on rd between Ri'o Marque/