~ ry The Libre -_ scientific Papers Natural History Museum The University of Kansas 24 September 199 Number 3:1—-41 Frogs of the Hyla armata and Hyla pulchella Groups in the Andes of South America, with Definitions and Analyses of Phylogenetic Relationships of Andean Groups of Hyla By . WILLIAM E. DUELLMAN, IGNACIO DE LA Riva!, AND ERIK R. WILD? Division of Herpetology, Natural History Museum and 8 2 Department of Systematics and Ecology, The University of Kansas s = Lawrence, Kansas 66045-2454, USA. oa ou 3 CONTENTS BS 2S) ANVBISTIRUN Go sete sce oS 9 Pan PARP OR eS eh nee EEE ae EOE 2 Bg TROTERSS IMATINIB Eee ArT PRESSE 2 28 TINSINR @)1D) Gala @) Ngee eee nae weet Pd, AO ee oes Soh ati E Sast me beeeee Re? 3 2 z INGKNOW BED MENTGU ESE ee bon tee lity AE Ee eek Where 6 Oe eee 3 5 fee IMUATPIBIRAVAILS /AINJID) IMDBEIEG KO) DIS) occ scscctocoancsbonsbssonscons0500¢560d604<650<00550s06005000000e0600200060n0000800000800550850050 3 5 JNINVANTIENESIIS) FH GIBUAIRVAN GANS RRS es irae cacosocece sone sass eeedecece crachneeccesecoatee ion a San eS cece SaoceocRS a eeo eS ecccooD 4 3 INFORPRIONIEMRICS eee ee eae ee aaa a cect Nee SaG aE Tact Pease ere eae ence Sab SURE RUNS dope ase ooee 4 = EXGHERNUATOIN [ORPETO LOG Verearesteees coe cea e area ae ee eTe REESE OMS S Lis Cate Se Noe nT Toe Ct Sere Saeed seseaseees 7 (GOVORATION ers RN PIN wae bat chet aea dan en ean cae eu Pasadesaesevate soot ese arescuteessotees 9 @STEOWO GY a SST EET ST SUDO TEE eas ON Tau eon eda na aan Sue Tas taese Sasso Se yokc can eeunesuek ae 9 PND VERTISENIE Nila GATES ee sees secse eas coe crerecnaea ees ete eae one ance dase sEALT Re ate Seneeteance eacevereetacaet 10 SIVA) OLS eae eee et ea a SE eee ne ees Pacers Nee aea eee Rete SB SEGTTL Hoe sake oa carotaees SETAE Suenos 10 ANINIDIBVAINT GIROUUIES (OU BE BA A \ cco seosencpbotireb deseobodeguscossdecacecdbeoc0eee Jed sic sete aa5e ced con Soa SO Rae RE SOp A ada 2 TE BALA ARINUAIBA (ARC TUT. ecnnacnaneceacactbecbreo0000 ibs cSEBEsn0neBUEbeo ENS EECIGDS0C EC TEGO GEOR AS RCE DXCODOCS DGCC ISR RELOR GOSS 12 TRIGA TRUTH CLETTIUAN GIAO TOO ss sactasbropeceoodebheeces00es0dod000c Cede eoA0 FERC GRRE CET ESR ASeSonCEEo5 CEPOLESC OR soca seo OcIEGSEEEC 15 FEINHANE OG OREN SISK GROUPE arrcee ec eeeeee eNeaae aati Stescattegsererssrneesnseeesces 30 JERGLA CIELO IROBIIEA (GROUT? ccccoonsococcnoooessoueonde6se09060000068908000500099000005549850066000408 00 3EE00 IDS UBD OSOODAGCOGASSODD 31 EL HANE OLUINIBIANTAN GROUP eee ereer ce eee Tee ee eee aneanctescteenincee sate acne 31 IBRALA CARAGGIEINGIG (GIRO IOI soconscacosoas0ac00d0ocacoosoEoJeRdAedes80020000006808e000000eT 050000 500SHN5d5HHADDIODoaESTEEEsOo%d00500 32) Present address: Museo Nacional de Ciencias Naturales, J. Gutiérrez Abascal 2, 28006 Madrid, Spain. : Present address: Department of Biology, University of Dubuque, 2000 University Avenue, Dubuque, Iowa 52001, USA. ISSN No. 1094-0782 © Natural History Museum, The University of Kansas 2 Screntisic Papers, NATURAL History Museum, THE UNIVERSITY OF KANSAS FLY EA NAB TALIS! GOROUD ccs crccssessc ora sos es soos SSS OS Seo COE ESSE S Sa SUTRAS CONTE ONC CST EEN eo 32 FAY ACBARINGPY GION) GROUP eccssorsseaec eso oos oes e See TES NTE OO ENS L ET TOE ae eign Pong 32 [PAE YA LOG] EUNTEMEN CG TRUBULYAUIKOUN ST BIDS} oot cos sacscnoneacnesee écosbboneboccobo debe soeanood Sednasonsbnaabbenbbacedonb6Ko0 33 RELATIONSHIPS AMONG ANDEAN GROUPS OF FIYLA ....0....c00c0cccoesscosenscsosesceessesscessccctssccsessssesseness 33 RELATIONSHIPS OFMIEEy lV AVPUINCEEN MAN GRO Ulzreerscsne see eeee ste eae eee eee 36 TDISGUSSTON ress ots esc tae ena RE es Ree 38 TT RATE GTR se ack ee cae ce creat eae eno ee a ic eR EU re ED RRO nv a ne 38 NPR END DXS SPECIMENS) ROCAIVIINED eee eee erate neeae e e 40 ABSTRACT Andean hylid frogs of the genus Hyla are reviewed. Eight groups of Hyla recognized in the Andes are defined; the species in two of these groups are treated in detail. The large H. armata, characterized by hypertrophied forelimbs and clumps of nuptial spines on the prepollex and humerus in males, and a stream-adapted tadpole with a LTRF up to 15/17, is the sole recognized species in the H. armata group, Accounts of this species and six species (one new) in the H. pulchella group include diagnoses, descriptions of adults, larvae, advertisement calls, and information on ecology, and distribution. Hyla ocapia Andersson, 1938, is placed in the synonymy of H. albonigra Nieden, 1923; H. callipleura Boulenger 1902, is placed in the synonymy of H. balzani Boulenger 1898, and H. pulchella andina Muller, 1924, is recognized as a distinct species. The H. pulchella group is characterized by hypertrophied forelimbs and projecting prepollical spines in adult males, and generalized, stream-adapted tadpoles. These characters are shared by frogs in the H. circumdata group in southeastern Brazil, and the two groups are combined herein into the H. pulchella group. The presence of sharp, projecting prepollical spines is a character shared with several groups of Hyla in the neotropical lowlands. Phylogenetic analysis reveals that the H. columbiana and H. labialis groups are sister groups; this clade is the sister of a clade containing several groups of Hyla having a diploid number of 30 chromosomes and inhabiting Andes (H. garagoensis Group) and the neotropical lowlands. Frogs in the H. bogotensis group are characterized by the presence of a mental gland in males; this derived character is shared with H. granosa in the Amazon Basin and the Guiana Region. No unique synapomorphies exist for the frogs assigned to the H. larinopygion group. Too few data are available for Hyla chlorostea to associate it with any lineages of Hyla. Key words: Hylidae; Hyla armata, bogotensis, chloroseta, columbiana, garagoensis, abialis, larinopygion, and pulchella groups; taxonomy; new species; phylogenetic relationships; South America; Andes. RESUMEN Se revisan los hilidos andinos de 1 género Hyla y se definen ocho grupos de especies. Las especies de dos de ellos son tratadas en detalle. Hyla armata es el unico miembro de su grupo Los adultos de este especie son grandes, caracterizados por tener brazos hipertrofiados y grupos de espinas nupciales en el prepollex y humero en los machos; las larvas estan adaptadas a corrientes rapidas, y cuya formula de filas de dientes labiales es de hasta 15/17. La informacion aportada sobre esta especie y otras seis especies (una nueva) de grupo de H. puchella incluye diagnosis, descripciones de adultos, larvas, y Ilamadas de advertencia, y datos sobre ecologia y distribucion. Hyla ocapia Andersson, 1938, es puesta en la sinonimia de H. albonigra Nieden, 1923; H. callipleura Boulenger, 1902, es puesta en la sinonimia de H. balzani Boulenger, 1898; H. pulchella andina Miller, 1924, es elevada a rango especifico. El grupo de H. pulchella se caracteriza por tener brazos hipertrofiados, machos con espinas sobresalientes en el prepollex, y larvas generalizadas, adaptadas a aguas corrientes. Estos caracteres son compartidos con las ranas del grupo H. circumdata, del sudeste de Brasil, por lo cual ambos grupos son combinados en el grupo de H. pulchella. La presencia de espinas sobresalientes y afiladas en el prepollex es un caracter compartido por varios grupos de Hyla en las tierras bajas neotropicales. Los analisis filogenénticos efectuados revelan que los grupos de H. columbiana e H. labialis son grupos hermanos. Este clado es el grupo hermano de un clado que contiene varios grupos de Hyla con numero diploide de 30 cromosomas y que habitan los Andes (grupo de H. garagoensis) y en las tierras bajas neotropicales. Las ranas del grupo H. bogotensis se caracterizan por la presencia de una glandula mentoniana en los machos; este caracter derivado es compartido con H. granosa, una especie de la Cuenca Amazonica y la region de las Guayanas. No hay sinapomorfias unicas para las ranas asignadas al grupo de H. larinopygion. Se dispone de muy poca informacion sobre H. chlorostea como para asociarla‘con algun grupo de Hyla. Palabras claves: Hylidae, Grupos de Hyla armata, bogotensis, chloroseta, columbiana, garagoensis, labialis, larinopygion, y pulchella; taxonormia; nueva especie; relaciones filogenéticas; Sudamérica; Andes. ANDEAN GROUPS OF HYLA 3 INTRODUCTION Many species of frogs of the immense, paraphyletic ge- nus Hyla in South America have been placed in several phenetic groups (Frost, 1985). These include the H. bogotensis Group (Duellman, 1972), H. columbiana Group (Duellman and Trueb, 1983), H. garagoensis Group (Kaplan and Ruiz, 1997), H. labialis Group (Duellman, 1989), and the H. larinopygion Group (Duellman and Hillis, 1990). Nine nominal species from the Andes of Argentina, Bolivia, and Peru have not been associated with any group; these spe- cies are H. albonigra Nieden, H. armata Boulenger, H. balzant Boulenger, H. charazani Vellard, H. chlorostea Reynolds and Foster, H. callipleura Boulenger, H. marianitae Carrizo, H. melanopleura Boulenger, and H. ocapia Andersson, although Frost (1985) placed the latter without comment in the Hyla ocapia Group. With the exception of the description of the tadpole of H. armata (Cadle and Altig, 1991), morphology of the tadpole of H. pulchella andina (Lavilla and Fabrezi, 1987), additional distributional records from Bolivia (De la Riva, 1990; Reynolds and Foster, 1992), and descriptions of calls of some species (Barrio, 1965; Cadle and Altig, 1991; Marquez et al., 1993), no new information has been pub- lished on any of these species since their original descrip- tions. Casual examination of specimens of these species revealed that all, except H. armata and H. chlorostea, have projecting prepollices that are sexually dimorphic; this character is shared with H. pulchella Duméril and Bibron, and presumably related species in southeastern Brazil, northeastern Argentina, and Uruguay, with isolated popu- lations in the Cordillera de Cordoba and in the Andes of northern and central Argentina and central Bolivia (Bar- rio, 1965, Lutz, 1973, Cei, 1980). Independent field work by De la Riva in Bolivia, and by Duellman and his associates in Argentina, Bolivia, and Peru has resulted in the acquisition of many specimens of these frogs and data on their ecology and life histories. The purposes of this paper are to: (1) present analyses of data on the external morphology and osteology of adults; (2) describe the tadpoles and advertisement calls; (3) syn- thesize all available information pertaining to the tax- onomy, ecology, and distribution of the species; (4) define the Hyla armata and Hyla pulchella groups; and (5) attempt to determine the phylogenetic relationships of the groups of Hyla in the Andes. ACKNOWLEDGMENTS For the loan of specimens, we thank Erick Ahlander (NHRM), José Cabot (EBD), Jonathan A. Campbell (UTA), Gustavo R. Carrizo (MACN), Javier Castroviejo (CET), Barry T. Clarke (BMNH), Guilano Doria (MSNG), Darrel R. Frost (AMNH), W. Ronald Heyer (USNM), and Anne M. Olher (MNHN). Duellman is grateful to Jesus Cordova (MHNSM), Alice G. C. Grandison (BMNH), Jean Guibé (MNHN) and the late Walter Hellmich (ZSM) and Greta Vestergren (NHRM) for the provision of working space in their respective collections. We thank Joseph R. Mendelson Ill for his assistance with the multivariate analysis and Hélio da Silva for his aid in the phylogenetic analyses. Amy Lathrop executed the drawings comprising Figures 4, 7, and 16. and John E. Simmons made the photographs com- prising Figures 5 and 8. Michael B. Harvey provided data on frogs collected by him in Bolivia, as did Lily O. Rodriguez for frogs collected by her in Peru. We are grate- ful to John D. Lynch, Robert P. Reynolds, and Linda Trueb for critically reading the manuscript. Duellman’s visits to European museums was made pos- sible by a grant (5063) from the Penrose Fund of the Ameri- can Philosophical Society and his field work was supported by a grant from the National Science Foundation (DEB- 7401998); he is grateful to his field companions—David C. Cannatella, the late Ismael Ceballos, Dana T. Duellman, Thomas H. Fritts, Oscar Ochoa M, John E. Simmons, and Linda Trueb. Furthermore, David C. Cannatella and Tho- mas J. Berger collected critical material in Boli- via.Appropriate permits were issued for Peru by Luis J. Cuento Aragon and Armando Pimentel Bustamente of the Direccion General Forestal y de Fauna. Logistic support in Peru was provided by Oscar Ochoa M. and in Argen- tina by Raymond F. Laurent. The research was completed as part of a project on pat- terns of speciation and distribution of anurans in the Andes supported by the National Science Foundation (BSR- 8805920; W. E. Duellman, principal investigator). De la Riva is indebted to the Asociacién Amigos de Donana for fund- ing field work in Bolivia and to personnel at the Museo de Historia Natural Noel Kempff Mercado and the Colecci6n Boliviana de Fauna for logistic support in Bolivia. MATERIALS AND METHODS Specimens used in this study are referenced by their museum catalogue numbers prefaced by the institutional codes given by Leviton et al. (1985) with the addition of the following: CET = Centro de Estudios Tropicales, Sevilla, Spain; EBD = Estacion Biologica Donana, Sevilla, Spain; MHNSM = Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima Peru; MNHNB = Museo Nacional de Historia Natural de Bolivia, La Paz, Bolivia*?; MNK = Museo de Historia Natural Noel Kempff >The collections in the MNHNB have been combined with the collection in the Instituto de Ecologia; the new collection is known as the Coleccion Boliviana de Fauna (CBF). This occurred subsequent to our examination of specimens in the MNHNB, and the catalogue numbers that we give for specimens in that collection probably have been changed. 4 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Table 1. Sexual dimorphism in snout-vent length (SVL) in Hyla armata and Andean species in the Hyla pulchella Group. n = males/females; in the columns of the sexes, the mean + one standard deviation is followed by the range in parentheses. Sexual dimorphism is the mean SVL of males divided by the mean SVL of females. Species N Males Females Dimorphism Hyla albonigra 9/11 50.5 + 3.15 (47.0-56.0) 54.9 + 4.06 (49.3-64.7) 92.0 Hyla andina 98/59 46.5 + 7.72 (34.6-57.6) 49.8 + 6.06 (37.0-62.7) 93.4 Hyla armata 28/22 59.6 + 4.82 (47.5-68.5) 64.3 + 5.93 (47.5-74.5) 92.7 Hyla balzani 61/8 40.4 + 3.66 (33.3-49.9) 48.5 + 2.73 (44.4-52.3) 83.3 Hyla marianitae 26/6 47.6 + 4.96 (36.5-56.8) 51.4 + 4.57 (45.0-56.5) 92.6 Hyla melanopleura 1/3 43.6 47.1 (46.4-47.7) 92.6 39.5 + 3.53 (36.2-50.4) 49.0 80.6 Hyla palaestes 25/1 Mercado, Santa Cruz de la Sierra, Bolivia. Specimens ex- amined and their localities and catalogue numbers are listed in the Appendix. Measurements of frogs and and tadpoles were made to the nearest 0.1 mm with dial calipers in the manner de- scribed by Duellman (1970). Forearm length was measured from the base of the thenar tubercle to the proximal tip of the radioulna; forearm breadth was measured at the wid- est part of the forearm. Snout-vent length is abbreviated SVL throughout. Stepwise discriminate function analyses were accomplished using the BMDP Statistical Software (Dixon, 1981). Webbing formulae follow the methodology of Savage and Heyer (1967) as modified by Myers and Duellman (1982). Dried skeletons were prepared by hand; cleared and stained specimens were prepared in the man- ner described by Dingerkus and Uhler (1977). Osteologi- cal terminology follows Trueb (1993). Tadpoles were staged according to Gosner (1960); descriptions generally follow those of Cadle and Altig (1991). The labial tooth row for- mula is abbreviated LTRF. Calls were recorded using por- table tape recorders (Sony WM D6C and Sennheiser di- rectional microphone model Me-80 or Uher with Uher mi- crophone) and analyzed by means of the Canary 1.2 soft- ware (Charis et al., 1995) on a Macintosh computer. Phylo- genetic analyses were accomplished using the DELTRAN option in PAUP software (Swofford, 1991). Credits for the color photographs in Figure 3 are: DCC = David C. Cannatella, IDIR = Ignacio de la Riva, WED = William E. Duellman. ANALYSIS OF CHARACTERS The following discussion includes only Hyla armata and those species in the Hyla pulchella Group that inhabit the Andes. For comments on the characteristics of extra- Andean species of the Hyla pulchella Group, see Phyloge- netic Relationships. MOorPHOMETRICS Mean SVLs of males vary from 39.5 mm in Hyla palaestes to 59.5 mm in H. armata, and the means of females vary from 47.1 in H. melanopleura to 74.5 mm in H. armata. Among those species for which series of adults of both sexes are available, sexual dimorphism in SVL is slight; SVL in males is more than 90% of that in females, except in H. balzani and H. palaestes (Table 1). Strong sexual dimor- phism exists in the relative breadth of the forearm, which is massive in males and normal in females. Measurements of other external features are given in Table 2 In order to determine morphometric distinctness, we performed a stepwise discriminate function analysis on measurements taken from 49 specimens of Hyla armata and 304 individuals of seven species of frogs in the Hyla pulchella group (sexes combined)—Hyla albonigra (n = 15), H. andina (n = 156), H. balzani (n = 69), H. “NANT Gi SB), ISI, melanopleura (n = 3), H. palaestes (n = 26), H. pulchella (n = 3). The two species with small sample sizes (H. melanopleura and H. pulchella) were treated as unknowns and, thus, were not used in determining the discriminate function. Twelve morphological variables were log transformed and in- cluded in the stepwise analysis, which identifies those variables that maximize separation of group means. The twelve variables are SVL, tibia length, foot length, head length, head width, interorbital distance, internarial dis- tance, eyelid width, eye diameter, tympanum diameter, eye-nostril distance, and total number of vomerine teeth. Stepwise discriminate function analysis was terminated at a model that included all 12 variables as significant (F > Fo0s15,330) = 2.2388). The approximate F-statistic rejected the null hypothesis of equality among means (P < 0.001). Mul- tiple F-tests of pairwise comparisons among all groups using Bonferroni protection for multiple tests (a = 0.001 for a = 0.05; Foooit12,330) = 2.8408) indicated that the twelve variables were able to separate all group means effectively except for Hyla melanopleura and H. albonigra (F = 1.47), H. melanopleura and H. andina (F = 1.70), H. melanopleura and H. marianitae (F = 2.22), H. melanopleura and H. pulchella (F = 2.57), and H. melanopleura and H. palaestes (F = 2.64). Overall jackknifed classification of individuals was 81% correct (not including individuals of species treated as ANDEAN Groups OF HYLA 5 Table 2. Morphometric data for specimens of Hyla armata and Andean species of the Hyla pulchella Group. In each section, first line is the mean + one standard deviation; second line is range. H. andina H. melanopleura H. armata H. albonigra H. balzani H. marianitae H. palaestes Character Sex 28 OO, 22 99 9 OG, 11 99 98 OG, 59 99 61 OG, 8 99 26 OG, 6 90 14,399 25 00,19 Tibia length OS 31.6+2.44 26.9 +2.52 24.5 + 2.95 21.6+2.11 26.1 + 2.83 21.6 21.9 +£1.77 26.1-34.7 23.4-31.0 173-3202. 17.8-27.9 19.2-31.8 21.6 19.1-26.9 QQ): 34.22.16 25.7 + 1.94 25.8 + 3.08 26.0 + 1.00 26.6 + 2.57 24.7 23.7 27.2-39.0 23.6-29.9 18.2-32.1 24.5-27.4 22.5-29.4 24.2-25.2 23.7 Foot length OS 28.3+42.63 23.1 42.21 21.7 + 2.90 18.0 + 2.37 ZARSEEDES9 WA 17-3} 1855, 21.8-32.0 19.6-26.5 13.3-29.5 14.6-29.4 15.4-24.9 W7E7. 14.8-22.1 eg. 31.1 +3.42 25.7 + 1.94 22.8 + 3.24 2210 FE 155 22.3 42.19 20.6 17.4 23.6-37.1 23.6-29.9 16.2-29.0 19.2-23.7 19.2-24.7 20.5-20.7 17.4 Head length OS 19.6+2.36 15.8 + 1.47 15.2 + 1.44 14.3 + 1.03 16.3 + 1.60 14.4 14.1 £0.97 15.8-29.7 13.2-17.2 11.6-18.9 11.9-17.3 13.0-19.8 14.4 13.0-16.8 eg «20.2 + 2.16 16.6 + 1.19 16.2 + 1.97 16.6 + 1.07 7.24 1.66 15.5 15.1 17.0-23.5 14.2-18.1 12.1-21.6 15.3-18.3 14.8-18.9 15.4-15.6 15.1 Head width OS 20.3+1.46 17.4+1.93 15.0 + 1.82 13.9 + 1.27 16.3 + 1.65 14.4 14.1 + 1.38 16.5-23.0 15.5-20.8 11.1-19.5 11.8-17.7 13.0-20.3 14.4 12.5-18.5 QQ. 21.8 + 1.70 18.5 + 2.61 15.9 + 2.25 16.6 + 0.87 17.141.18 15.5 14.8 17.8 + 24.2 12.8-22.2 11.7-20.7 15.1-18.0 15.5-18.5 15.4-15.6 14.8 Interorbital distance CoS 6.0+0.57 4.9 + 0.63 4.4 + 0.45 4.5 + 0.47 5.4 + 0.53 4.2 4.3 + 0.62 4.8-7.1 4.2-6.3 3.5-5.6 Si7=57, 4.3-6.5 4.2 3.4-6.2 C9 = 6.640.87 5.0 + 0.45 4.6 +0.59 5.0 + 0.28 5.3 + 0.59 4.6 4.5 5.3-8.0 4.5-5.9 3.6-6.0 4.7-5.5 4.5-5.9 4.5-4.7 4.5 Eye diameter OS 604088 5140.80 4.3 + 0.50 4.3 + 0.46 4.6 + 0.52 3.9 4.4+0.24 3.8-7.3 4.3-6.5 3.1-5.6 3.4-5.5 3°7=9:9 3.9 3.8-4.7 °9.—Cs«6.7 £0.81 5.4 + 0.59 4.4 + 0.59 4.7 + 0.45 4.9 + 0.72 5.0 4.5 5.1-8.4 4.9-6.4 3.2-6.1 3.9-5.3 S959 4.8-5.1 45 Tympanum diameter OS 2.1+0.41 2.7 + 0.20 2.6 + 0.32 2.2 + 0.33 2.5 + 0.36 2.4 2.4 + 0.26 1.43.4 215-341 1.9-3.2 1.6-3.2 1.8-3.1 2.4 U9=2:9 QQ. =. 2.2 + 0.46 3.0 + 0.25 2.8 + 0.48 2.7 + 0.22 2.9 + 0.32 3.0 27 1.6-3.3 2.6-3.4 7319 2-5-2.9 2.4-3.3 2.7—=3:3 27 unknowns). Hyla armata was classified with 96% correct. Ayla andina, H. marianitae, and H. palaestes were all classi- fied with over 80% correct, whereas H.balzani and H. albonigra were lower, with 62% and 40% correctly classi- fied, respectively. Of the 67 cases misclassified, 50 can be considered borderline cases (posterior probability of group membership < 0.75). Of the nonborderline misclassifica- tions, H. andina and H. albonigra had the greatest frequency with seven individuals of H. andina classified as H. albonigra and one H. albonigra classified as H. andina. Some individu- als of H. balzani and H. marianitae also were confused, with three H. balzani classified as H. marianitae, and one H. marianitae classified as H. balzani. Other nonborderline misclassifications include two H. palaestes classified as H. marianitae, two H. balzani classified as H. palaestes, and one H. balzani classified as H. andina. Classification of individu- als for the two species treated as unknowns placed all three H. pulchella with H. andina (two with posterior probabili- ties of group membership > 0.75). Hyla melanopleura was classified as H. andina (one with high and one with low posterior probability of group membership) or H. albonigra (low posterior probability of group membership). The first two canonical axes displayed 95% of the total dispersion (Fig. 1), both with eigenvalues greater than the mean; other axes had eigenvalues lower than the mean and contributed little to the cumulative proportion of the total dispersion. Values for canonical variables evaluated 6 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS H. palaestes (@) ‘O H. balzani @) H. marianitae H. albonigra H. pulchella mann H. andina WW H. melanopleura sf Canonical Variable 2 -4 By A 0 1 2 3 4 5 6 Canonical Variable 1 Fig. 1. Plot of canonical discriminant scores for Hyla armata and species of the Hyla pulchella Group. Closed symbols indicate group mean scores. All individual scores for H. melanopleura (inverted triangles) and H. pulchella (squares) are plotted as open symbols. 95% confidence inter- vals around mean scores are provided for the six other species (the 95% confidence interval for H. andina is smaller than the solid symbol; for the group mean). at group means indicated that the first canonical axis was able to separate most groups. The least differentiated on the first axis were H. andina and H. melanopleura, although these are separated by the second axis. Examination of the standardized (by pooled within-group variances) coeffi- cients for canonical variables shows that tympanum di- ameter was the most important variable on the first axis. The second axis had SVL as the most important variable, followed by foot length, head width, and eye-nostril dis- tance. These standardized coefficients for canonical vari- ables were: Canonical Axis I (Canonical Axis II) = SVL: — 0.40 (1.45), foot length: 0.07 (-0.88), head length: —-0.01 (0.15), head width: 0.57 (0.75), interorbital distance: 0.48 (0.46), internarial distance: 0.34 (0.33), eyelid width: 0.15 (0.05), eye diameter: 0.29 (0.08), tympanum diameter: -1.14 (0.21), eye-nostril distance —0.03 (0.69), VT: 0.40 (0.42). In summary, the discriminate function model involv- ing 12 variables could discriminate Hyla armata, and most of the species of the Hyla pulchella Group. The function was unable to differentiate group means of H. melanopleura from several other species (H. albonigra, andina, marianitae, palaestes, and pulchella). This is likely a result of the small sample size of H. melanopleura. The ability of the model to classify was not especially high (81%); however, most misclassified individuals were borderline cases and may represent outliers, such as young individuals. Of those misclassifications that were not borderline cases, nearly half involved misclassified individuals of H. andina and H. albonigra. The first two axes displayed most of the varia- tion among groups (95%). The first, in which tympanum diameter was by far the most important variable, showed H. armata to be distinct with the smallest tympanum. The first axis was not effective in discriminating H. andina from H. melanopleura. The second axis, in which SVL was the most important variable, improved the ability to distin- guish these two species. In order to determine morphometric differences among individuals of Hyla andina from different regions and to ascertain the possible existence of cryptic taxa, especially in Bolivia, stepwise discriminate function analysis was performed on measurements taken from 156 specimens from four regions (sexes combined): Bolivia (n = 84), and the provinces of Catamarca (1 = 35), Jujuy (1 = 24), and Tucuman (1 = 13) in Argentina. The same 12 morphologi- cal variables used in the analysis of the Hyla pulchella Group were log transformed and included in the present stepwise analysis. Stepwise discriminate function analysis was ter- minated at a six variable model that included snout-vent length, foot length, interorbital diameter, internarial dis- tance, eye-nostril distance, and number of vomerine teeth as significant (F > F, 9.5 47) = 26656). The approximate F- statistic rejected the null hypothesis of equality among means (P < 0.001). Multiple F-tests of pairwise compari- sons among all groups using Bonferroni protection for multiple tests (a = 0.001 for a = 0.05; Berean = 3.986) indi- cated that the six variables were able to separate all group means effectively except those of Tucuman and Jujuy (F = 1.33), and those of Tucuman and Bolivia (F = 3.07). Overall jackknifed classification of individuals was 56% correct. Specimens from Catamarca were correctly classi- fied most frequently (69% correct), followed by those from Bolivia (60% correct). Specimens from Jujuy and Tucuman were classified with much lower success, 38% and 31%, respectively. The failure of the function to classify correctly individuals is even more evident when examining only those correct classifications that had high posterior prob- abilities of group membership (> 0.75): Catamarca 26%, Bolivia 17%, Tucuman 8%, and Jujuy 0%. Furthermore, there appears to be no evident pattern of misclassification among any of the localities and of the 69 cases misclassified; only five were not borderline cases (posterior probability of group membership > 0.75), which emphasizes the in- ability of the function to discriminate most individuals of the four groups. The first two canonical axes displayed 96% of the total dispersion (Fig. 2). The first axis had an eigenvalue greater than the mean; the eigenvalue of the second axis was slightly less than the mean, and the third axis had a very low eigenvalue and contributed little to the cumulative proportion of the total dispersion. Values for canonical vari- ables evaluated at group means indicated that the first ca- nonical axis was able to separate populations from ANDEAN Groups OF HYLA V Canonical Variable 2 -4 -3 -2 -1 0 1 2 3 Canonical Variable 1 Fig.2. Plot of canonical discriminant scores for individuals (closed symbols; n = 156) of Hyla andina. Open symbols indicate group mean scores. Squares = Bolivia, circles = Catamarca, triangles = Jujuy, and in- verted triangles = Tucuman, Argentina. Catamarca and Tucuman, whereas those from Bolivia and Jujuy were more similar, although these are separated by the second axis, as were Catamarca and Tucumaén . Exami- nation of the standardized (by pooled within-group vari- ances) coefficients for canonical variables revealed that snout-vent length was the most important variable on the first axis, followed by eye-nostril distance. The second axis had foot-length as the most important variable, followed by snout-vent length. These standardized coefficients for canonical variables were: Canonical Axis I (Canonical Axis II) = SVL: -2.14 (1.17), foot length: 0.72 (—1.60), interorbital distance: 0.34 (—0.44), internarial distance: 0.82 (0.70), eye- nostril distance 1.04 (0.53), number of vomerine teeth: 0.40 (-0.61). In summary, the discriminate function model involv- ing six variables was able to discriminate means of some of the geographic groups of Hyla andina. The function was unable to differentiate group means of Tucuman and Bo- livia, and Tucuman and Jujuy. The ability of the model to classify was poor (56%), with the majority of individuals classified, both correctly and incorrectly, with low confi- dence (low posterior probabilities of group membership). There was no pattern of misclassification evident among the localities. The first two.axes displayed most of the varia- tion among groups (96%). The first, in which snout-vent length was by far the most important variable, discrimi- nated Catamarca and Tucuman, but was not effective at distinguishing Bolivia and Jujuy. The second axis in which foot-length was the most important variable, improved the ability to distinguish all groups. The ability of the func- tion to distinguish the mean of Catamarca from the other groups and the relatively high success of classification of individuals from Catamarca suggest that it is the most dis- tinctive group, but the differences from the other groups is minimal. The failure to separate all group means coupled with the abysmal ability to classify individuals confidently, indicate that the morphological variables included in this analysis do not adequately distinguish the individuals from the four localities. In other words, the groups do not show sufficient morphometric differences to consider them distinct, although there are some differences in SVL. EXTERNAL MORPHOLOGY The body is moderately robust; head width and length are about equal, and the snout is short, usually rounded in dorsal view and bluntly rounded in profile (in dorsal view, nearly truncate in Hyla armata and subacuminate in H. marianitae). In transverse sectional profile, the canthus rostralis is acutely rounded to angular, and the lips are rounded. The internarial area usually is depressed slightly between protuberant nostrils that are directed dorsolaterally at a level just posterior to the anterior mar- gin of the lower jaw. The top of the head is flat, and the interorbital distance usually is greater than the width of the upper eyelid. The tympanum is round with a distinct, smooth annulus; the supratympanic fold obscures the upper edge of the tympanum and abruptly curves down- ward to a point above the insertion of the arm. In breeding males, the forearm is massive owing to greatly hypertrophied musculature. A small prepollical protrusion is evident in females, but in males of all species except Hyla armata, the prepollex is elongated and pro- trudes medially. Males of Hyla armata have a cluster of keratinous spines on the inside of the thumb and a similar cluster on the proximal ventral surfaces of the humerus. The relative lengths of the fingers are I < II < IV < Ill. Fin- gers II-IV bear distinct lateral fringes and have large, round discs. The fingers are about one-third to one-half webbed. The palmar tubercle is low, indistinct, and bifid; the th- enar tubercle is low and elongately ovoid. On both hands and feet, the subarticular tubercles are moderately large and round; the supernumerary tubercles are round to subconical, numerous proximally and readily evident in most specimens. The inner metatarsal tubercle is prominent and ovoid; an outer metatarsal tubercle is absent, except in H. armata, in which it is low and about one-fourth the size of the inner metatarsal tubercle. A tar- sal fold extends the full length of the tarsus; the fold is conspicuously elevated in H. albonigra, weak in H. marianitae, and low in the other species. The relative lengths of the toes are I < II < III < V < IV; all toes bear distinct lateral fringes and moderately large, round discs. The toes are three-fourths to four-fifths webbed. The skin on the dorsum and flanks is coarsely granular in Hyla albonigra, weakly granular to smooth in H. andina (oe) SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Hyla armata, KU 173219, male, 61.2 mm SVL. WED Hyla armata, KU 163341, female, 68.6 mm SVL. WED Hyla balzani, KU 139212, female, 46.8 mm SVL. WED Hyla balzani, KU 183425, male, 42.0 mm SVL. DCC narianitae, CET A109, male, 54.9 mm SVL. IDIR Hyla palaestes, KU 163305, male, 46.3 mm SVL. WED Fig.3. Frogs of the Hyla armata and Hyla pulchella groups in the Andes. ANDEAN Groups OF HYLA 9 and H. armata, and smooth in the other species. The skin on the throat, chest, belly, and proximal posteroventral sur- faces of the thighs is coarsely granular, and the skin on other ventral surfaces is smooth. The cloacal opening is directed posteroventrally near the upper level of the thighs and covered by a short sheath; tubercles and folds in the cloacal region are absent. The vocal sac is single, median, and subgular; the vocal slits extend posterolaterally from the midlateral base of the tongue. The tongue is cordiform, shallowly notched posteriorly, and barely free behind. The dentigerous processes of the vomers are transverse to slightly oblique between the posterior margins of the rounded choanae and bear numerous teeth. COLORATION The dorsal coloration in these Andean species usually is brown to gray, but some individuals of H. andina, callipleura, and marianitae are green; normally, the mark- ings on the dorsum are darker shades of the ground color. These markings vary from round spots in H. armata to nar- row transverse bars in H. balzani, irregular dashes in H. albonigra, andina, melanopleura, and palaestes, and vermicu- lations in some H. marianitae (Figs. 3, 4). The venter is creamy white to creamy gray or pale yellow in all species, but the vocal sac is brown or dark gray. In most species, the most distinctive markings are on the flanks and hid- den surfaces of the thighs. These markings consist of ver- tical black bars or irregular spots in H. andina and H. albonigra, large dark spots in H. armata, and pale spots on irregular marks on dark flanks in H. balzani and H. marianitae. The flanks and hidden surfaces of the thighs in H. melanopleura and H. palaestes are unpatterned, except for small pale dots on the ventrolateral edge of the flanks in H. melanopleura. A transverse white, supracloacal stripe is characteristic of most species (absent in H. albonigra). Faint white dorso- lateral, ulnar, and tarsal stripes are present in Hyla balzani and H. melanopleura; these stripes are bold in some indi- viduals of H. andina, some populations of which are char- acterized by blue in the axilla and groin in life. OsTEOLOGY The relatively few skeletal preparations of only a few taxa preclude a thorough analysis of skeletal features. In- stead, a brief description is given of principal osteological features of Hyla armata and of three members of the Hyla pulchella Group (H. andina, balzani, and palaestes). Ayla armata Skull.—The skull is about as long as wide, and the snout region is short and blunt. The nasals are widest anteriorly and separated medially by a distance about equal to about one half of their greatest width. The nasal is not in contact with the frontoparietal and it bears a short maxillary pro- cess approximating, but not articulating with, the preor- bital processes of the maxilla. The sphenethmoid is heavily ossified and broader than long in dorsal view; it underlies the nasals and anterior ends of the frontoparietals, and Fig. 4. Ventral views of right arm and hand of Hyla armata. Top, external view, KU 173219, male. Bottom, cleared-and-stained, KU 173220, male. Line = 10 mm. 10 ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS forms the anterior margin of the frontoparietal fontanelle. The septum nasi is ossified and extends anteriorly beyond the anterior margins of the nasals. The frontoparietals are slender and approximate one another only posteriorly. The frontoparietal fontanelle is short and oval. The prooti and exoccipital are incompletely fused to one another, and the prootic is elaborated anteriorly. The crista parotica is mod- erately broad and narrowly separated by cartilage from the squamosal. The squamosal is robust; the zygomatic ramus is short, and has a lanceolate cartilaginous process distally. The premaxilla has a high, vertical alary processes. The maxilla has a high facial process and a long, slender preorbital process, but no postorbital process. The vomers are broad and juxtaposed medially; each bears a massive transverse dentigerous processes. Each slender neopalatine bears a ventral ridge; the bone articulates with the maxilla and sphenethmoid. The parasphenoid is slender anteri- orly, where the tip is nearly at the level of the neopalatines; posteriorly, it is wider with irregular (feathered) lateral margins. The alary processes of the parasphenoid are nar- row and directed posterolaterally; the posteromedial pro- cess is short and does not reach the margin of the foramen magnum. The medial ramus of the pterygoid is short and robust; it articulates with the anterolateral surface of the otic capsule. The anterior ramus of the pterygoid is long and slender; it is in broad contact with the maxilla anteri- orly to the level of the neopalatine. Postcranial skeleton.—With a few exceptions, the axial and appendicular skeletons are typical of hylid frogs. A notable exception is the complete ossification of the su- prascapulae. Also, major differences exist in the forelimbs of adult males. In an adult male (KU 173200, SVL 60.5 mm), both the humerus and the radioulna are massive, and the former has greatly expanded flanges (Fig. 4). On the hu- merus, the crista lateralis is expanded laterally into a cres- cent-shaped process; the anterior part of the crescent bears numerous spines on the ventral surface. The cristae me- dialis and ventralis are greatly flared medially. In the hand, the prepollex is enlarged and supports an elliptical osseous plate medially. The first metacarpal is noticeably robust and distally supports a round osseous plate. Both plates support spines of soft tissue that are keratinized. In an adult female (KU 164085, SVL 71 mm) and a juvenile (KU 163334, SVL 32.4 mm), the humeri, radioulnae, and first metacar- pals are not noticeably enlarged; the cristae on the humeri are not expanded. The prepollices in both specimens are elongate, triangular structures; they are cartilaginous in the juvenile and ossified in the adult female. Hyla pulchella Group Skull.—The skull is about as long as wide, and the snout region is short and bluntly rounded. The nasals are qua- drangular and widely separated; the nasals are not in con- tact with the frontoparietals. Each nasal has a short maxil- lary processes approximating, but not articulating with, the preorbital process of the maxille. The sphenethmoid is - moderately ossified and broader than long in dorsal view. The septum nasi is not ossified anteriorly between the anteromedial margins of the nasals. The sphenethmoid does not underlie the nasals, but it does underlie the ante- rior ends of the frontoparietals. The frontoparietals are slen- der and approximate one another only posteriorly. The fr- ontoparietal fontanelle is large and elongate. The prootic is completely fused with the exoccipital and is not elabo- rated anteriorly; the crista parotica is robust and narrowly separated by cartilage from the squamosal. The entire squa- mosal is rather slender; the zygomatic rami is short and lacks a cartilaginous process distally. The premaxill has a high, vertical alary processes. The maxilla has a high fa- cial process and long, slender preorbital process, but no postorbital process. The vomers are moderately ossified and narrowly separated posteromedially; each bears a small, arched dentigerous process. The neopalatine is slen- der and smooth; it articulates with the maxilla and sphenethmoid. The parasphenoid is slender anteriorly, where the tip does not extend to the level of the neopalatines; at about midlength of the orbit the parasphenoid becomes abruptly wider and has smooth lateral margins. The alary processes of the parasphenoid are narrow and directed posterolaterally; the posterome- dial process is short in H. balzani and long and slender in H. andina and H. palaestes; it does not reach the margin of the foramen magnum. The medial ramus of the pterygoid is moderately long and slender; it articulates with the anteroventral surface of the otic capsule. The anterior ra- mus of the pterygoid is long and slender; it is in broad contact with the maxilla anteriorly nearly to the level of the neopalatine. Postcranial skeleton.—The axial and appendicular skel- etons are typical of hylid frogs. The suprascapulae are poorly ossified. The forelimbs of adult males have humeri with slightly expanded cristae medialis; the prepollex is enlarged and curved medially with a sharp distal point. ADVERTISEMENT CALLS The advertisement calls are known for five of the spe- cies, but few recordings are available. The call of Hyla andina was described by Barrio (1965) and Marquez et al. (1993), and that of H. armata was described by Cadle and Altig (1991) and Marquez et al. (1993), who also described the call of H. marianitae (as H. callipleura). Herein we present descriptions of the calls of H. andina, armata, balzani, marianitae, and palaestes in the individual species accounts. The calls of the Andean members of the Hyla pulchella Group are rather soft and low-pitched; they consist of a single note, as in H. armata, to a series of notes, as in H. andina (Table 3). 4 TADPOLES Tadpoles of four species (Hyla andina, armata, balzani, and palaestes) are known. The tadpoles of H. andina and H. ANDEAN GROUPS OF HYLA 11 Table 3. Characteristics of advertisement calls of Hyla armata and four Andean species in the Hyla pulchella Group. Ranges are in first row, and means are in parentheses in second row; m = number of calls. Notes/ Notes / Note Pulses/ Frequency Species n call minute duration (sec) second Dominant (kHz) Fundamental (kHz) Modulation H. armata' 3) 1 2-6 0.16-0.24 59-64 2.5—4.7 2.4-2.5 Up (4) (0.20) (61.5) (3.6) (2.45) H. andina? 2 2-3 21-24 0.01-0.06 110-120 1.7-2.3 1.1-1.2 Up (2.5) (22.5) (0.04) (115) (2.0) (1.15) H. balzani 2 1 2.5-4.0 0.17 80 1.15 1.15 No (3.25) H. marianitae® 5 1 15-23 0.25-0.59 44-45 0.84-0.93 0.84-0.93 No (18.7) (0.36) (44.6) (0.88) (0.88) H. palaestes il 1 9 0.2-0.8 42-44 4.43-4.45 4.43-4.45 No (0.36) (43) (4.44) (4.44) ' Based on data from Cadle and Altig (1991) and Marquez at al. (1993). ? Only recordings from Quebrada Grande, Prov. Jujuy, Argentina. 3 Based on data in Marquez et al. (1993). armata were described by Lavilla (1984) and Cadle and Altig (1991), respectively, whereas those of H. balzani, and H. palaestes are described herein for the first time. Tadpoles of these four species are Type-IV tadpoles (Orton, 1953) characterized by: (1) large size (> 40 mm at stages 37-38); (2) labial tooth rows of at least 2/4; (3) modest jaw sheaths with upper widely arched and lower broadly V-shaped; (4) marginal papillae; (5) submarginal papillae or flaps lat- erally; (6) sinistral spiracle; (7) dextral vent opening from a large saccular structure that underlies the limb buds; (8) uniformly pigmented, depressed bodies; (9) dorsal or dorsolateral eyes; (10) oval nares which are slightly me- dial to center of eyes, recessed, and weakly rimmed; (11) dorsal fin terminating at dorsal extent of tail-body junc- tion; and (12) tail fins with blotches (Table 4). Tadpoles of these four species exhibit considerable di- versity, particularly in the elements of the oral disc; this variation seems to be associated with habitat preference. The tadpole of H. armata is easily distinguished from the other three species by its large size, TL = 78.2 at Stage 37 (Cadle and Altig, 1991), which is much greater than that of the tadpole of H. andina, the next largest with TL = 57.4 at Stage 38. The most striking feature of H. armata is the labial tooth row arrangement (LTRF) of 13(13)/16(1) (Cadle and Altig, 1991) which is radically different from the other three species, none of which possesses more than three upper and four lower rows of teeth. The oral disc of H. armata also possesses complete marginal papillae, is slightly emarginate, and has jaw sheaths with shelves. In the other three species, the oral disc has a median gap in the marginal papillae of the upper labium, is not emargin- ate, and lacks shelves on the jaw sheaths. The caudal mus- culature of H. armata is uniformly darkly pigmented, whereas that of the other species at least has diffuse blotches. The tadpole of Hyla palaestes is similar to that of H. armata in having dorsally positioned eyes and an acutely rounded snout in profile. Both H. balzani and H. andina have dorsolaterally positioned eyes, and broadly and bluntly rounded snouts, respectively. As in both H. balzani and H. andina, the tadpole of H. palaestes possesses small flaps, which resemble broad papillae, laterally on the oral disc, and as in H. andina, these possess minute teeth. Although all these tadpoles except those of H. armata have a median gap in the marginal papillae of the upper labium, that of H. palaestes is narrow,whereas that of H. balzani and H. andina is wider. The LTRF of H. palaestes, 2(1)/4(1) also dif- fers from that of H. balzani and H. andina; the former vari- ably possesses a third interrupted upper row, and in the latter, the fourth row is variably present. The tadpoles of H. palaestes differ from those of the other species in having an acutely rounded snout in dorsal view and pigment on the dorsum of the tail musculature extending for only half the length of the tail; the others have broadly or bluntly rounded snouts and tail musculature pigmentation on the dorsum for the entire length of the tail. The body of H. palaestes possesses olive-green mottling and the blotches of the tail are very distinct, differing from the non-blotched bodies and diffuse tail blotches of the other species. The tadpoles of Hyla balzani and H. andina are the most similar of the four tadpoles. Both of these species have moderately wide gaps in the marginal papillae of the up- per labium, dorsolateral eyes, dorsal body pigment extend- ing posterior on the dorsum of the tail musculature for entire length of tail, white flecks on dorsum of body and tail fins, and diffuse blotches on the tail musculature and 12 ScreNTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Table 4. Comparisons of selected features of tadpoles of Hyla armata and the Hyla pulchella Group. Features for Hyla pulchella are from Lutz (1973) with some modification of terminology. __Character : H. andina A. armata H.balzani H. palaestes H. pulchella Total length 57.4mm 78.2 mm 41.7 mm 55.2 mm 71mm (Stage 38) (Stage 37) (Stage 38) (Stage 37) (maximum) Body shape Elongately ovoid Elongately ovoid Oval Oval Oval Eye position Dorsolateral Dorsolateral Dorsolateral Dorsal Dorsolateral Spiracle Terminally free Terminal third free Terminally free Completely attached — Fin terminus Pointed Rounded Pointed Pointed Pointed Ventral gut pigmentation Little Pigmented Little Little _ LTRF 2(2)/4(1) 15(15)/17(1) 3(1,3)/4(1) 2(2)/4(1) 2(2)/3 Shelf on upper jaw sheath None Prominent None None — Gap in anterior marginal papillae Middle third None Middle fifth Middle seventh Present fins—all features in which H. armata and H. palaestes dif- fer. Features unique to H. balzani include its small size (TL = 41.7 at Stage 38), short spiracle that is directed posteri- orly, broadly rounded snout in profile, white flecks on ven- ter, a dark brown interorbital bar in life, and the variable presence of a third row of labial teeth on the upper labium. The tadpole of H. andina is unique in having a bluntly rounded snout in both dorsal view and profile, finely ser- rate jaw sheaths, a variably present fourth row of teeth on the lower labium with small teeth, lateral flaps with teeth forming a broken vertical row, dark brown spots scattered throughout dorsum of body, and a deep bronze belly with silvery lichenous marks in life. ANDEAN GROUPS OF HYLA The foregoing analysis of characters leads us recognize Hyla armata as being highly distinctive from the other spe- cies in the southern Andes; the latter we associate with the Hyla pulchella group. Six other phenetic groups of Hyla are recognized in the Andes—viz., Hyla bogotensis Group (Duellman, 1972), Hyla chlorostea (Reynolds and Foster, 1993), Hyla columbiana Group (Duellman and Trueb, 1983), Hyla garagoensis Group (Kaplan and Ruiz, 1997)Hyla labialis Group (Duellman, 1989), and Hyla larinopygion Group (Duellman and Hillis, 1990)*. In the following definitions of these groups, characters are arranged in the same nu- *Hyla charazani was described from Charazani, 3200 m in the Andes of “Depto. Murieca” (= Prov. Saavedra, Depto. La Paz) in northwestern Bolivia by Vellard (1970), who reported that the specimen was in a stream. Vellard’s description of the unique holotype (an adult male 55 mm SVL) mentioned only a rudiment of a prepollex, broad dorsolateral stripes, granular skin, and bilateral vocal sac. The accompanying pho- tographs of the holotype show the pale dorsolateral stripe, no enlarge- ment of the forearm, and no indication of a protruding prepollical spine. We have not been able to examine the holotype, which was stated to have been deposited in the Museo Argentino de Ciencias Naturales (Vellard, 1970:16). However, the specimen was never deposited in that institution (Jorge Cranwell, in litt. to De la Riva). Subsequently, De la Riva was informed by Jehan Vellard (in litt.) that the specimen was in Vellard’s home in Buenos Aires. If the description and illustrations por- tray the specimen accurately, it is not a member of the Hyla pulchella Group. merical order in each account; accounts of species are pro- vided for taxa in the Hyla armata and Hyla pulchella groups. In the species accounts, we provide detailed information on each species in the Hyla armata and Hyla pulchella groups in the Andes. For ease of comparisons, the same seven char- acters are listed numerically in the diagnoses; these are followed by a paragraph discussing differences among similar species. Each description also is presented in a con- sistent format. Because of the great amount of variation in color pattern in some species, we provide many descrip- tions of living specimens. Criteria for synonymies and designation of lectotypes are given in the section on Re- marks. HYLA ARMATA GROUP Definition.—(1) Large, stream-breeding frogs with males attaining SVLs of 68 mm and females 73 mm; (2) round mental gland absent; (3) preorbital ridge and lat- eral folds absent; (4) dorsum brown with darker brown spots or cream reticulations; (5) flanks pale with large dark spots; hidden surfaces of thighs uniform brown; (6) ven- ter uniformly pale; (7) digits bearing large discs; (8) clus- ter of spines on prepollex and on proximal ventral surface of humerus (9) forearms of adult males greatly hypertro- phied; (10) axillary membrane absent; (11) toes three- ANDEAN Groups OF HYLA 13 fourths webbed; (12) nasals moderately broad, separated medially; (13) frontoparietal fontanelle small; (14) sphenethmoid broad; septum nasi ossified anteriorly; (15) quadratojugal articulating with maxillary; (16) dentiger- ous processes of vomers long, transverse, posterior to choa- nae, barely separated medially; (17) tadpoles having long, muscular tails, low fins, and rounded terminus; (18) distal one third of spiracle free from body wall; (19) larval mouth large, ventral, with complete rows of marginal papillae, broad plate below serrations on upper jaw sheath, and a LTRE of 15/17; (20) advertisement call a single, high- pitched note. Content.—One species: Hyla armata Boulenger, 1902. Distribution.—Cloud forests at elevations of 1700-2400 m on the eastern slopes of the Andes from central Peru to central Bolivia. Remarks.—Two series of tadpoles (KU 181882-83 from 25.5 and 28.3 km, respectively, SSW of Leimebamba, Depto. Amazonas, Peru) are like those of Hyla armata in having a large mouth directed ventrally, two complete rows of mar- ginal papillae, a broad plate below the serrations on the upper jaw sheath (shelf of Cadle and Altig, 1991) and a round terminus to the tail. These tadpoles differ from those of H. armata in having the spiracular tube attached for its entire length to the body wall and fewer rows of labial teeth (LTRF 3/7(1)). These tadpoles were taken from pools in cascading streams at elevations of 3300 and 3450 m in the northern part of the Cordillera Central; no frogs have been associated with these tadpoles, which possibly rep- resent the existence of a species related to H. armata. The presence of a cluster of spines on the prepollex and on the proximal ventral surface of the humerus in adult males is unique in the Hylidae. Similar, but less conspicuous, clus- ters of spines on the prepollex occur in breeding males of some species of Ptychohyla, plus Hyla echinata, lancastert, and pachyderma in Middle America (Duellman, 1970). Ayla armata Boulenger Hyla armata Boulenger, 1902:394.—Syntypes: BM 1947.2.13.59—-60 from “La Paz,” Bolivia. Diagnosis.—A species in the Hyla armata Group char- acterized by: (1) snout bluntly rounded in dorsal view and in profile; (2) skin on dorsum smooth to weakly granular; (3) width of disc on Finger III about twice diameter of tym- panum; (4) tarsal fold low; (5) dorsal color pattern consist- ing of moderately large round, dark spots, cream reticula- tions of cream spots; (6) flanks with large dark spots; (7) posterior surfaces of thighs dark brown. Hyla armata differs from species in the Hyla pulchella group by its larger size, proportionately smaller tympa- num, and in coloration—viz., the absence of dark bars or white or creamy-yellow spots on the flanks and posterior surfaces of the thighs. Moreover, the species is unique in the genus by having a large cluster of black spines on the inner surface of each thumb and on the proximal ventral surface of the humerus in adult males. Fig.5. Variation in dorsal color pattern in female Hyla armata from Peru. KU 166341, 68.8 mm SVL; KU 166342, 74.5 mm SVL; KU 166343, 57.3 mm SVL; KU 166344, 61.2 mm SVL (all from Rio Cosnipata, Depto. Cuzco); KU 166345, 68.0 mm SVL, from Tutumbaro, Depto. Ayacucho. Description.—n = 36 dc, 20 99. Body robust; SVL in males to 68.5 mm, in females to 74.5 mm; head width about equal to head length, not as wide as body; head width 30.9- 37.4% (X= 33.9) of SVL; head length 28.4-36.5% (x = 32.3) of SVL; snout short, bluntly rounded (nearly truncate) in dorsal view, bluntly rounded in profile; interorbital dis- tance 96-146% (X= 118) of width of eyelid; diameter of eye 104-177% (x = 144) its distance from nostril; tympanum separated from eye by distance about twice diameter of tympanum; diameter of tympanum 23.2-50.0% (xX = 34.8) of diameter of eye in males, 23.9-40.3% (X = 33.2) in. Breadth of forearm 66.2—74.2% (X= 69.2, n = 10) of forearm length in males, 36.5-44.2% (x =40.0, n = 10) in females; row of low, indistinct ulnar tubercles; webbing formula for fingers: II (2—2*)—(3-3°) II (3—3)—(2-2%) IV; width of disc on Finger III at least twice diameter of tympanum; cluster of black nuptial spines on inner surface of thumb and on proximal ventral surface of forearm in breeding males (Fig. 4). Hind limb moderately robust; tibia length 47.2-58.6% (x = 53.0) of SVL; foot length 43.0-60.8% (x = 47.9) of SVL; inner tarsal fold low, barely curved; inner metatarsal tubercle ovoid, barely visible from above; outer metatarsal tubercle low, round, about one fourth size of inner metatarsal tubercle; webbing formula for toes: I (I— 1%)—(1%-2>) I (1-1*)—(1%-2) TI (1*-1%)— 2-2") IV2— 2*)—(1-1°*) V. Skin on dorsum and flanks smooth to finely granular. Vomerine odontophores long, transverse, nar- rowly separated medially, posteromedial to rounded choa- nae, bearing total of 10-27 (x = 17.5) teeth. Coloration in preservative: Dorsum usually brown with darker brown or cream irregular markings (few large marks, many large rounded spots, or mottling) outlined, or not, by tan or black (Fig. 5); flanks uniform brown or tan or gray with dark brown spots or lateral extensions of 14 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS — 0 5 10 15 20 Kilohertz dorsal markings; variable number of broad to narrow trans- verse marks on limbs—1-3 on forearm, 3-6 on thigh, 2-5 on shank, 3-6 on tarsus; anterior and posterior surfaces of thighs uniform brown or more commonly marked with extensions of transverse marks on dorsal surfaces of thighs; ventral surfaces grayish brown to dark brown with or with- out cream streaks or reticulations on throat. Coloration in life: In living individuals, the coloration is even more variable than in preservative, as evidenced from Figure 3 and the following exerpts of field notes: KU 163341-44 from Rio Cosnipata, 4 km SW Santa Isabel, Depto. Cuzco, Peru—Dorsum bronze tan mottled with dark brown and creamy yellow; flanks creamy yel- low with dark brown and black spots and ocelli; axilla, groin, and posterior surfaces of thighs lavender; webbing pale lavender-gray; venter grayish brown; iris pale gray (W. E. Duellman field notes, 11 February 1975). KU 163345 from Tutumbaro, Rio Piene, Depto. Ayacucho, Peru—Dorsum tan with dark brown spots; ven- ter gray; iris olive-tan (W. E. Duellman field notes, 20 Feb- ruary 1975). KU 173219-22 from Buenos Aires, Depto. Cuzco, Peru.— At night, uniform tan. By day: 173219, dorsum dark brown with olive-green spots; 173220-22 dull reddish brown with slightly darker bars on hind limbs; in all of above, venter dull grayish brown; spines on chest and thumbs black; webbing dull brown; iris dull olive-cream with fine black reticulations (W. E. Duellman field notes, 19 January 1977). KU 183299-313 from 68.8 km SW Villa Tunari, 1860 m, Depto. Cochabamba, Bolivia—Dorsum dark brown with olive markings; ventral surfaces dark brown with yellow- olive markings; vocal sac gray-brown, iris chocolate-brown (D. C. Cannatella field notes, 21 January 1979). MNK AM963 from La Siberia, 2300 m, Depto. Cochabamba, Bolivia.— Dorsum beige with dark, irregu- lar markings; venter purplish-brown with yellow dots. Iris pale green (I. De la Riva field notes, 16 February 1994). One of the most striking variations is the color of the iris, which ranges from olive-tan in Ayacucho, Peru, to ol- ive-cream or pale gray in Cuzco, Peru; more striking vari- ants are known from Bolivia, where individuals having the iris chocolate-brown were found NW of Villa Tunari, reddish brown at Serrania Bellavista, and red or green at La Siberia (field notes and color transparencies of I. de la Riva). Life history.—Calls of this species were recorded at Amaybamba, Depto. Cuzco, Peru (Cadle and Altig, 1991) and at Serrania de Bellavista, Depto. La Paz, Bolivia, (Marquez et al., 1993). The call was described as a high- pitched “whirrr” by Cadle and Altig (1991). Analyses of these two recordings reveal the call to consist of a single monophasic, well-modulated note repeated at a rate of 2— 6 notes per min and having a duration of 0.16—0.24 sec and a pulse rate of 59-64 pulses per sec. Three harmonics— 2.4, 3.6, and 4.7 kHz (dominant)—are evident in the Peru- vian call, whereas only two harmonics—2.5 (dominant) Decibels Kilohertz Pascals Seconds Fig. 6. Advertisement call of Hyla armata recorded at Serrania de Bellavista, Depto. La. Paz, Bolivia. Spectrum, audiospectrogram, and waveform; arrow in audiospectrogram indicates point of spectrum. and 5 kHz—are evident in the Bolivian recording (Fig. 6). Both calls show a slight upward frequency modulation. Tadpoles of this species from Amaybamba, Depto. Cuzco, Peru, were described by Cadle and Altig (1991). They were characterized by being large (total length 78.2 in Stage 37) and having complete marginal papillae, slightly emarginate oral disc, jaws sheaths with shelves, and a labial tooth row formula (LTRF) of 13(13)/16(1). One tadpole (CET A483) from 40 km N Caranavi, Depto. La Paz, Bolivia, and two tadpoles (CET A484-85) from Carahuasi, Depto. Cochabamba, Bolivia, were taken in streams inhabited by adults of Hyla armata and are assigned to that species. In nearly all features they are like the tad- poles described by Cadle and Altig (1991), except that two of them have more rows of labial teeth (Fig. 7). The stages, sizes, and LTRFs of these tadpoles are: CET A483—Stage 25, body length 35.0 mm, total length 81.0 mm, LTRF 15(15)/17(1); CET A484—Stage 37, body length 29.0 mm, total length 73.5 mm, LTRF 14(14)/16(1); CET A485—Stage 25, body length 32.0 mm, total length 75.5 mm, LTRF 13(13)/16(1). The LTRF of 15/17 is close to the largest re- ported for any hylid frog; the highest LTRF is 14/21 in H. inparquesi (Ayarzaguena and Senaris, 1993). Twenty-one juveniles (some with remnants of clusters of melanophores near the tip of the coccyx) were found on 10 February 1974 at the Rio Cosnipata, 4 km W of Santa Isabel, Depto. Cuzco, Peru. Data were taken on 18 of these, which have SVLs of 28.0-33.3 mm (x = 31.0). Coloration in preservative is highly variable (Fig. 8). Eleven individuals have distinct bars on the limbs; four have pale dorsolat- ANDEAN Groups OF HYLA 15 Fig. 7. Tadpole of Hyla armata, CET A483, Stage 25, total length 80.8 mm. eral stripes. The dorsum of the body has distinct dark spots in five individuals, whereas in the other 13, the dorsum is brown with darker brown flecks. In all individuals, the venter is cream with a dusky suffusion on the throat. In life, the dorsum is tan with or without darker brown blotches and creamy-yellow dorsolateral stripes and bars on limbs; the venter is creamy white, and the iris is pale dull bronze with fine black reticulations (W. E. Duellman field notes, 10 February 1974). Distribution and ecology.—Hyla armata is known from cloud forests at elevations of 1700-2400 m on the eastern slopes of the Andes from Depto. Ayacucho, Peru, south- eastward to Depto. Santa Cruz, Bolivia (Fig. 9). The spe- cies always is associated with cascading streams, where adults perch on boulders and cliffs in the spray zone of waterfalls, and adults and juveniles are on logs, branches, and leaves of plants usually within 1 m of the water, but some individuals have been found as high as 2 m above streams. At Amaybamba, Peru, and at Carahuasi, Bolivia, tad- poles were in pools in streams, but at 40 km N Caranavi, Bolivia, a tadpole was in a small, high-gradient stream. Fig.8. Variation in dorsal color pattern in juvenile Hyla armata from the Rio Cosnipata, Depto. Cuzco, Peru. The tadpoles are capable of sustained swimming in strong currents (Cadle and Altig, 1991). Remarks.—The type locality of “La Paz” (Boulenger, 1902:394) most certainly is incorrect and probably was the point of shipping from Bolivia. The city of La Paz lies at elevations of about 3500-4000 m, at least 1500 m higher than the known altitudinal distribution of Hyla armata. Geographic variation seems to be apparent in the color of the iris in life. No red or green is evident in Peruvian specimens, but these colors and chocolate-brown are present in the iris in some populations in Bolivia. Also, the dorsal skin tends to be more granular in Bolivian speci- mens than in those from Peru. These slight differences sug- gest that possibly Hyla armata, as treated here, may repre- sent more than one species. This idea is not novel, in that J. E. Cadle (pers. comm.) “thought” that specimens he col- lected at the Rio Quiquira, Depto. Puno, Peru, had a dif- ferent call from specimens that he recorded in Depto. Cuzco, Peru. Although both Duellman and De la Riva in- dependently examined these specimens (USNM 299310- 38) and confirmed their identity as H. armata, we were not given the opportunity to take data on them and to include them in our analyses. HYLA PULCHELLA GROUP Definition.—(1) Moderately large, stream-breeding frogs with males attaining SVLs of 58 mm and females 63 mm; (2) round mental gland absent; (3) preorbital ridge and lateral folds absent; (4) dorsum brown, tan, green, or gray, usually with darker markings consisting of transverse bars, spots, or reticulations; (5) groin and hidden surfaces of thighs pale with black spots, bars, or reticulations, or dark with pale spots; (6) venter uniformly pale; (7) digits bearing large discs; (8) nuptial excrescences absent in males; enlarged, projecting prepollex present; (9) forearms of adult males hypertrophied; (10) axillary membrane ab- sent; (11) toes three-fifths to three-fourths webbed; (12) nasals slender, widely separated medially; (13) frontopa- rietal fontanelle large; (14) sphenethmoid broad; septum nasi not ossified anteriorly; (15) quadratojugal articulat- ing with maxillary; (16) dentigerous processes of vomers medium in length to moderately long, transverse, poste- rior to choanae, narrowly separated medially; (17) tadpoles having moderately long tails, moderately low fins, and pointed terminus; (18) spiracular tube attached to body wall for all, or nearly all, of its length; (19) larval mouth moderately small, directed anteroventrally with a median gap in the labial papillae on the anterior labium, no plate below serrations on upper jaw sheath, and a LTRF of 2-3/ SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS H. albonigra H. andina H. armata H. balzani H. marianitae H. melanopleura H. palaestes 300 Kilometers 1000 meters — 3000 meters Fig.9. Distribution of Hyla armata and species of the Hyla pulchella Group in the Andes. ANDEAN Groups OF HYLA LF 3-5; (20) advertisement calls consisting of series of soft, bell-like notes. Content.—Eleven species: Hyla albonigra Nieden, 1923 (H. zebra Duméril and Bibron, 1841, and H. ocapia Andersson, 1938, are synonyms); H. andina Muller, 1924; H. balzani Boulenger, 1898 (H. callipleura Boulenger, 1902 is a synonym); H. cymbalum Bokermann, 1963; H. marginata Boulenger, 1887; H. marianitae Carrizo, 1992; H. melanopleura Boulenger, 1912; H. palaestes new species; H. prasina Burmeister, 1856; H. pulchella Duméril and Bibron, 1841 (H. leucomelas Duméril and Bibron, 1841; H. vauterii Bibron, 1843°; H. agrestis Bibron, 1843; H. leucotaenia Burmeister 1861; H. bracteator Hensel, 1867; and Hylodes orientali: Philippi, 1902, are synonyms); H. semiguttata Lutz, 1925. Distribution.—In the Andes, elevations of 500-3416 m in cloud forest, dry forest, subparamo, and paramo from the Cordillera Oriental in central Peru to precordilleras in central Argentina; also in the Sierra de Cordoba in Argen- tina and in the Platine region of northeastern Argentina, Uruguay, and southeastern Brazil, where it ascends to el- evations of 1100 in the Brazilian Highlands (Lutz, 1973). Remarks.—The extra-Andean species include Hyla cymbalum, marginata, prasina, pulchella, and semiguttata, which is the only species having a LTRF of 3/5 (Faivovich, 1996). Although the Hyla pulchella Group was recognized and defined by Barrio (1965), Lutz (1973), and Cei (1980), the only Andean species recognized here that was referred to the group was H. andina (as H. pulchella andina). As noted by Lutz (1973), the taxonomic status of some members of this group in southeastern Brazil is questionable. (For fur- ther comments on the taxonomy, see Remarks in account of H. andina.) The forelimbs in males are nearly as hypertrophied as those in Hyla armata, whereas the forelimbs in females are not enlarged (Fig. 10). Hyla albonigra Nieden Hyla zebra Duméril and Bibron, 1841: 575.—Syntypes: MNHN 4817 (2 speci- mens) from “Buenos Ayres” in error; Tacopaya, Departamento Chuquisaca, Bolivia (d’Orbigny, 1847). (Preoccupied by Rana zebra Shaw, 1802 = Rana (Hyla) boans Linnaeus, 1758). Hyla albonigra Nieden, 1923:278.—New name for Hyla zebra Dumeéril and Bibron, 1841. Hyla ocapia Andersson, 1938:19.—Syntypes: NHRM 1873 (7 specimens) from Puka Khara, Bolivia. New synonymy. Diagnosis.—A large species in the Hyla pulchella Group characterized by: (1) snout rounded in dorsal view and bluntly rounded in profile; (2) skin on dorsum coarsely granular; (3) width of disc on Finger III equal to diameter of tympanum; (4) tarsal fold elevated; (5) dorsal color pat- tern consisting of small irregular spots; (6) flanks with dark vertical bars; (7) posterior surfaces of thighs with dark vertical bars. >Hyla vauterii was recognized as a species distinct from H. pulchella by Klappenbach and Langone (1992). Fig. 10. Ventral views of the right arm and hand of Hyla andina. Top left, external view, KU 160418 GC; top right, external view, KU 160412, 9; bottom, cleared-and-stained, KU 182961, C. Line = 10 mm. The only other species with dark vertical bars on the hidden surfaces of the thighs is Hyla andina, which differs from H. albonigra by having small dark spots on the flanks and ventral surfaces of the shanks (shanks unmarked in H. albonigra), a transverse white supracloacal stripe (ab- sent in H. albonigra), width of disc on Finger III less than diameter of tympanum, and a low tarsal fold. Also the skin on the dorsum is not so coarsely granular in H. andina as in H. albonigra. Description.—1 = 9 dC, 11 99. Body moderately robust; SVL in males to 56.0 mm, in females to 64.7 mm; head width slightly greater than head length, not as wide as body; head width 28.5-38.7% (X= 34.3) of SVL; head length 24.6-34.8% (x = 30.3) of SVL; snout rounded in dorsal view, bluntly rounded in profile; internarial region not depressed; in- terorbital distance 91-162% (x = 120) of width of eyelid; 18 ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Fig. 11. Hyla albonigra, MNK A846, adult male, 48.7 mm SVL. diameter of eye 105-142% (x = 123) its distance from nos- tril; tympanum separated from eye by distance slightly less than diameter of tympanum; diameter of tympanum 43.1— 63.3% (X = 53.8) of diameter of eye. Breadth of forelimb 60.2-63.8% (x = 62.1, n = 4) of forearm length in males, 32.7-38.5% (X = 34.7, n = 3) in females; ulnar fold proxi- mally, formed by coalesced tubercles; webbing formula for fingers: II (2—2)—3 III (2-3-)—(2 —2) IV; width of disc on Finger III about equal to diameter of tympanum, nuptial spines absent; nuptial excrescences absent in breeding males. Hind limb slender; tibia length 49.4-57.7% (x= 53.9) of SVL; foot length 41.3-49.3% (x = 46.1) of SVL; inner tar- sal fold elevated, slightly curved; webbing formula for toes: I 1%—(2-2) If (1-17) —(2-2") MI (1—-1%4)—(2*-2%4) IV (2- 2*)—1 V. Skin on dorsum, at least on head and anterior part of body, coarsely granular; flanks smooth. Dentiger- ous processes of vomers short, transverse, narrowly sepa- rated medially, posteromedial to rounded choanae, each bearing total 6-14 (x = 9.1) teeth. Coloration in preservative: Dorsum gray to brown with few to many dark gray to dark brown marks in the form of small spots or, more commonly, streaks or short trans- verse marks; limbs with narrow transverse bars, usually faint (Fig. 11); flanks cream with 2-5 black marks usually in the form of vertical bars; anterior and posterior surfaces of thighs cream to grayish tan with narrow, vertical black bars—2-5 on anterior surfaces, 3-8 (usually 6 or 7) on pos- terior surfaces; 2-5 bars on inner surfaces of shanks; ven- ter creamy yellow, except throat dusky in females and dark grayish brown in males. One female (EBD 26926) with only two small black spots posteriorly on flanks, one spot proxi- mally on anterior surface of thigh, and no marks on inner shank. Coloration in life: José Cabot noted specimens from the Rio Julpe, Depto. Cochabamba, Bolivia, as being green in life. Life history—Nothing is known. Distribution and ecology.—The few localities from which this species is known are at elevations of 2000-3416 m on the eastern front of the Andes in southern Bolivia (Fig. 9). At the Rio Julpe, Bolivia, individuals were perched in small trees and bushes at the edge of a stream at night. Remarks.—Duméril and Bibron (1841:575) stated that the two syntypes of Hyla zebra were sent to Paris from “Buenos Ayres” by d’Orbigny. The same locality informa- tion was given by Boulenger (1882; 1887) in his redescrip- tion of the species. Berg (1896) pointed out that Duméril and Bibron gave an incorrect locality; d’Orbigny (1847) re- ported that he collected these specimens at Tacopaya, Bo- livia. Tacopaya (19°10' S, 64°40' W) is in a region of high, arid mountains between Sucre and Rio Grande in Departamento Chuquisaca, Bolivia. When the syntypes of Hyla zebra were examined by Duellman on 3 July 1969, the specimens were soft and nearly completely bleached; however, the pattern of verti- cal dark bars on the posterior surfaces of the thighs was evident. Duméril and Bibron (1841:575) observed the granular nature of the skin (“Téte couverte de petites glandules.”) and noted the coloration on the flanks: “... grandes taches noires sur les flanc, ...” Examination of the seven syntypes of Hyla ocapia by Duellman on 17 July 1969 revealed that they were in good condition with distinct color patterns. The dorsum was pale gray with darker gray, irregular, small dark gray marks. The anterior and poste- rior surfaces of the thighs were creamy yellow with dark brown vertical marks; the throat and chest were creamy white; the belly and ventral surfaces of the limbs were creamy yellow. The skin on the dorsum was distinctly granular. The specimens closely match Andersson’s (1938) description and illustration. Examination of the type speci- mens of both species convinces us that Hyla ocapia Andersson, 1938, is a junior synonym of Hyla zebra Dumeéril and Bibron, 1841. Andersson (1938) reported that one of the syntypes of Hyla ocapia (a male) had about 40 eggs covered with mu- cus and attached to the posterior part of the belly and com- mented about the differences with other hylid frogs, such as Fritziana (= Flectonotus) carrying eggs. We conclude that Andersson was mistaken in thinking that he witnessed an example of parental care, because he also mentioned that the frogs had been kept in a jar for several days and that the females spawned therein. Thus, it is most likely that part of a clutch adhered to the male, which was preserved “carrying” eggs in this fashion. Moreover, examination of the syntypes reveals that one of the females also has rem- nants of mucus adherent on the belly. Reproductive modes of frogs having parental investment in the form of egg- carrying, or egg-brooding is correlated with a low number of eggs (Duellman and Trueb, 1986). One gravid female (EBD 28926) contained 750 pigmented eggs, 1.5-1.6 mm ANDEAN Groups OF HYLA 19 in diameter; the number, sizes, and pigmentation of the eggs are consistent with aquatic deposition of eggs. Hyla andina Muller (New combination) Hyla raddiana andina Miiller, 1924:77.—Syntypes: ZSM 18/1922 (3 specimens) from Nacimiento, Provincia Catamarca, Argentina. Hyla pulchella andina—Barnio, 1965:124. Diagnosis.—A moderate-sized species in the Hyla pulchella Group characterized by: (1) snout rounded in dorsal view and bluntly rounded in profile; (2) skin on dorsum smooth to granular; (3) width of disc on Finger III less than diameter of tympanum; (4) tarsal fold low; (5) dorsal color pattern consisting of small irregular spots; (6) flanks with small dark spots; (7) posterior surfaces of thighs with dark vertical bars. The only other species with dark vertical bars on the hidden surfaces of the thighs is Hyla albonigra, which dif- fers from H. andina by having vertical dark bars on the flanks and lacking dark spots on the ventral surfaces of the shanks (shanks with dark spots in H. andina), no trans- verse white supracloacal stripe (present in H. andina), width of disc on Finger III equal to diameter of tympanum, and an elevated tarsal fold. Also the skin on the dorsum is more coarsely granular in H. albonigra than in H. andina. Description.—1 = 98 do, 59 99. Body moderately ro- bust; SVL in males to 57.6 mm, in females to 62.7 mm; head width slightly greater than head length, about as wide as body; head width 28.0-36.0% (x= 32.0) of SVL; head length 29.0-37.0% (X= 33.0) of SVL; snout rounded in dorsal view, bluntly rounded in profile; internarial region not de- pressed; interorbital distance 84-192% (x = 121) of width of eyelid; diameter of eye 86-167% (xX = 122) its distance from nostril; tympanum separated from eye by distance about equal to diameter of tympanum; diameter of tym- panum 42.0-76.0% (x = 62.0) of diameter of eye. Breadth of forearm 49.4—60.2% (X = 55.3, n = 10) of forearm length in males, 34.4-39.0% (X= 36.7, = 10) in females; no row of tubercles on forearm; webbing formula for fingers: I] 2— (3-3*) III 3 —(2-2*) IV; width of disc on Finger III slightly smaller than diameter of tympanum; nuptial spines ab- sent; nuptial excrescences absent in breeding males. Hind limb slender; tibia length 47.0-59.0% (x= 52.0) of SVL; foot length 38.0-62.0% (x = 46.0) of SVL; inner tarsal fold low, slightly curved; webbing formula for toes: I (I-1*)—(2- 2 MN @=12)—(2=2>) MIN (—2) —(2'—3) IV (22%) — (1-172) V. Skin on dorsum weakly granular; flanks smooth. Den- tigerous processes of vomers short, transverse, narrowly separated medially, between posterior margins of rounded choanae, each bearing total 3-14 (x = 8.2) teeth. Coloration in preservative: Dorsum tan, grayish brown, or brown with darker brown markings—small, irregular spots on head and body with or without dashed white or cream dorsolateral line; some individuals also with white flecks or irregular streaks on dorsum or only with such markings; narrow transverse dark marks usually evident on dorsal surfaces of limbs—3 or 4 on forearm, 4-6 on thigh, 6 or 7 on shank, and row of dark spots on tarsus and Toes IV and V; flanks tan or brown anteriorly and cream poste- riorly, with many round to irregular dark brown to black spots, most numerous in groin; anterior and posterior sur- faces of thighs cream to tan with vertical black bars or ir- regular black marks—3-5 on anterior surface and 4-10 on posterior surface; inner and ventral surfaces of shank cream with irregular black spots; venter dull cream; vocal sac grayish brown. Coloration in life: The color pattern is highly variable and seems to show some geographic variation. (See Re- marks.) This variation is evident from the photographs (Fig. 3) and the following descriptions of living individu- als. CET A122-126 from Rio Corani, Depto. Cochabamba, Bolivia—Dorsum brown with irregular dark blotches; a barred pattern consisting of narrow yellow and black trans- verse stripes on the groin, and on the anterior and poste- rior surfaces of thighs, shanks, and tarsi; dorsolateral stripes not distinct; no blue in groin and on hind limbs; by day, dorsum mostly greenish yellow with dark blotches (I. De la Riva field notes, 22 May 1988). CET A134-135 from La Siberia, Depto Cochabamba, Bo- livia.—Similar to specimens from Rio Corani, but with blue in groin and on anterior and posterior surfaces of hind limbs (I. De la Riva field notes, 7 January 1989). CET A144-145 from La Siberia, Depto. Cochabamba, Bo- livia.—- Dorsum uniformly green; flanks and groin blue with black spots; thighs and shanks with black and blue transverse bars (I. De la Riva field notes, 4 January 1990 and 9 March 1990). CET A116 from 5 km E Samaipata, Depto. Santa Cruz, Bolivia.—Dorsum uniformly brown; no blue in groin or on hind limbs (I. De la Riva field notes, 18 May 1988). Varia- tion in specimens from this locality was discussed by Kohler et al. (1995). CET A117-121 from Comarapa, Depto. Santa Cruz, Bo- livia——Dorsum brown with irregular dark blotches and irregular dorsolateral creamy-white stripes; no blue in groin or on hind limbs (I. De la Riva field notes, 19 May 1988). CET A146 from Vallegrande, Departamento Santa Cruz, Bolivia—Dorsum green with white reticulations; no dor- solateral stripes; instead of typical barred pattern in groin and on anterior and posterior surfaces of hind limbs, these surfaces pale blue with small black and white spots (I. De la Riva field notes, 6 January 1990). : KU 160143 from 3 km S Epizana, Depto. Cochabamba, Bolivia —Dorsum green with yellow flecks and black and gold bars on flanks; posterior surfaces of thighs grayish lavender with pale creamy-yellow and black vertical lines; belly pinkish cream; vocal sac bronze-tan; ventral surfaces of limbs pinkish gray; webbing and tympanum bronze- tan; iris bronze (W. E. Duellman field notes, 11 November 1974). KU 183385-92 from 2.9 km S Epizana, 3070 m, Depto. Cochabamba, Bolivia.—Dorsum dark green to pale green; 20 pale yellow lines on flanks, forelimbs, thighs, and shanks; belly off-white; lateral portions of chin green, central re- gion yellowish brown; dark blue markings on concealed surfaces of groin, thighs, shanks, and axilla, also in post- tympanic region; pale yellow bands fading to off white on concealed surfaces of thighs; upper lip from below eye to angle of jaw pale yellow; tympanum and iris gold; ventral surfaces of thighs, shanks, and forearms gray blue; web- bing colorless; supracloacal tubercles pale yellow; all fin- gers and Toes IV and V green; Toes [III grayish green; tongue very pale green (D. C. Cannatella field notes, 16 December 1977). KU 160195 from 6 km W Betanzos, Depto. Potosi, Bo- livia——Dorsum green with golden-tan and reddish-brown transverse marks; groin and posterior surfaces of thighs pale blue with black and orange-cream bars; venter creamy white; iris and tympanum bronze (W. E. Duellman field notes, 13 November 1974). KU 160236-39 from 12.5 km SSE Tupiza, Depto. Potosi, Bolivia.—Dorsum green with or without black dashes; la- bial, canthal, and supratympanic stripes bronze; ground and anterior and posterior surfaces of thighs pale blue with black bars; venter cream; webbing grayish brown; tympa- num bronze-brown; iris dull reddish bronze (W. E. Duellman field notes, 15 November 1974). KU 160337-41 from Rio Grande, 10 km N Tilcara, Prov. Jujuy, Argentina —Dorsum green with bronze dorsolateral stripe and labial stripe; flanks creamy blue with black spots; posterior surfaces of thighs pale blue with black dashes; vocal sac yellowish tan; rest of venter white; iris dark red- dish bronze (W. E. Duellman field notes, 18 November 1974). KU 160342-60 from Arroyo Los Naranjos, 8.3 km SSW Perico del Carmen, Prov. Jujuy, Argentina.—At night, dor- sum tan with or without brown spots; flanks and hidden surfaces of hind limbs creamy white to pale bluish white with black spots and flecks; vocal sac dull yellow; rest of venter creamy white; webbing brown; some individuals have creamy-bronze canthal, dorsolateral, and tarsal stripes or rows of dashes; iris reddish brown. By day, some individuals changed to pale creamy tan and others to pale greenish bronze (W. E. Duellman field notes, 19 Novem- ber 1974). KU 182964-67 from Finca Jakulika, Prov. Salta, Argen- tina——Dorsum green or pale green; stripes dark blue on off-white background; vocal sac yellowish green. (D. C. Cannatella field notes, 15 December 1978). KU 160365 from 21 km SSE Tafi del Valle, Prov. Tucuman, Argentina —Dorsum pale green; stripes on lips and limbs cream; groin and posterior surfaces of thighs pale blue with black marks; iris reddish copper-bronze (W. E. Duellman field notes, 21 November 1974). KU 160410-21 from Arroyo El Durazno, 19 km N La Merced, Prov. Catamarca, Argentina——Dorsum green in most individuals; in others dorsum tan with or without brown spots; vocal sac yellowish tan; labial stripe pale ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS -60 Decibels — 2 4 6 8 10 Kilohertz -80 Kilohertz 0.0 Pascals 2.0 24 2.2 2.3 Seconds Fig. 12. Advertisement call of Hyla andina, KU Tape 1208, recorded at 19°C at Quebrada Grande, 10 km N Tilcara, Prov. Jujuy, Argentina. Spectrum, audiospectrogram, and waveform; arrow in audiospectrogram indicates point of spectrum. greenish yellow; rows of flecks dorsolaterally on body and on shank and tarsus cream to bronze; iris copper-bronze (W. E. Duellman field notes, 3 December 1974). Life history.—The advertisement call of individuals from Horco Molle, Prov. Tucuman, Argentina, was re- corded at 13°C by Barrio (1965), who reported as many as seven notes per call and a dominant frequency at about 2400 Hz. Ata site 2.9 km S of Epizana, Depto. Cochabamba, Bolivia, the call was described as a metallic “clink-clink” (D. C. Cannatella field notes, 16 January 1979). Reynolds and Foster (1992) described the call of individuals from Departamento Cochabamba, Bolivia, as containing one or two notes that are frequency modulated and have domi- nant frequencies of 1420-2000 Hz. Marquez et al. (1993) reported two tonal notes produced by individuals at Tablasmontes, Depto. Cochabamba, Bolivia. Two individu- als were recorded at 18° and 19°C at Quebrada Grande, 10 km N of Tilcara, Prov. Jujuy, Argentina. The call is a series of tinkle-like notes. The call of one individual consisted of three notes per call, and the calls were repeated at a rate of 21 calls per min; the second individual produced only two notes per call at a rate of 24 calls per min. Analysis of two recordings (KU Tapes 1208 and 1210) reveals a pulse rate of 110-120 pulses per sec. The duration and modulation of notes is variable within a given call. For example in KU Tape 1208, the first note has a duration of 0.06 sec and one harmonic at 1940 Hz; the second note has a duration of 0.02 sec, and the third note has a duration of 0.05 sec, a harmonic at 1400 Hz, and a dominant harmonic at 1710 Hz (Fig. 12). In KU Tape 1210, duration of the first note in ANDEAN Groups OF HYLA 21 each pair of notes is much shorter (0.01 sec) than the sec- ond note (0.06 sec); in these the dominant frequencies are at 2110 and 2270 Hz (Fig. 12). The tadpole of Hyla andina was described and illustrated (as H. pulchella andina) by Cei (1980) and Lavilla (1984); the latter provided data on variation in tadpoles in some Ar- gentine populations. Lavilla and Fabrezi (1987) described the chondrocranium and oral anatomy. The more complete description provided below (inconsistencies with Cei [1980] noted) is based on 35 specimens (Stages 27-41, KU 160144) from 3 km S Epizana, Cochabamba, Bolivia, 2825 m. A specimen in Stage 38 (KU 160144) has the following measurements: total length 57.4, body length 19.5, tail muscle height at base 6.3, tail muscle width at base 5.0, maximum dorsal fin height 4.4 located 24.6 from body ter- minus, maximum ventral fin height 3.9 located 26.0 from body terminus, body width 11.9, body height 9.9, eye di- ameter 2.7, pupil diameter 1.1, interorbital distance 4.3, greatest and least nostril diameters 0.9 x 0.5, internarial distance 3.0, snout-nostril 3.1, snout-eye 5.7, snout-spiracle 11.5, eye-nostril 1.7, and transverse oral disc diameter 4.9. Other major characteristics include a ventral oral disc, marginal papillae with median gap on upper labium, spi- racle sinistral, eyes dorsolateral, vent dextral, dorsal fin terminating at the dorsal extent of the tail-body junction. The LTRF is 2(2)/4(1). The outer wall of the spiracle is longer than the inner wall, and emerges from the ventrolateral surface of the body at midlength. The inner wall of the spiracular tube is attached to the body for most of its length; terminally it is free and hence the tube is free. The tube is directed posterodorsally and narrows at a round, posteriorly di- rected aperture. The vent tube is dextral, and is positioned at the posterolateral surface of a large saccular structure that underlies the limb buds and is contiguous with the ventral fin. The vent aperture is directed posteroventrally. The eyes are positioned dorsolaterally and directed later- ally and slightly dorsally. Cei (1980) described the eyes as dorsal, but it is apparent from his figure that their position is dorsolateral in our terminology. The nares are oval, slightly medial to the center of the eyes, recessed, and weakly rimmed; the rim of the medial edge is raised slightly. The snout is bluntly rounded in dorsal view and profile. The body is elongately ovoid in dorsal view; in lateral view depressed dorsally and ventrally, becoming deeper posteriorly toward the bulging gut. The oral disc is slightly emarginate posterolaterally and is completely circumscribed by marginal papillae except for a median gap in the middle third of the upper labium; several other specimens have fewer marginal papillae an- teriorly on the upper labium and medially on the lower labium. Laterally, there are few submarginal papillae, but there are flaps that possess keratinized teeth forming bro- ken vertical rows. The LTRE is 2(2)/4(1), with long, nar- row, closely spaced teeth on all rows except P-4 which possesses much smaller teeth. The labial tooth rows of the upper labium do not vary within the series, but only three rows on the lower labium are present in some specimens, in one of which the third row has a medial gap. This varia- tion is consistent with the LTRF reported by Cei (1980) of 2(1-2)/3(1) with the exception of the gap in the upper an- terior row of his specimen. The upper jaw sheath is mod- erate in width, widely arched, and finely serrate. The lower sheath is narrow, broadly V-shaped, and finely serrate. The caudal musculature is highest at the tail-body junc- tion and is the same height posteriorly to midlength, where it narrows gradually to a pointed tail terminus. Cei (1980) figured the tail musculature as terminating prior to the end of the tail; this is not the case in any of our specimens and is attributed to an error in his figure. In lateral view, the dorsal fin terminates at the dorsal tail-body junction, is highest on posterior third of the tail, and is shorter than musculature at midlength (i.e., taller posteriorly, shorter anteriorly). The ventral fin is shorter than the dorsal fin, except for the anterior and posterior extremes which are the same height as the dorsal fin. The ventral fin gradu- ally increases in height from anterior to posterior until it abruptly narrows at the tail terminus. The ventral fin is shorter than the musculature at midlength (taller only on posterior third). The greatest height of tail is along the posterior third of the length. The tail terminates as a pointed tip. In preservative, the dorsum of the body is uniformly pigmented with brown chromatophores with a few, tiny white flecks and larger, distinct, dark brown spots scat- tered throughout. The venter is transparent white with few melanophores, mostly laterally; the intestine is clearly vis- ible. Dorsal pigmentation continues onto the dorsum of caudal musculature lateral to the base of the dorsal fin for the entire length of the tail. The remainder of the tail mus- culature is opaque yellowish brown, with many scattered melanophores, some of which form diffuse blotches. The fins are transparent with scattered white flecks and few melanophores, some of which form many small blotches. In life, the body is olive-tan above and deep bronze with silvery lichenous marks below; the tail is olive-tan with brown streaks, and the iris is bronze (W. E. Duellman field notes, 11 November 1974). Distribution and ecology.—Hyla andina is widely dis- tributed in the eastern Andes and the Andean precordilleras from northern Bolivia southward to Prov. Catamarca, Argentina (Fig. 9). In the northern part of its range it occurs at elevations of 1650-3400 m, but in the southern part (provincias Tucuman and Catamarca), it occurs at elevations of 500-1640 m. At high elevations, the frog inhabits montane grassland; in the northern part of its range it descends into cloud forest and temperate forest, whereas in the southern part of its range it occurs in premontane deciduous forest. Throughout its range, Hyla andina is always near water, either lotic or lentic—viz., lagoons, ponds, large or small streams, or roadside seepages. By day, individuals perch on low (usually less than 1 m above the ground) branches D2. ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS of trees or bushes, or they are on the ground at the edges of streams, especially above treeline. Occasionally they are on tall grasses or under stones. Individuals often are in direct sunlight. In contrast to most other tree frogs, they jump into the water when disturbed during the day. At night, males call from branches above or at the edge of water, stones in streams, the ground at the edge of water, or in shallow water in ponds, marshes, or pools in streams. At some places, especially above treeline and in temper- ate forests, the species is abundant, whereas it seems to be much less common along streams in cloud forest, which it seems to colonize via roadside seepages. In cloud forest, the frogs seem to prefer still or very slow-moving water, in contrast to the usual fast-moving water in most streams. Tadpoles have been found in shallow ponds, marshes, roadside ditches, and quiet pools in streams. These obser- vations generally concur with the results of a detailed study of the ecology of tadpoles of this species in the Sierra de Metan, Prov. Salta, Argentina by Lavilla (1980). Hyla andina apparently occurs in sympatry with H. marianitae at Carahuasi, Depto. Cochabamba, and in the vicinity of Samaipata, Depto. Santa Cruz, Bolivia; also, it has been reported from Cochabamba, Depto., Cochabamba, Bolivia, a locality from which H. albonigra has been reported. Remarks.—According to the catalogue in the Zoologisches Staatsammlung Munchen, there were origi- nally five syntypes (ZSM 5/1922 from Argentina; ZSM 21/ 1922 and 23/1922 from Caspinchango, Prov. Catamarca, Argentina; ZSM 22/1922 from Famabalasto, Prov. Catamarca, Argentina; ZSM 38/1922 from northern Argen- tina) that are no longer extant. The three existing syntypes (ZSM 18/1922) were collected by F. Weiser in November 1921 at Nacimiento, Prov. Catamarca, Argentina. When ex- amined on 30 July 1969, these specimens were in fairly good condition. The dorsum was gray with creamy-tan, irregular dashes and streaks; a white transverse supracloacal stripe and dark bars in the groin and on the posterior surfaces of the thighs were evident. The polytypic species Hyla pulchella was reviewed by Barrio (1965), who recognized five subspecies based on size, coloration, and advertisement call. Lutz (1968) rec- ognized another subspecies, H. pulchella joaquini in Santa Catarina, southeastern Brazil. However, one of these sub- species, H. pulchella prasina Burmeister from moderate el- evations in southeastern Brazil, was recognized as a dis- tinct species by Lutz (1973). As presently recognized, the nominate subspecies is distributed in mesic lowlands in extreme southeastern Brazil, Uruguay, and northeastern Argentina; H. pulchella cordobae Barrio is restricted to the isolated Sierra de Corboba in north-central Argentina, and the other two subspecies are in the Andes and Andean precordilleras (Barrio, 1965; Cei, 1980). Specimens lacking black bars on the flanks and thighs from the Andean slopes in Provincia La Rioja and extreme southern Provincia Cajamarca, Argentina, were referred to H. pulchella riojana Koslowsky by Barrio (1965), who recognized Andean populations from elsewhere in Provincia Catamarca north- ward into Bolivia as H. pulchella andina. Structurally, all of these frogs are similar, but differences in color patterns and advertisement calls suggest that each might be distinct species. This suggestion is supported by discontinuities in distribution, although the ranges of H. pulchella riojana and H. pulchella andina may be parapatric or overlap. By recog- nizing H. andina as a species distinct from H. pulchella, we are inferring that H. pulchella riojana also is specifically dis- tinct from H. andina. On the other hand, if future studies show that these two taxa are conspecific but distinct from H. pulchella, the oldest available name is Hyla riojana Koslowsky (1895). - As noted in the description of coloration, there is a vast array of color patterns in Bolivian populations. In some cases, consistent interpopulational differences seem to ex- ist. However, we have limited confidence in the designa- tion of coloration in preserved specimens, and we have not been able to quantify color patterns. Our assignment of all of these populations is supported by the discrimi- nant functions analysis of morphometrics. Populations considered to have the typical pattern of Hyla andina (dor- sum green with or without dark blotches and irregular white stripes on shanks and dorsolateral regions of body) in Central Bolivia occur only in localities above treeline (i.e., La Paz, Departamento La Paz; Colomi, Lopez Mendoza, and Tiraque, Departamento Cochabamba). Be- low treeline (around 3000 m, depending on the areas con- sidered), patterns and color vary dramatically. Populations living in the same general area below treeline may vary depending on the altitude, as is evident in those occurring in the area of La Siberia, Depto. Cochabamba. In Argen- tina, both green and brown individuals are known for the latitudinal length of the range of the species. Also, speci- mens from the southern part of the range (provincias Tucuman and Catamarca, Argentina) tend to have a white stripe on the outer edge of the shank and tarsus. Hyla balzani Boulenger Hyla balzani Boulenger, 1898:132.—Holotype: MCSN 28872 from “Prov. Yungas, 1600 metres altitude” (= Coroico and Chulumani, Depto. La Paz), Bolivia. Hyla callipleura Boulenger, 1902.—Syntypes: BM 1947.2.13.64-73 from Charuplaya, 1350 m, Bolivia, and BM 1947.2.13.74 from San Ernesto, Bo- livia. BM 1947.2.13.65 here designated as lectotype. New synonymy. Diagnosis.—A moderate-sized species in the Hyla pulchella Group characterized by: (1) snout rounded in dorsal view and in profile; (2) skin on dorsum smooth; (3) width of disc on Finger II equal to or slightly less than diameter of tympanum; (4) tarsal fold low; (5) dorsum unicolor or having a pattern consisting of transverse dark marks or broad irregular dark mark enclosing pale spots; (6) flanks brown with small pale spots; (7) posterior sur- faces of thighs usually brown with small cream spots. One other species, Hyla marianitae, is like H. balzant in having dark flanks with pale spots. It differs from H. balzani by having a subacuminate snout in dorsal view, usually ANDEAN Groups OF HYLA 23 irregular blotches or reticulations on the dorsum, cream and brown mottling on the posterior surfaces of the thighs, and a weak tarsal fold. Hyla melanopleura has dark brown flanks with small pale dots only along the ventral edge of the flanks. Description.—n = 61 do, 8 99. Body moderately robust; SVL in males to 50.4 mm, in females to 50.9 mm; head width about equal to head length, usually not as wide as body; head width 32.0-37.0% (X= 34.0) of SVL; head length 32.0-— 39.0% (X = 35.0) of SVL; snout rounded in dorsal view and in profile; interorbital distance 98-177 % (x = 128) of width of eyelid; diameter of eye 93-159% (x = 124) its distance from nostril; tympanum separated from eye by distance about equal to diameter of tympanum; diameter of tym- panum 37.0-70.0% (xX = 53.0) of diameter of eye. Breadth of forearm 58.7—71.8 (X = 65.0, 1 = 10) of forearm length in males, 38.8-41.2 (X= 39.8, n = 4) in females; ulnar fold proxi- mally and row of low, round tubercles distally on fore- arm; webbing formula for fingers: II 2-—(3—3*) HI 2%2—2* IV; width of disc on Finger III equal to or slightly less than diameter of tympanum; prepollex blunt, projecting at right angle in males, short and blunt in females; nuptial spines absent. Hind limb slender; tibia length 49.0-58.0% (x= 54.0) of SVL; foot length 41.0-66.0% (x = 45.0) of SVL; inner tar- sal fold low; inner metatarsal tubercle ovoid, widest proxi- mally, not visible from above; webbing formula for toes: I (1-1%)—(2—2) I (1-1*) —(2-2*) MI (1-144) (2-2) Tv (2— 2*)—(1-1*) V. Skin on dorsum and flanks smooth. Denti- gerous processes of vomers short, transverse, narrowly separated medially, posteromedial to rounded choanae, bearing total of 7-24 (x = 14.2) teeth. Coloration in preservative: Side of head and dorsal sur- faces of head, body, and limbs tan, beige, or grayish brown with or without irregular dark brown, usually transverse marks; as many as five such marks including interorbital bar (in some individuals, V-shaped with apex posteriorly, connected or not to transverse mark in scapular region); markings transverse across entire dorsum and more or less parallel to sinuous (in some individuals interconnected); some individuals with only irregular marks on dorsum; flanks usually dark brown (sharply demarcated from dor- sal color) usually with creamy white, irregular spots; dor- sal surfaces of limbs usually with dark brown transverse bars; posterior surfaces of thighs tan to dark brown with creamy-white blotches or spots; cream transverse stripe above cloacal opening; venter dull creamy white or gray- ish brown, with or without small brown flecks on throat, chest, and limbs; throat usually paler than belly; palmar and plantar surfaces tan or brown. Coloration in life: The dorsum is brown or green, usu- ally with darker markings; the flanks and posterior sur- faces of the thighs are dark brown or green with creamy- white spots. The variation in coloration is evident from the following descriptions of living individuals (Fig. 3). KU 139212 from 4 km SW Santa Isabel, Depto. Cuzco, Peru.—Dorsum tan with brown flecks and blotches nar- rowly outlined with creamy tan; chin creamy white with black flecks; venter creamy white anteriorly becoming pale gray posteriorly on belly and legs; side of head below cream canthal and supratympanic stripe bronze-brown, bordered below by metallic green; labial stripe dusty cream; flanks and posterior surfaces of thighs dark brown with cream spots; supracloacal stripe and stripes on outer edges of feet creamy white; plantar surfaces and webbing gray; iris deep bronze, darkest anteriorly and posteriorly; palpebrum clear (W. E. Duellman field notes, 16 January 1971). KU 163315 from 4 km SW Santa Isabel, Depto. Cuzco, Peru.—Dorsum tan with faintly darker brown markings; side of head and anterior flanks pale green; posterior sur- faces of thighs and flanks dark brown with creamy-white spots; venter pale bluish gray; iris pinkish brown (W. E. Duellman field notes, 10 February 1975). USNM 332412-14 from Cerros de Tavara, Depto. Puno, Peru.—Dorsum green or brown with white dorsolateral line bordered by reddish brown; irregular white flecks on flanks; white heel and supracloacal stripes; hidden sur- faces of thighs mustard; webbing on feet yellow; toe pads green; iris yellow (Lily O. Rodriguez field notes, 1-2 June 1992) KU 183326-28 from 58.1 km W Villa Tunari, Depto. Cochabamba, Bolivia.—Dorsum tan or brown with dark brown markings; ventral surfaces dark brown or brown- ish green, chin paler; flanks black with tan markings; pos- terior surfaces of thighs black with tan spots; ventral sur- faces of hands and feet dark brown; ventral surfaces of toe pads pale green; bones green; iris pale gold (D. C. Cannatella field notes, 21 January 1979). KU 183330-34 from 4.4 km E Chulumani, Depto. La Paz, Bolivia—Dorsum brown or olive-brown with darker brown markings in some individuals; dark brown stripe (with touch of green) from nostril to tympanum and onto flanks; portion of upper lip below eye gold; flanks and anterior and posterior surfaces of thighs dark brown with pale tan markings; belly and chin off-white “gray”; ven- tral surfaces of limbs gray brown; pads greenish; iris gold with dark brown horizontal stripe; bones green; one indi- vidual (KU 183331) with white supracloacal and tarsal stripes gold (D. C. Cannatella field notes, 21 January 1979). KU 183335-36 from 33.9 km E Unduavi, Depto. La Paz, Bolivia.—Dorsum brown with olive and rust markings; belly off-white; chin and ventral surfaces of limbs dark gray; flanks and posterior surfaces of thighs black with yellowish-white markings; webbing brown; supracloacal and tarsal stripes off-white; upper lip gold in KU 183335 gold (D. C. Cannatella field notes, 30 January 1979). KU 18333740 from Coroico, Depto La Paz, Bolivia.— Dorsum green or brown with green blotches; flanks and posterior surfaces of thighs dark brown with off-white markings; brown to brown and green stripe from nostrils to flanks; belly off-white; chin yellowish white; ventral sur- faces of limbs pale gray; supracloacal and heel stripes off- white; bones green; iris pale gold (D. C. Cannatella field notes, 31 January 1979). 24 a ® 2 -70 rs) ® ia) —80 2 4 6 8 10 Kilohertz a Oo 6 S 4 SZ 2 Se 0 2.0 2 w 2 0 © a -2.0 2.0 2.2 2.4 2.6 Seconds Fig. 13. Advertisement call of Hyla balzani, KU Tape 1229, recorded at 21°C at 4 km WSW Santa Isabel, Depto. Cuzco, Peru. Spectrum, audiospectrogram, and waveform; arrow in audiospectrogram indicates point of spectrum. KU 18334146 from Coroico, 1750 m, Depto La Paz, Bo- livia—Dorsum brown, olive, or yellowish green (D. C. Cannatella field notes, 4 February 1979). Likewise, Reynolds and Foster (1992) commented on the variation in coloration of specimens from Departamento Cochabamba, Bolivia (USNM 257819-33). They stated that the dorsum was reddish brown to golden brown with pale to dark markings, and that the flanks and posterior surfaces of the thighs were mottled dark choco- late-brown and cream. Also, they described newly meta- morphosed young as being opalescent greenish brown dorsally with small dark spots and stripes and the limbs as yellow green. The bones are green. Life history.—The advertisement call consists of a short, low note repeated at a rate of 2.54 notes per minute. Analy- sis of two recordings (KU Tapes 1228-29) made at 17° and 21°C at the Rio Cosnipata, Peru, revealed a note duration of 0.17 sec, a pulse rate of about 80 pulses per sec, and harmonics at 1150, 1400, 1650, and 1900 Hz, with the low- est harmonic being dominant (Fig. 13). Reynolds and Fos- ter (1992) described the call as a sharp, moderately high- pitched “tink.” The recording tentatively identified as Hyla callipleura (= H. balzani) by Marquez at al. (1993) is actually H. marianitae. Sixty-four specimens (Stages 25-45) of two series of tad- poles (KU 164052, 164053-54) are from 4 km SW Santa Isabel, Departamento Cuzco, Peru, 1700 m; one individual (KU 164054) of the latter series was allowed to develop to Stage 45 before being preserved four days after collection. A specimen in Stage 38 (from KU 164053) has the fol- lowing measurements: total length 41.7, body length 14.8, SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS tail muscle height at the base 5.3, tail muscle width at base 4.2, maximum dorsal fin height 3.5 located 20.9 from body terminus, maximum ventral fin height 3.0 located 20.0 from body terminus, body width 10.3, body height 7.6, eye di- ameter 1.9, pupil diameter 0.7, interorbital distance 3.4, greatest and least nostril diameters 0.5 x 0.2, internarial distance 3.3, snout-nostril 3.0, snout-eye 4.8, snout-spiracle 8.5, eye-nostril 1.5, transverse oral disc diameter 4.3. Other major characteristics include a ventral oral disc, marginal papillae with median gap on upper labium, sinistral spi- racle, dorsolateral eyes, dextral vent, and the dorsal fin terminating at the dorsal extent of the tail-body junction. The LTRF is 3(1,3)/4(1). The outer wall of the spiracle is longer than the inner wall, and emerges from the ventrolateral surface of the body at midlength. The inner wall of the spiracular tube is attached to the body for most of its length; terminally it is free and hence the tube is free. The tube is directed poste- riorly and narrows to forma round, posteriorly and slightly dorsally directed aperture. The vent tube is dextral, and is positioned at the posterolateral surface of a large saccular structure that underlies the limb buds and is contiguous with the ventral fin. The vent aperture is directed posteroventrally. The eyes are positioned dorsolaterally and directed laterally and slightly dorsally. The nares are oval, slightly medial to the center of the eyes, recessed, and weakly rimmed, with the rim of the medial edge slightly raised. The snout is broadly rounded in dorsal view and profile. The body is oval in dorsal view; in lateral view depressed dorsally and ventrally becoming deeper poste- riorly toward the bulging gut. The oral disc is slightly emarginate posterolaterally and is completely circumscribed by marginal papillae except for a median gap in the middle fifth of the upper labium; there are fewer marginal papillae medially on the lower labium. There are many submarginal papillae and flaps laterally. The LTRF is 3(1,3)/4(1) with long, narrow, closely spaced teeth. There is intraseries variation in the LTRF that seems to coincide with development. In KU 164053, tad- poles early in Stage 25 have a LTRF of 2(2)/3(1). A fourth lower row is gained later during Stage 25, this formula of 2(2)/4(1) is seen in specimens as advanced as Stage 30. A third upper row with a medial gap, thereby resulting in a LTRF 3(1,3)/4(1), is seen as early as, but usually later than, Stage 26. In KU 164052, tadpoles early in Stage 25 have a LTRF of 2(1)/4(1); later in this stage, tadpoles possess the gap in the other upper row, 2(2)/4(1). One specimen at Stage 31 of this series has the LTRF 3(1,3)/4(1) seen in ad- vanced specimens of the first series. The upper jaw sheath is moderately wide, widely arched, and moderately ser- rate. The lower sheath is narrow, broadly V-shaped, and moderately serrate (Fig. 14). The caudal musculature is highest at the tail-body junc- tion and is the same height to midlength where it narrows gradually to a rounded (narrow and pointed in most other specimens) tail terminus. In lateral view, the dorsal fin ter- minates at the dorsal tail-body junction, is highest on the ANDEAN Groups OF HYLA D5) Fig. 14. Tadpole of Hyla balzani, KU 164053, Stage 38, total length 41.7 mm. posterior fourth of tail, and is shorter than the muscula- ture at midlength (taller posteriorly, shorter anteriorly). The ventral fin is shorter than the dorsal fin on the posterior half of the tail, taller for the anterior half; it does not vary in height until the posterior fourth of the tail where it is taller until narrowing abruptly at the tail terminus. The ventral fin is shorter than the musculature at midlength (taller only on posterior fourth). The greatest height of the tail is on the posterior fourth of the tail. The tail terminates as a rounded tip. (The tail tip is narrow and pointed in most other individuals of the series.) In preservative, the dorsum of the body is uniformly pigmented with dark brown chromatophores and few white flecks anterior to the level of the spiracle. The ven- ter has little or no pigment laterally, and possesses scat- tered white flecks; the intestine is clearly visible. The dor- sal pigmentation continues posteriorly onto the dorsum of the caudal musculature lateral to the base of the dorsal fin posteriorly to tail tip. The remainder of the caudal mus- culature is opaque yellowish brown with scattered mel- anophores; some are located in patches forming diffuse, pale brown blotches. The fins are transparent gray with few scattered melanophores forming small diffuse blotches on the dorsal fin, but not on the ventral fin. Both fins have few scattered white flecks. In life (KU 164052), the body is grayish brown above with a dark brown interorbital bar, and the belly is silvery gray. The caudal musculature is tan with dark brown bars dorsally, and the fins are trans- lucent with brown flecks; the iris is reddish bronze (W. E. Duellman field notes 10 February 1975). Distribution and ecology.—Hyla balzani primarily is an inhabitant of cloud forests at elevations of 1200-2210 m on the eastern slopes of the Andes from central Peru to cen- tral Bolivia (Fig. 9), but the species is known from lower elevations (500 m at San Ernesto, Depto. La Paz, Bolivia; 620 m at La Union, Depto. Puno, Peru; 900 m at Cerros de Tavara, Depto. Puno, Peru; and 925 m at Huancabamba, Depto. Pasco, Peru). One specimen (USNM 98928) from “16 km E Loja City, 3000 m, Prov. Loja, Ecuador” is part of a collection received from Clodoveo Carrion; in the USNM catalogue, the date of collection as “Aug. 22, 1926.” We are skeptical of this record, because (1) the locality is about 500 km NNW of the northernmost locality of H. balzani(Huancabamba, Depto. Pasco, Peru); (2) the local- ity is north of the Huancabamba Depression, a major dis- tributional barrier to anuran inhabitants of cloud forest (Duellman, 1979; Duellman and Wild, 1993); (3) the given elevation is 790 m higher than any other known for the species; and (4) extensive collections from the Abra de Zamora, 15-16 km E Loja contain no specimens of this species. Although it seems to be primarily a forest-dweller, the species can tolerate a high degree of environmental dis- turbance. It has been found mainly along streams, ponds, and roadside seepages. At the Rio Corani, Depto. Cochabamba, Bolivia, the species was common in a small pond (about 25 m diameter) close to the river. There, males called at night from stones in the middle of the pond, or perched on stones or vegetation (<1.5 m above ground) at the shore; also, they were observed on stones in the river. At Cerro Uchumachi, Depto. La Paz, Bolivia, individuals were found in a sloping, flooded meadow with scattered bushes; similarly, frogs were found in a flooded field at Coroico, Depto. La Paz, Bolivia. However, at most locali- ties, males call from stones in, or vegetation above, small streams with moderate current. Tadpoles have been found in grassy pools, shallow roadside ditches, and pools in streams. Hyla balzani has been found syntopically with Hyla armata in some streams in Bolivia and southern Peru— Paracti and Serrania Bellavista in Bolivia and the Rio Cosnipata in Peru. Remarks.—Although the holotype of Hyla balzani (MSNG 28872) is a small male with a SVL of 32.9 mm, it has vocal slits and well-developed prepollical spines. Com- parison of this specimen with series of frogs that had been identified as H. callipleura revealed that they represent one taxon, the earliest name for which is H. balzani Boulenger, 1898. Thus, H. callipleura Boulenger, 1902, is a junior syn- onym of H. balzani Boulenger, 1898. In the type descrip- tion of Hyla callipleura, Boulenger (1902) compared the spe- cies only with H. crepitans and made no mention of H. balzani. In the type description of H. balzani, Boulenger (1898:132) reported the specimen from “Prov. Yungas, 1600 metres altitude,” Bolivia. However, Boulenger (1898:128) listed the localities where L. Balzan had collected. The only locality in the Yungas is “Coroico and Chulumani, Prov. Yungas, 1600 metres alt.” This provides a reasonably pre- cise locality, because Coroico and Chulumani are only a few kilometers apart on the slopes of the Andes in Depto. La Paz, Bolivia. 26 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS When the type series of Hyla callipleura was examined by Duellman on 16 June 1969, all specimens were in rea- sonably good condition. BMNH 1947.2.13.65 is an adult male with a SVL of 40.4 mm and the best-preserved color pattern; therefore, it is designated as the lectotype. Two other syntypes are adult males with SVLs of 37.2 and 43.3 mm; one is a gravid female with a SVL of 40.0 mm, and the other specimens are immature. One specimen (CET A1453) from 10 km NW of Masicuri, Depto. Santa Cruz, Bolivia, may belong to this species. It is a male with a SVL of 34.8 mm, but it lacks a distinct tarsal fold. In preservative, the dorsum is pale brown with darker brown markings—a distinct triangular mark in the occipital region with the apex between the orbits and faint transverse marks on the body. The sides of the head and flanks are dark brown with three vertical pale marks on the posterior part of the flanks. A wide, pale cream labial stripe is present., and a pale cream stripe extends along the canthus rostralis, outer edge of upper eyelid, supratympanic fold, and along the dorsolateral surface of the body to the groin. White stripes are present on the heels and above the cloacal opening. The anterior and posterior surfaces of the thighs are creamy-tan, and the venter is creamy white. Color photographs of the living individual show the dorsum to be dull tan with faint brown mark- ings, the flanks dark brown, and the iris dull tan; the la- bial, dorsolateral, supracloacal, and heel stripes are pale creamy gray. Ayla marianitae Carrizo Hyla marianitae Carrizo, 1992:18.—Holotype: MACN 32330 from the Rio Pescado, Sierra de las Pavas, Parque Nacional Bariti, Departamento Oran, Provincia Salta, Argentina. Diagnosis.—A moderate-sized species in the Hyla pulchella Group characterized by: (1) snout subacuminate in dorsal view and rounded in profile; (2) skin on dorsum smooth; (3) width of disc on Finger III less than diameter of tympanum; (4) tarsal fold low, weak; (5) dorsal color pattern consisting of irregular blotches or reticulations; (6) flanks brown with pale spots; (7) posterior surfaces of thighs with cream and dark brown mottling. One other species in the Hyla pulchella group has dark flanks with pale spots. Hyla balzani differs by having a rounded snout in dorsal view, a more distinct tarsal fold, transverse dark marks on the dorsum, and the posterior surfaces of the thighs dark brown with discrete pale spots. Hyla melanopleura has dark brown flanks with small pale dots along the ventral edge of the flanks. Description.—1 = 26 Go, 6 99. Body robust; SVL in males to 56.8 mm, in females to 56.5 mm; head width about equal to head length, not as wide as body; head width 31.0- 37.0% (x = 34.0) of SVL; head length 32.0-36.0% (x = 34.0) of SVL; snout slightly subacuminate in dorsal view, rounded in profile; interorbital distance 114-183% (x = 147) of width of eyelid; diameter of eye 102—151% (x = 119) its distance from nostril; tympanum separated from eye by distance about equal to diameter of tympanum; diameter of tympanum 41.0-70.0% (xX = 55.0) of diameter of eye. Breadth of forearm 64.3-73.9 (x = 68.1, n = 10) of forearm length in males, 40.5-44.3 (x= 43.2, n = 4) in females; ulnar fold proximally and row of low, round tubercles distally on forearm; webbing formula for fingers: II (2—2*)—(3— 3*) WT 24%—(2-2*) IV; width of disc on Finger III slightly less than diameter of tympanum; prepollex blunt, project- ing at right angle in male, short, blunt in female; nuptial spines absent and nuptial excrescences absent in breeding males. Hind limb moderately slender; tibia length 50.0— 59.0% (x = 54.0) of SVL; foot length 41.0-52.0% (x= 44.0) of SVL; inner tarsal fold low, weak, curved; inner metatarsal tubercle ovoid, widest proximally, not visible from above; webbing formula for toes: I 1—(1%-2>) If 1—(2-2*) IM (— 1%)—(2-2*) IV (2-2*)—1 V. Skin on dorsum and flanks smooth. Dentigerous processes of vomers moderately long, transverse, narrowly separated medially, posteromedial to rounded choanae, each bearing total 6-22 (x= 12.3) teeth. Color in preservative: Side of head and dorsal surfaces of head, body, and limbs tan, gray, or brown, uniform to boldly patterned with irregular small dark markings (form- ing network pattern in some individuals), irregular large dark blotches, or transverse dark bars; flanks usually with dark markings (encircling creamy-white spots in some in- dividuals); same kind of pattern usually present on ante- rior and posterior surfaces of thighs; dorsal surfaces of limbs usually unpatterned, but with indistinct transverse dark bars in some; creamy-white labial stripe usually present; transverse cream stripe above cloacal opening; venter cream, tan, or gray; throat and chest with or with- out brown flecks; palmar and plantar surfaces same color as limbs. Color in life: The dorsal ground color usually is tan or brown, but a few individuals are vivid green (Fig. 3). This variation is apparent from the following descriptions of living individuals. CET A89 from Arroyo Las Juntas, 8 km NW Samaipata, Depto. Santa Cruz, Bolivia —Dorsal surfaces of body and limbs yellowish tan with some indistinct, irregular dark blotches; flanks and posterior surfaces of thighs gray, en- circling dots of the same color as dorsum; belly pale yel- low, throat yellow; ventral surfaces of limbs gray; iris golden-yellow (I. De la Riva field notes, 7 March 1990). CET A88 from Carahuasi, Depto. Cochabamba, Bo- livia.—Dorsal surfaces of body and limbs mostly uniformly tan; flanks cream with large dark brown, connected blotches (I. De la Riva field notes, 9 March 1990). CET A109 from Rio Cheyo, Amboro National Park, Depto. Santa Cruz, Bolivia—Dorsal surfaces of body and limbs mostly uniformly pale green; flanks brown, encir- cling creamy white blotches; iris beige (I. De la Riva field notes, 16 March 1990). Life history.—The advertisement call was described (as Hyla callipleura) by Marquez et al. (1993). The call consists of a single note repeated at a rate of 15-23 (x = 18.7) notes per min; the notes have a duration of 0.025-0.059 (x= 0.036) ANDEAN GRoupPs OF HYLA Di Decibels 4 f=) foe) [e) 10 Kilohertz pr) [o) ats [o) N 1= oO ds re) NZ 0.0 0.5 1.0 1.5 2.0 Seconds Fig. 15. Advertisement call of Hyla marianitae, recorded at Tablasmontes, Depto. Cochabamba, Bolivia. Spectrum, audiospectrogram, and waveform; arrow in audiospectrogram indicates point of spectrum. sec. Analysis of recordings of five calls made at La Siberia, Depto. Santa Cruz, Bolivia, revealed a pulse rate of 44-45 (x = 44.4) pulses per second and harmonics at about 880, 1250, and 1700 Hz, of which the lowest is the dominant frequency (Fig. 15). The tadpoles are unknown; presumably they develop in the streams along which adults have been found. Distribution and ecology.—Hyla marianitae is known from elevations of 700-2650 m on the eastern slopes of the Andes and Andean precordilleras from central Bolivia to northern Argentina (Fig. 9). It inhabits cloud forests, low montane rainforest, subhumid, and even dry temperate forests, where it is found in the immediate vicinity of streams. Some of these streams are permanent, but those in temperate valleys can become intermittent or dry by the end of the dry season. At night, individuals perch on, and call from, tree branches near the streams or, more of- ten, on large stones in the middle or at the edge of streams. By day, individuals can be found under stones on the banks of streams. The species is common at some localities. At Carahuasi, Depto. Cochabamba, Bolivia, many individu- als and amplectant pairs were observed on 9 March 1990. A female of a pair kept in a plastic bag laid pigmented eggs 2.6 mm in diameter. Males commonly have many scars on the dorsum; presumably these are injuries caused by prepollical spines during male-male encounters. Hyla marianitae is allopatric to H. balzani. On the other hand, H. marianitae is sympatric with H. andina at Carahuasi, Depto. Cochabamba, Bolivia; it also occurs sym- patrically with H. armata at El Chapé and Quebrada Huertillas, Depto. Santa Cruz, Bolivia. Remarks.—In describing this species from two speci- mens from Parque Nacional Baritu in northwestern Ar- gentina, Carrizo (1992) apparently was unaware that it was widely distributed to the north and west in Bolivia. More- over, although Carrizo (1992) described and illustrated the prepollical spine, he did not associate the new species with the Hyla pulchella Group. Although the holotype (MACN 32330) is slightly faded, a dorsal pattern of irregular dark markings is perfectly vis- ible; on the flanks, irregular pale spots extend from the axilla to the groin. The snout is subacuminate in dorsal view and rounded in profile; the fingers are about one- third webbed, and the feet are nearly fully webbed. The tarsal fold is not discernible. Our comparison of the holo- type of Hyla marianitae with specimens from Bolivia con- vinces us that they are conspecific. Hyla melanopleura Boulenger Hyla melanopleura Boulenger, 1912:185.—Syntypes: BM 1947.2.13.54—58 from Huancabamba, Departamento Pasco, Peru; BM 1947.2.13.56 designated as lectotype. Diagnosis.—A small species in the Hyla pulchella Group characterized by: (1) snout rounded in dorsal view and bluntly rounded in profile; (2) skin on dorsum smooth; (3) width of disc on Finger III less than diameter of tympa- num, (4) tarsal fold low; (5) dorsal color pattern consisting of triangular interorbital mark and irregular dashes on body; (6) flanks dark brown with small pale dots along the ventral edge; (7) posterior surfaces of thighs uniform dark brown. The only other species in the Hyla pulchella Group lack- ing large marks on the flanks is H. palaestes, in which the flanks and posterior surfaces of the thighs are not darker than the dorsum and the dorsal color pattern consists of irregular, narrow transverse marks. Moreover, in H. palaestes, the width of the disc on Finger III equals the di- ameter of the tympanum. Description.—n = 1 &, 4 99. Body moderately robust; SVL in males to 43.6 mm, in females to 47.7 mm; head width equal to head length, not as wide as body; head width and head length 32.7-33.2% (x = 33.0) of SVL; snout rounded in dorsal view, bluntly rounded in profile; interorbital dis- tance 124-136% (x = 131) of width of eyelid; diameter of eye 108-155% (x = 126) its distance from nostril; tympa- num separated from eye by distance equal to diameter of tympanum; diameter of tympanum 56.3-64.7 % (x = 60.8) of diameter of eye. Breadth of forelimb 41% of forearm length in male, 33% in females; ulnar fold proximally and row of low, round tubercles distally on forearm; webbing formula for fingers: Il (2—2)—(3-3) II 24%4—(2-—2") IV; width of disc on Finger III slightly less than diameter of tympanum; prepollex pointed, projecting at right angle in male, short, blunt in females; nuptial spines absent; nup- tial excrescence brown on inner side of base of thumb in Fig. 16. mm SVL. Hyla melanopleura, BMNH 1947.1.13.54, adult female, 47.7 breeding male. Hind limb slender; tibia length 51.8-63.3% (X= 56.0) of SVL; foot length 40.6-44.4% (x = 42.7) of SVL; inner tarsal fold low, curved; inner metatarsal tubercle ovoid, widest proximally, not visible from above; webbing formula for toes: I (1-1%)—(2-2"*) II (1-1*)—(2*-2') III (1- 12)—(2-2'%) IV (2—2*)—(1-1*) V. Skin on dorsum and flanks smooth. Vomerine odontophores short, transverse, nar- rowly separated medially, between ovoid choanae, each bearing total 3-6 (x = 4.5) teeth. Color in preservative: Dorsum tan with irregular brown markings—BMNH 1947.2.13.54: narrow triangle on head connected by narrow middorsal mark to long inverted V extending from scapular region to sacrum; irregular dashes postsacrally (Fig. 16). BMNH 1947.2.13.56: interorbital bar; irregular paravertebral marks extending from scapular region to sacrum; irregular dashes dorsolaterally and postsacrally. BMNH 1947.2.13.58: triangle with base be- tween inner edges of eyelids and apex in scapular region; ovoid mark with tan center between scapular region and sacrum; diamond-shaped mark postsacrally. Irregular, transverse brown marks on forearms, shanks, and tarsi; flanks and anterior and posterior surfaces of thighs dark brown (with small white flecks on flanks in BMNH 1947.2.13.58). Side of head brown; creamy-white labial stripe (only posteriorly from point below orbit in BMNH 1947.1.13.54); white transverse stripe above vent (absent in BMNH 1947.2.13.54). Venter dull creamy white with small brown flecks on throat, chest, undersides of arms, and ventral surfaces of tarsi; webbing on hand tan, on feet, brown. Color in life: Unknown. Boulenger (1912:186) described the color of the syntypes as follows: “Greyish or reddish brown above, sides blackish; the back may be spotted or dotted with brown, the sides dotted with white; limbs with irregular dark cross-bands; lower parts dirty white. Life history—Nothing is known. ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS Distribution and ecology.—This species is known only from the type locality, Huancabamba in the Departamento Pasco, Peru (Fig. 9). Huancabamba is at an elevation of 925 m on the Amazonian slopes of the Andes. Remarks.—When the type series was examined on 17 June 1969, the specimens were soft, but the color pattern was discernible. Of the five syntypes, BM 1947.2.13.56 is a female having a SVL of 46.4 mm; it shows the color pat- tern better than the other specimens and thus is designated as the lectotype. See remarks in following account. Hyla palaestes new species Holotype.—KU 163305, an adult male, one of a series collected at south edge of Tutumbaro (12°42' S, 73'54' W, 1840 m), 5.4 km (by road) SW Ayna, Provincia La Mar, Departamento Ayacucho, Peru, on 20 February 1975 by William E. Duellman and Linda Trueb. Paratypes.—KU 163306-14 collected with the holotype; KU 196844—53, and LSU 26002-15 from San José, Rio Santa Rosa, 1005 m, Provincia La Mar, Departamento Ayacucho, Peru. Referred specimens.—KU 164051 (tadpoles), KU 164092 (skeleton). Diagnosis.—A moderate-sized species in the Hyla pulchella Group characterized by: (1) snout rounded in dorsal view and bluntly rounded in profile; (2) skin on dorsum smooth; (3) width of disc on Finger III equals di- ameter of tympanum; (4) tarsal fold low; (5) dorsal color pattern consisting of irregular dark, transverse marks; (6) flanks uniform tan; (7) posterior surfaces of thighs uni- form tan. The only other species in the Hyla pulchella Group lack- ing large marks on the flanks is H. melanopleura, in which the flanks and posterior surfaces of the thighs are darker than the dorsum and the dorsal color pattern consists of a triangular interorbital mark and irregular dashes on the body. Also, in H. melanopleura, small, pale spots are present ventrolaterally on the flanks, and the width of the disc on Finger III is less than the diameter of the tympanum. Description.—n = 29 dd, 1 9. Body moderately robust; SVL in males to 50.4 mm, in females to 50.9 mm; head width about equal to head length, not as wide as body; head width 32.0-38.4% (x= 35.4) of SVL; head length 33.8-37.7% (x = 35.5) of SVL; snout rounded in dorsal view, bluntly rounded in profile; interorbital distance 103-159% (x= 128) of width of eyelid; diameter of eye 108-138% (X = 123) its distance from nostril; tympanum separated from eye by distance about equal to diameter of tympanum, diameter of tympanum 48.9-67.5% (x = 55.3) of diameter of eye. Breadth of forearm 41.9-51.3% (X= 46.2, n = 10) of forearm length in males, 32% in female; ulnar fold proximally and row of low, round tubercles distally on forearm; webbing, formula for fingers: I] (2—2)—(3-3) III 24%—(2-2) IV; width of disc on Finger III about equal to diameter of tym- panum; prepollex blunt, projecting at right angle in male, shorter in female; nuptial spines absent; nuptial excres- ANDEAN Groups OF HYLA 29 Decibels Kilohertz Seconds Fig. 17. Advertisement call of Hyla palaestes, KU Tape 1298, recorded at 14°C at Tutumbaro, Depto. Ayacucho, Peru. Spectrum, audiospectrograph, and waveform; arrow in audiospectrogram indicates point of spectrum. cences absent in breeding males. Hind limb slender; tibia length 49.9-58.8% (x= 53.2) of SVL; foot length 39.8-47.7% (X= 43.6) of SVL; inner tarsal fold low, straight; inner meta- tarsal tubercle ovoid, widest proximally, visible from above; webbing formula for toes: I (1-142)—(2-2%) II (I- 1*)—(2-2") IIT (1-14%4)—(2-2%) IV (2-2*)—(1-1*) V. Skin on dorsum and flanks smooth. Vomerine odontophores short, transverse, narrowly separated medially, posteromedial to rounded choanae, each bearing total 4-9 (x = 6.9) teeth. Coloration in preservative: Dorsum and flanks tan to brown with or without indistinct darker markings (most distinct in tan individuals) consisting of interorbital bar and four or five transverse marks on body and usually with narrow, transverse lines in interspaces between pri- mary marks; limbs with dark brown transverse bars—two or three on forearm, three or four (indistinct) on thigh, and four or five on shank; digits and webbing same color as dorsal ground color; anterior and posterior surfaces of thighs uniform pale brown; cloacal region dark brown with narrow, transverse creamy-white stripe above cloacal re- gion (most distinct in smaller individuals); margin of up- per lip creamy tan; venter uniform creamy tan. Juvenile (24.8 mm SVL) with narrow cream canthal and dorsolat- eral stripes bordered below by narrow, dark brown stripe; subadult male (32.5 mm SVL) with indistinct brown dor- solateral stripe. Coloration in life: The dorsum is brown to tan with faint markings; the flanks lack contrasting markings, and the venter is white (Fig. 3). KU 163305-13 from Tutumbaro, Depto. Ayacucho, Peru.—At night, dorsum of adults pale reddish brown to olive-tan with faintly darker transverse markings on body and limbs; groin and hidden surfaces of hind limbs pur- plish gray; webbing brown; venter creamy white with tint of olive-green on vocal sac; supracloacal and heel stripes creamy white. By day, some individuals changed to pale greenish tan with olive-green markings. Juveniles colored like adults by day except for a greenish-white labial stripe, pale yellow canthal and dorsolateral stripes bordered be- low by narrow reddish-brown line; groin and hidden sur- faces of hind limbs pale green. Iris pale creamy bronze (W. E. Duellman field notes, 20 February 1975). Measurements of holotype: SVL 46.3, tibia length 24.3, foot length 20.1, head length 15.4, head width 16.7, inter- orbital distance 6.2, internarial distance 3.2, width of eye- lid 3.9, eye-nostril distance 3.4, diameter of eye 4.7, diam- eter of tympanum 2.6. Life history.—The advertisement call consists of a se- ries of soft, low, trill-like notes of variable length. Analysis of one recording (KU Tape 1298) made at 14°C at the type locality reveals a call rate of nine calls per minute. Each call consists of four or five notes having durations of 0.02— 0.08 sec (x = 3.4) and 42-44 pulses per sec (x = 43). Only a single harmonic at 4.43-4.45 kHz (x = 4.44) is evident (Fig. 17). The first and last notes in a call are shorter than the intermediate ones; in one call, the lengths of sequential notes are 0.02, 0.03, 0.08, 0.03 sec, and in another they are 0.02, 0.04, 0.05, 0.02, 0.02 sec (Fig. 17). Sixteen tadpoles (Stages 37-43; KU 164051) are from the Rio Piene, Tutumbaro, Departamento Ayacucho, Peru. A specimen in Stage 37 (KU 164051) has the following mea- surements: total length 55.2, body length 18.5, tail muscle height at the base 6.2, tail muscle width at the base 5.6, maximum dorsal fin height 4.3 located 20.8 from body ter- minus, maximum ventral fin height 3.3 located 25.0 from body terminus, body width 13.0, body height 9.9, eye di- ameter 2.4, pupil diameter 0.7, interorbital distance 3.3, greatest and least nostril diameters 0.7 x 0.5, internarial distance 3.1, snout-nostril 3.3, snout-eye 6.0, snout-spiracle 9.8, eye-nostril 1.6, and transverse oral disc diameter 4.9. Other major characteristics include a ventral oral disc, mar- ginal papillae with median gap on upper labium, sinistral spiracle, dorsal eyes, dextral vent, and dorsal fin termi- nating at the dorsal extent of the tail-body junction. The LTRF is 2(2)/4(1) (Fig. 18). The outer wall of the spiracle is longer than the inner wall, and emerges from the ventrolateral surface of the body at midlength. The inner wall of the spiracular tube is attached to the body for its entire length; thus, none of the tube is free. The tube is directed posterodorsally and nar- rows to form a round, posteriorly directed aperture. The vent tube is dextral, and is positioned at the posterolateral surface of a large saccular structure that underlies the limb buds and is contiguous with the ventral fin. The vent ap- erture is directed posteroventrally. The eyes are positioned dorsally and directed dorsolaterally. The nares are oval, slightly medial to the center of the eyes, recessed, and weakly rimmed with the rim of the medial edge slightly 30 Fig. 18. 2mm Tadpole of Hyla palaestes, KU 164051, Stage 37, total length nN 5 raised. The snout is acutely rounded in dorsal view, and acutely rounded in profile, gradually sloping pos- terodorsally. The body is oval in dorsal view; in lateral view depressed dorsally and ventrally becoming deeper poste- riorly to bulging gut. The oral disc is slightly emarginate posterolaterally and is completely circumscribed by marginal papillae except for a median gap in the middle seventh of the upper la- bium; there are fewer marginal papillae medially on the lower labium. Submarginal papillae are scarce and present laterally in addition to flaps, some of which possess minute teeth. The LTRF of most individuals is 2(2)/4(1) with long, closely spaced teeth, but a few have gaps in A-1, P-2, or P- 3. The upper jaw sheath is moderately wide, widely arched, and moderately serrate. The lower jaw sheath is narrow, broadly V-shaped, and moderately serrate. The caudal musculature is highest at the tail-body junc- tion and is the same height to the midlength, where it nar- rows gradually to a long, narrow tail terminus. In lateral view, the dorsal fin terminates at the dorsal tail-body junc- tion, is highest at about two-thirds length of tail, and is equal to musculature height at midlength of the tail (taller posteriorly, shorter anteriorly). The ventral fin is shorter than the dorsal fin at the midportion of tail (equal for pos- terior fourth, taller for anterior fourth). The ventral fin is shorter than the musculature at midlength (taller posteri- orly, shorter anteriorly). The greatest height of the tail is at about midlength. The tail terminates as a long, narrow, pointed tip. SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS In preservative, the dorsum of the body is uniformly pigmented with brown chromatophores. The venter has little, if any pigment laterally, and the intestine is clearly visible. Dorsal pigmentation continues posteriorly onto dorsum of anterior half of caudal musculature. The fins are transparent, and the musculature is opaque yellowish brown; both have scattered discrete brown blotches; the blotches interconnect along the midheight of the muscu- lature and are limited to the anterior part of the dorsal fin. In life, the dorsum is olive-tan with olive-green mottling, flanks greenish silver, and belly silver-white. The caudal musculature is grayish tan with brown spots which are also on the transparent fins; the iris is creamy bronze, and the bones are green (W. E. Duellman field notes 20, Febru- ary 1975). Distribution and ecology.—This species is known defi- nitely only from two localities—San José on the Rio Santa Rosa 1005 m and Tutumbaro on the Rio Piene 1840 m—in the drainage of the Rio Apurimac on the eastern face of the Andes in central Peru (Fig. 9). The localities are about 12 km (airline) apart. Males were calling from dense vegetation within 0.5 m of a low-gradient stream after a rain of 12 mm on the night of 20 February 1975. The site is at an elevation of 1840 m at the south edge of the village of Tutumbaro in the valley of the Rio Piene, a tributary of Rio Apurimac. Tutumbaro is 5 km 5 of Ayna on the road to Ayacucho. The site is in dis- turbed cloud forest with bamboo, ferns, and elephant ear plants but no tree ferns and few bromeliads. Other anurans found at the same site include Bufo veraguensis, Cochranella phenax, Hyalinobatrachium bergeri, Gastrotheca testudinea, and Ayla armata. Etymology.—The specific name, palaestes, is a Latin noun meaning wrestler; the name is used in reference to the robust arms of adult males. Remarks.—This species is similar in size and colora- tion to Hyla melanopleura, which also lacks bold markings on the flanks, but has small pale spots ventrolaterally (ab- sent in H. palaestes). However, in H. melanopleura, the flanks are much darker than the dorsum, whereas they are the same color as the dorsum in H. palaestes. The relative size of the disc on Finger III (smaller than tympanum in H. melanopleura and equal to diameter of tympanum in H. palaestes) may be an artifact of preservation. HYLA BOGOTENSIS GROUP Definition.—(1) Moderate-sized, stream-breeding frogs with males attaining SVLs of 45 mm and females 53 mm; (2) round mental gland in males; (3) preorbital ridge and lateral folds absent; (4) dorsum green or pale brown with or without dark flecks; (5) no conspicuous marks on flanks or hidden surfaces of thighs; (6) venter uniformly pale; (7) digits bearing small discs; (8) nuptial excrescences and projecting prepollical spines absent in males; (9) forearms of adult males not hypertrophied; (10) axillary membrane absent; (11) toes about three-fourths webbed; (12) nasals small, widely separated medially; (13) frontoparietal fon- ANDEAN Groups OF HYLA 3] tanelle large; (14) sphenethmoid broad; septum nasi not ossified anteriorly; (15) quadratojugal usually articulating with maxillary; (16) dentigerous processes of vomers trans- verse, between posterior margins of choanae, narrowly separated medially; (17) tadpoles having long, muscular tails, low fins, and pointed terminus; (18) spiracle attached to body wall for its full length or free distally; (19) larval mouth large, anteroventral, complete rows of marginal papillae, no plate or narrow plate below serrations on up- per jaw sheath, and a LTRF usually of 4-5/6-10 (maxi- mum of 14/15 in Hyla jahni); (20) known advertisement calls consisting of series of short, loud peeps (trill in H. jahni). Content.—Fourteen species: Hyla alytolylax Duellman, 1972; H. bogotensis (Peters, 1882); H. callipeza Duellman, 1989; H. colymba Dunn, 1931 (H. alvaradoi Taylor, 1952, is a synonym); H. denticulenta Duellman, 1972; H. jahni Rivero, 1961; H. lascinia Rivero, 1969; H. lynchi Ruiz-Carranza and Ardila-Robayo, 1991; H. palmeri Boulenger, 1908; H. phyllognatha Melin, 1941; H. piceigularis Ruiz-Carranza and Lynch, 1982; H. platydactyla Boulenger, 1905 (H. paramica Rivero, 1961, is a synonym); H. simmonsi Duellman, 1989; H. torrenticola Duellman and Altig, 1978. Distribution.—Most species have allopatric distribu- tions at elevations of 800-2600 m in cloud forests on the eastern and western flanks of the Andes in Colombia (also Cordillera Central) and Ecuador, and two species (Hyla jahni and H. platydactyla) occur sympatrically in the Mérida Andes in Venezuela (La Marca, 1985). Hyla bogotensis oc- curs in subparamo and paramo to elevations of 3600 m (Ruiz-Carranza and Lynch, 1982). On the Amazonian slopes of the Andes, H. phyllognatha ranges from southern Colombia to central Peru; it is the only species known to occur south of the Huancabamba Depression. Hyla palmeri is known from elevations of 100-1600 m in the Chocoan lowlands and on the western slopes of the Cordillera Oc- cidental in Colombia and Ecuador (Ruiz-Carranza et al., 1996) and from one locality in Central Panama, where it is sympatric with H. colymba, which has a discontinuous range from central Costa Rica to extreme eastern Panama (Myers and Duellman, 1982). Remarks.—The Hyla bogotensis group was reviewed by Duellman (1972) and redefined by Duellman (1989); sub- sequently, H. lynchi was described by Ruiz-Carranza and Ardila-Robayo (1991). HYLA CHLOROSTEA GROUP Definition.— (1) Moderately small frog with only known male having a SVL of 39.7 mm; (2) round mental gland absent; (3) preorbital ridge and lateral folds present; (4) dorsum pale brown with darker brown markings; (5) no conspicuous marks on flanks or hidden surfaces of thighs; (6) venter uniformly pale; (7) digits bearing mod- erately large discs; (8, 9) unknown; (10) axillary membrane absent; (11) toes about three-fourths webbed; (12-15) un- known, (16) dentigerous processes of vomers short, slightly curved posteromedially, posterior to choanae; (17-20) un- known. Content.—One species: Hyla chlorostea Reynolds and Foster, 1992. Distribution.—Known only from Parjacti at an eleva- tion of 2044 m in Departamento Cochabamba, Bolivia. Remarks.—This highly distinctive species is known from only a subadult male. De la Riva and Duellman in- dependently examined the holotype and concur with Reynolds and Foster (1992) that this frog cannot be as- signed to any known group of Hyla. HYLA COLUMBIANA GROUP Definition.— (1) Small, pond-breeding frogs with males attaining SVLs of 26 mm and females 32 mm; (2) round mental gland absent; (3) preorbital ridge and lateral folds absent; (4) dorsum tan or brown with or without darker markings and pale dorsolateral stripes; (5) axilla and groin yellow, orange, or blue; no bold marks on hidden surfaces of thighs; (6) venter pale with darker spots; (7) digits bear- ing small discs; (8) nuptial excrescences and projecting prepollical spines absent in males; (9) forearms of adult males not hypertrophied; (10) axillary membrane present; (11) toes two-thirds to three-fourths webbed; (12) nasals small, widely separated medially; (13) frontoparietal fon- tanelle short, narrow; (14) sphenethmoid broad; septum nasi not ossified anteriorly; (15) quadratojugal absent; (16) dentigerous processes of vomers inclined posteromedially between choanae, moderately to widely separated medi- ally; (17) tadpoles having moderately short tails, with high fins, and xiphicercal terminus; (18) spiracle attached to body wall for its entire length; (19) larval mouth small, terminal, directed anteroventrally, with broad median gap on upper labium, no plate below serrations on upper jaw sheath, and a LTRF usually of 1/2; (20) advertisement calls consisting of short, moderately pitched notes followed by shorter secondary notes Content.—Three species: Hyla carnifex Duellman, 1969 (H. bogerti Cochran and Goin, 1970, and H. charlesbogerti Goin, 1970, are synonyms); H. columbiana Boettger, 1892 (H. variabilis Boulenger, 1896, is a synonym); H. praestans Duellman and Trueb, 1983. Distribution.—Allopatric distributions at elevations of 975-2580 m on the Pacific slopes of the Cordillera Occi- dental in Ecuador and Colombia (H. carnifex), the upper Rio Cauca (H. columbiana) and Magdalena (H. praestans) valleys, and northeastern slopes of the Cordillera Oriental (H. garagoensis and H. stingi) in Colombia. Remarks.—The group was defined and reviewed by Duellman and Trueb (1983), who hypothesized that on the basis of larval characters and a diploid number of 30 chro- mosomes the Hyla columbiana group was the sister group of the H. minuta, marmorata, parviceps, leucophyllata, and microcephala groups inhabiting tropical lowlands in Middle and South America. Kaplan (1991) noted in his descrip- tion of H. garagoensis that the larvae of H. praestans are unknown and the adults of that species closely resemble those of H. garagoensis; subsequently, Kaplan (1994) de- os) scribed H. stingi, which he considered to be an indepen- dent lineage between the H. columbiana and H. minuta groups. HYLA GARAGOENSIS GROUP Definition.—(1) Small, pond-breeding frogs with males attaining SVLs of 30 mm and females 34 mm; (2) round mental gland absent; (3) preorbital ridge and lateral folds absent; (4) dorsum down with darker or paler marks and pale dorsolateral stripes; (5) groin and hidden surfaces of thighs yellow or orange; (6) venter pale with or without dark flecks on chest; (7) digits bearing small discs; (8) nup- tial excrescences and projecting prepollical spines absent in males; (9) forearms of adult males not hypertrophied; (10) axillary membrane brief or absent; (11) toes one-half to two-thirds webbed; (12) nasals small, widely separated medially; (13) frontoparietal fontanelle short and poste- rior or long and slender; (14) sphenethmoid broad; sep- tum nasi not ossified anteriorly; (15) quadratojugal absent; (16) dentigerous processes of vomers small, inclined posteromedially between choanae, moderately separated medially; (17) tadpoles having moderately short tails, with high fins and xiphicercal terminus; (18) unknown; (19) lar- val mouth small, terminal, directed anteroventrally, with broad median gap on upper labium, no plate below serra- tions on upper jaw sheath, and a LTRF of 0/0; (20) adver- tisement calls not reported. Content.—Three species: Hyla garagoensis Kaplan, 1991; H. padreluna Kaplan and Ruiz, 1997; H. virolinensis Kaplan and Ruiz, 1997. Distribution —Allopatric distributions at elevations of 1750-2200 m on the eastern and western slopes of the Cor- dillera Oriental in central Colombia. Remarks.—The group was defined by Kaplan and Ruiz (1997), who assumed that the species have 2N = 30 chro- mosomes; in their phylogenetic analysis, the Hyla garagoensis Group formed a trichotomy with the Hyla marmorata Group and the clade containing the H. parviceps, leucophyllata, and microcephala groups. We have examined no specimens of this group and have relied on data pub- lished by Kaplan (1991) and Kaplan and Ruiz (1997), who noted that males of all three species were calling but ne- glected to describe the calls. HYLA LABIALIS GROUP Definition.—(1) Moderate-sized, pond- and slow- stream-breeding frogs with males attaining SVLs of 50 mm and females 55 mm; (2) round mental gland absent; (3) preorbital ridge and lateral folds absent; (4) dorsum green with or without diffuse black spots; (5) groin and hidden surfaces of thighs blue or yellow, mottled with black; (6) venter uniformly pale; (7) digits bearing small discs; (8) nuptial excrescences and projecting prepollical spines ab- sent in males; (9) forearms of adult males not hypertro- phied; (10) axillary membrane present or absent; (11) toes one-half to two-thirds webbed; (12) nasals small, slender, widely separated medially; (13) frontoparietal fontanelle 2 ScrENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS large; (14) sphenethmoid broad; septum nasi ossified an- teriorly; (15) quadratojugal absent; (16) dentigerous pro- cesses of vomers inclined posteromedially between choa- nae, narrowly separated medially; (17) tadpoles having moderately short tails, high fins, and pointed terminus; (18) spiracular tube attached to body wall for its entire length; (19) larval mouth small, directed anteroventrally with broad median gap on upper lip, no plate below ser- rations on upper jaw sheath, and a LTRF of 1/2; (20) ad- vertisement calls consisting of short series of guttural notes. Content.—Three species: Hyla labialis Peters, 1863 (H. vilsoniana Cope, 1882, H. creolica Werner, 1899, H. servalina Werner, 1899, H. gularis Werner, 1916, and H. vilsoniana krausi Hellmich, 1940, are synonyms); H. meridensis Rivero, 1961; H. pelidna Duellman, 1989. Distribution.—Allopatric distributions in upper cloud forest, subparamo, and paramo at elevations of 2180-3400 m in the Cordillera Oriental in Colombia and the Mérida Andes in Venezuela. Remarks.—The Hyla labialis group was defined by Duellman and Trueb (1983) and Duellman (1989). A dip- loid number of 30 chromosomes was reported for H. labialis by Bogart (1973). HYLA LARINOPYGION GROUP Definition.—(1) Large, stream-breeding frogs with males attaining SVLs of 68 mm and females 77 mm; (2) round mental gland absent; (3) preorbital ridge and lat- eral folds absent; (4) dorsum brown, gray, or black with or without pale markings; (5) groin and hidden surfaces of thighs black or marked with black bars or mottling; (6) venter black with or without pale spots or mottling; (7) digits bearing large discs; (8) nuptial excrescences absent in males; enlarged, nonprojecting prepollex present; (9) forearms of adult males not hypertrophied; (10) axillary membrane absent; (11) toes one-third to two-thirds webbed; (12) nasals long, slender, narrowly separated medially; (13) frontoparietal fontanelle small; (14) sphenethmoid broad; septum nasi ossified anteriorly; (15) quadratojugal articulating with maxillary; (16) dentiger- ous processes of vomers long, transverse, posterior to choa- nae, abutting medially; (17) tadpoles having long, muscu- lar tails, low fins, and pointed terminus; (18) distal one- third of spiracular tube free from body wall; (19) larval mouth large, ventral, with complete rows of marginal pa- pillae, no plate below serrations on upper jaw sheath, and a LTRE of 4-8/6-10; (20) advertisement calls consisting of series of whistles or ticklike notes. Content.—Nine species: Hyla caucana Ardila-Robayo, Ruiz-Carranza, and Roa-Trujillo, 1993; H. larinopygion Duellman, 1973; H. lindae Duellman and Altig, 1978; H. pacha Duellman and Hillis, 1990; H. pantosticta Duellman and Berger, 1982; H. psarolaima Duellman and Hillis, 1990; H. ptychodactyla Duellman and Hillis, 1990; H. sarampiona Ruiz-Carranza and Lynch, 1982; H. staufferorum Duellman and Coloma, 1993. ANDEAN Groups OF HYLA 33 Table 5. Character states of seven groups of Andean Hyla. Characters Group A a? B e D E F : G Ee : I J K L M N Outgroup 0 0 0 0 0 0 0 0 0) 0 0 0 0 armata 0) 1 0 0 1 1 1 2 1 1 1 1 0 bogotensis 0 1 0 0 1 1 0 0 1 1 1 1 0 0 columbiana 1 0 0 1 0 1 0 0 3 0 0 2 (0) 1 garagoensis 1 0 0 0 0 1 0 0 3 0 0 2 0 1 labialis 1 0 0 0 0 1 0 0 0 0 0 2 0 1 Distribution.—Most species are inhabitants of the im- mediate vicinity of cascading streams in cloud forests at elevations of 1900-2700 m on the slopes of the Cordillera Occidental and Cordiliera Oriental in Ecuador and south- ern Colombia, but two species occur in the Cordillera Cen- tral in Colombia—H. caucana at 2400 m and H. larinopygion at elevations of 2200-3100 m (Ardila-Robayo et al., 1993). Most of the species are known from only one or two lo- calities, but H. larinopygion is widely distributed in the Cordillera Central in Colombia, and H. psarolaima has an elevational range of 1950-2660 m on the Amazonian slopes of the Cordillera Oriental in southern Colombia and Ec- uador where it occurs sympatrically with H. lindae and H. pacha (Duellman and Hillis, 1990). Remarks.—The group was defined by Duellman and Hillis (1990), who provided a key to the species and a cla- distic analysis of allozymes of five species. Subsequently, additional species were described by Ardila-Robayo et al. (1993) and Duellman and Coloma (1993). PHYLOGENETIC RELATIONSHIPS Hypothesizing phylogenetic relationships among spe- cies in the large and morphologically diverse genus Hyla is hampered by the large number of species (294; updated from Duellman, 1993), inadequate knowledge about many species, and absence of reliable hypotheses of higher-level relationships. For example, it is unlikely that all groups of Andean Hyla are closely related. Thus, the results of our analyses are used more to provide a framework for dis- cussing the distribution of characters rather than hypoth- esizing phylogenetic relationships. Herein we employ the methodology of phylogenetic systematics (Wiley, 1981) and the principles of parsimony and outgroup comparison (Farris, 1982; Watrous and Wheeler, 1981). Ascertaining polarity of characters is problematic. Nev- ertheless, certain characters seem to be plesiomorphic within the genus. These are: (1) quadratojugal present and articulating with maxilla; (2) dentigerous processes of vomers inclined posteromedially and separated from one another medially; (3) prepollex not modified into a pro- jecting spine; (4) axillary membrane absent; (5) mental gland absent; (6) nuptial excrescences present; (7) forearms of adult males not hypertrophied; (8) clusters of spines on thumb and humerus absent; (9) tadpoles with high caudal fins and pointed tail and caudal musculature not robust; (10) larval oral disc directed anteroventrally; (11) larval oral disc having median gap anteriorly in marginal papillae; (12) LTRF 2/3; (13) larval mouth lacking plate below ser- rations on upper jaw sheath; (14) diploid chromosome number 24. These characters are present in many neotropical species of Hyla, including such diverse spe- cies as H. loquax, punctata, and smithii, as well as most spe- cies of such diverse genera as Pseudacris, Scinax, and Smilisca (Duellman, 1970; Duellman and Wiens, 1992; Gaudin, 1974). Thus, these taxa can be used collectively as an outgroup for comparisons with the groups of Andean Ayla. RELATIONSHIPS AMONG ANDEAN GROUPS OF HYLA Herein we make no attempt to analyze the Andean Hyla at the species level. Instead, each of the groups previously defined is considered as an operational taxonomic unit (OTU). Insofar as is known, the characters listed above are consistent within six of the groups; thus, each can be treated as a terminal taxon. Apomorphies support the monophyly of five of the groups. The group represented solely by Hyla chlorostea is not included in the analysis because only three of the 14 characters could be coded for that species, which differs from all other Hyla in having two derived charac- ter states—a conspicuous preorbital ridge and heavy lat- eral folds. The following characters were coded for the six groups of Andean Hyla (Table 5). These characters and the polar- ity (0 = primitive; 1, 2, and 3 derived) of their states are: A. Quadratojugal—tThe quadratojugal usually articu- lates anteriorly with the posterior tip of the maxilla and posteriorly with the pars articularis of the palatoquadrate; thus it forms the posterior part of the maxillary arcade. In some species (e.g., Hyla leucophyllata and H. parviceps), the quadratojugal is reduced in size and does not articulate with the maxilla; in other species (e.g., H. carnifex and H. labialis), the element is absent. Apparently, progressive re- duction terminates in the absence of the element; thus, the states are coded: 0 = present, articulating with maxilla; 1 = not articulating with maxilla. B. Dentigerous processes of vomers.—Dentigerous processes of the vomers (vomerine odontophores) usually are short, posteromedially inclined, and separated medi- ws) ally by a distance equal to about half the breadth of a pro- cess. A derived condition is the lengthening of the pro- cesses in a transverse plane and consequent reduction of the distance between the processes. The states are coded: 0 = detigerous processes short, posteromedially inclined, broadly separated medially; 1 = dentigerous processes long, transverse, narrowly separated or abutting medially. C. Prepollex.—tIn most hylids, the prepollex is some- what enlarged and provides support for the nuptial ex- crescences on the thumb. In some species (e.g., Hyla lindae), the prepollex is greatly enlarged but does not have a pro- jecting spine, whereas in other taxa (e.g., H. pulchella and H. boans groups) the prepollex in males has a projecting spine. The transformation series is ordered: 0 = prepollex not greatly enlarged; 1 = prepollex greatly enlarged but not having a projecting spine in males; 2 = prepollex greatly enlarged with a projecting spine in males. D. Axillary membrane.—Usually, no web of skin (axil- lary membrane or patagium) is present between the pos- terior edge of the upper arm and the flank, but this mem- brane is present in many species (e.g., members of the Hyla columbiana and H. leucophyllata groups). The states are coded: 0 = axillary membrane absent; 1 = extensive axil- lary membrane present. E. Mental gland.—Among Andean hylids, a round, glandular structure (mental gland) is present on the chin only in males of the Hyla bogotensis group. The states are coded: 0 = mental gland absent; 1 = mental gland present. FE. Nuptial excrescences.—Keratinized nuptial excres- cences normally are present on the medial surface of the thumb in pond-breeding hylids and in some species that breed in streams (e.g., Hyla bistincta group). The absence of nuptial excrescences is considered to be a derived con- dition; thus the states are coded: 0 = nuptial excrescences present; 1 = nuptial excrescences absent. G. Hypertrophy of forelimbs.—Usually, the arms of adult males are not noticeably more massive than those of females, but the arms are greatly hypertrophied in adult males of some species (e.g., members of the Hyla armata and H. pulchella groups). The states are coded: 0 = fore- arms not hypertrophied; 1 = forearms hypertrophied. H. Spines on thumb and humerus.—Among hylids, keratin-covered bony spines are present on the thumb and proximal ventral surface of the humerus only in Hyla armata. The states are coded: 0 = spines absent; 1 = spines present. I. Larval tail—The generalized tadpole of Hyla and most other genera within the family has a tail no more than twice the length of the body; the caudal fins are much higher than the caudal musculature at midlength of the tail, and the tail terminates ina pointed tip. In many groups of stream-breeding hylids (e.g., H. bogotensis and H. larinopygion groups), the tail is more than twice the length of the body, the fins are only slightly higher or equal to the height of the caudal musculature at midlength of the tail, and the tip of the tail is pointed. Hyla armata has a tail like 4 SCIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS that of members of the H. bogotensis and H. larinopygion groups, but the tip of the tail is bluntly rounded. Some pond-breeding Hyla (e.g., H. columbiana and H. leucophyllata groups) have tadpoles with high caudal fins and the tail terminating a long, slender tip (xiphicercal condition). The states are coded: 0 = short tail with high fins and pointed tip; 1 = long tail with low fins and pointed tip; 2 = long tail with low fins and bluntly rounded tip; 3 = short xiphicercal tail. The transformation is 3 <— 0 > 1 > 2. J. Orientation of larval oral disc.—In most pond-breed- ing hylids, the oral disc is directed anteroventrally, whereas in diverse groups of stream-breeding Hyla (e.g., H. bogotensis, claresignata, laririopygion, and rivularis groups), the oral disc is enlarged and directed ventrally. The states are: 0 = disc directed anteroventrally; 1 = disc enlarged, directed ventrally. K. Marginal papillae on larval oral disc.—The plesiomorphic condition is the existence of at least one row of marginal papillae with a broad gap in the papillae on the anterior labium of the oral disc. In most species hav- ing an enlarged, ventral oral disc, the marginal papillae extend across the anterior labium. The states are coded: 0 = gap in marginal papillae on upper labium; 1 = marginal papillae extending completely across upper labium. L. Labial tooth row formula (LTRF).—In most pond- breeding hylids, the standard tooth-row formula in tad- poles is 2/3, but it is not uncommon to have formulae of 3/4. Insome stream-breeding Hyla with large, ventral oral discs (e.g., H. bogotensis and H. larinopygion groups) the LTRF is much larger, whereas in some pond-breeding Hyla (e.g., H. columbiana and H. garagoensis groups), the LTRF is reduced to 1/2 or even 0/0. The states are coded: 0 = LTRF 2/3 or 3/4; 1 = LTRF 4/6 or greater; 2 = LTRF 1/2; 3 = LTRF 0/0. The transformation is 3 — 2<-0- 1. M. Larval upper jaw sheath.—Normally, the serrate edge forms the posterior margin of the upper jaw sheath, but in Hyla armata, the serrated edge is bordered posteri- orly by a broad, smooth plate (shelf of Cadle and Altig, 1991). The states are coded: 0 = broad plate posterior to serrations absent; 1 = broad plate posterior to serrations present. N. Chromosome number.—The normal diploid num- ber of chromosomes is 24 in frogs of the subfamily Hylinae. Based on the conclusions of Bogart (1973) and Morescalchi (1973) that 30 chromosomes in some groups of Hyla evolved by centric fission, we follow Duellman and Trueb (1983) in considering 30 chromosomes to be a uniquely derived condition in Hyla. This diploid number is known in only two groups (H. columbiana and H. labialis) in the Andes, as well as in several groups (e.g., H. leucophyllata and H. parviceps groups) in the neotropical lowlands. The states are coded: 0 = 24 chromosomes; 1 = 30 chromosomes. An analysis of the seven groups using the 14 characters with states unordered resulted ina tree having a length of 20 steps and a consistency index of 0.90 (Fig. 19). An analy- sis with the states polarized as noted above resulted in ANDEAN Groups OF HYLA 35 G (01) H (0+ 1) I (12) M (0+1) C(0>1) J (0>1) K (01) L (01) I (03) C (02) A) G (0+ 1) L (02) N (01) B (0O>1) I (0>1) ws = Apomorphies [4 Convergences Fig. 19. Cladogram showing distribution of, and shifts in, character states within Andean groups of Hyla. The tree was generated by PAUP analysis using DELTRAN character optimization and has a length of 20 steps and a consis- tency index of 0.90. See text for states of characters A-N. three equally parsimonious trees, each 21 steps long and having consistency indices of 0.857; the topology of one of these trees is like the one resulting from unordered char- acter states. The differences among the three trees result- ing from the analysis of polarized states are in the arrange- ment of the H. bogotensis, larinopygion, and armata groups; the only character that fluctuates in these topologies is Character C, enlargement of the prepollex and modifica- tion of the prepollex into a projecting spine in males. All four trees are consistent in sharing Character F, absence of nuptial excrescences, thereby distinguishing the Andean groups of Hyla from the outgroup and three characters (A, quadratojugal absent; L, labial tooth rows reduced; and N, 30 chromosomes) shared by the clade containing the H. labialis, columbiana, and garagoensis groups. The former is not supported by apomorphies, whereas the clade contain- ing the H. columbiana and H. garagoensis groups is sup- ported by one apomorphy and (I, xiphicercal tail in lar- vae). Within this clade, the H. columbiana Group has one apomorphy (D, axillary membrane present), and the H. garagoensis Groups has one apomorphy (L, LTRF reduced to 0/0). The other major clade containing four groups is consistently supported by two characters (B, dentigerous processes of vomers long, transverse, and narrowly sepa- rated medially; and I, larval tail with low fins and pointed tip). Within this clade, the H. pulchella Group is supported by one apomorphy (C, prepollex enlarged with a project- ing spine in males) and one character transformation (G, hypertrophied forelimbs in males) also present in the H. armata Group. The clade containing the other three groups (H. bogotensis, larinopygion, and armata) is supported by three synapomorphies—all characters of the larval oral disc (J, disc enlarged and directed ventrally; K, marginal pa- pillae complete across upper labium; and L, increased num- ber of labial tooth rows). Of these three groups, the H. bogotensis Group has a single apomorphy (E, mental gland present). The H. larinopygion Group is not supported by apomorphies, whereas H. armata is distinguished by three apomorphies (H, spines on thumb and humerus; I, larval tail with a rounded tip; and M, broad plate posterior to Fig. 20. Hypothesized phylogenetic relationships among Hyla having a diploid number of 30 chromosomes; based on Duellman and Trueb (1983), as modified by Kaplan (1994), Kaplan and Ruiz (1997), and data presented herein. Numbered synapomorphies are: | = 30 chromosomes, 2 = reduced quadratojugal, 3 = LTRF 1/2, 4 = nuptial excrescences absent, 5 = larval tail xiphicercal, 6 = larval mouth anterior, 7 = LTRF 0/1. 8 = LTRF 0/0, 9 = one ventral row of small labial papillae in larvae (two rows in some H. padreluna), 10 = extensive axil- lary membrane, 11 = longitudinal stripes on hind limbs of larvae, 12 = one ven- tral row of large papillae in larvae, 13 = larval body violin-shaped, 14 = larval body depressed, 15 = labial papillae absent in larvae. serrations on upper jaw sheath of larvae); H. armata also has one character transformation (G, hypertrophied fore- limbs in adult males) that also is present in the H. pulchella Group. The Hyla columbiana and H. labialis groups (and presum- ably the H. garagoensis Group) differ from other Andean Hyla by having 30 chromosomes; this seems to relate them to several groups of species in the lowlands. This relation- ship is supported by the reduced features on the oral discs of the tadpoles of the H. columbiana and H. labialis groups and further reduction in the H. garagoensis Group and the lowland groups. On the assumption that a diploid num- ber of 30 chromosomes is a synapomorphy for a major clade of neotropical Hyla, Duellman and Trueb (1983) pro- vided a cladistic analysis which was based primarily on larval characters (Fig. 20). If, as they proposed, species of Hyla having 30 chromosomes represent a single phyletic lineage, the H. columbiana, garagoensis, and labialis groups are not related to other Andean Hyla. However, chromo- some number must be used with caution, because the karyotypes are not known for all of the species. Among Andean Hyla, 30 chromosomes have been reported only for H. labials (Bogart, 1973) and H. carnifex (Duellman and Trueb, 1983). Preparations of tissues of H. armata and H. callipleura from Peru by David C. Cannatella revealed a we) diploid number of 24 chromosomes in these species. The chromosome number is not known for species in the H. bogotensis and H. larinopygion groups. There is no compelling evidence that the other groups of Andean hylids are closely related. In the present analy- sis, they are united by two characters (B, dentigerous pro- cesses of vomers long, transverse, and narrowly separated medially; and I, larval tail with low fins and pointed tip). The latter character, together with Characters J (oral disc enlarged and directed ventrally), K (marginal papillae com- plete across upper labium), and L (increased LTRF), are common modifications in tadpoles among diverse groups of hylids inhabiting lotic environments. All species in sev- eral groups of stream-breeding hylids in Mexico and Cen- tral America have long, muscular tails with low fins with pointed tips and enlarged, ventral mouths with one or more rows of marginal papillae complete across the up- per labium; the species of Hyla have LTRFs of 2/3 to 7/11 (Duellman, 1970), whereas those of Plectrohyla have a LTRF of 2/3 (Duellman and Campbell, 1992) and those of Ptychohyla have LTRFs of 2/5 to 6/9 (Campbell and Smith, 1992). Aside from Andean frogs in the H. armata, bogotensis, and larinopygion groups, few tadpoles adapted for life in torrential streams are known from South America. All of them have long muscular tails with low fins, large mouths directed ventrally, marginal papillae complete across the upper labium, and increased numbers of rows of labial teeth. These species are H. claresignata with a LTRF of 9/14 (Lutz and Orton, 1946), and H. clepsydra with a LTRF of 7/ 13 (Bokermann, 1972) in southeastern Brazil, and Hyla aromatica and H. inparquesi with LTRFs of 13/18 and 14/ 21, respectively, in the Guiana Highlands of southern Ven- ezuela (Ayarzagiiena and Senaris, 1993). It is unlikely that these larval characters are indicative of relationships; in- stead, they seem to represent convergent adaptations to lotic environments. Similar adaptive types of tadpoles are known in Australo-Papuan pelodryadine hylids of the genera Litoria and Nyctimystes, African Heleophryne and ranids of the genera Conraua, Petropedetes, and Trichobatrachus, and Asiatic pelobatids of the genus Scutiger and ranids of the genus Rana (Altig and Johnston, 1989; Duellman and Trueb, 1986). Given the independent evolution of the same stream adaptations in tadpoles of unrelated species groups, gen- era, and even families of anurans, it is unlikely that the Hyla armata, bogotensis, larinopygion, and pulchella groups in the Andes had an immediate common ancestor. Of these groups, the H. bogotensis Group is unique in the presence of a mental gland in males, but a similar reniform or oval gland occurs in the lowland H. granosa, which breeds in ponds and has typical lentic tadpoles with a LTRF of 2/3 (Hoogmoed, 1979); thus, H. granosa may be a lowland rela- tive of the Andean H. bogotensis Group, but detailed study of the glands must be undertaken to ascertain their ho- mology. Aside from the general stream adaptations of the tadpoles and the long, transverse dentigerous processes of the vomers narrowly separated or abutting medially 6 ScrENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS (shared with the H. larinopygion Group), there are no char- acters to unite members of this group with other Andean Hyla. The latter also applies to the H. larinopygion Group, males of which have an enlarged, nonspinous prepollex and no nuptial excrescences. Similarly, Mexican frogs of the H. bistincta Group also have enlarged, nonspinous prepollices, but these frogs have keratinized nuptial ex- crescences (Duellman, 1970; Duellman and Campbell, 1992). Hyla armata also has an enlarged nonspinous prepollex, but the prepollex (and humerus) bears kerati- nized spines in adult males, and the larvae are unique among Andean Hyla in having a bluntly rounded tail and a broad plate posterior to the serrations on the upper jaw sheath. Although no sister groups are apparent, it is pos- sible that the H. armata and H. larinopygion groups evolved independently from separate lineages in the lowlands. However, Hyla armata shares Character G (hypertro- phied forelimbs) in males with the H. pulchella and H. circumdata groups. (See following section.) The extent of the hypertrophied forelimbs is much greater in H. armata than in the other taxa. Hypertrophied forelimbs also oc- cur in adult males of the Middle American H. bistincta Group and Plectrohyla; in most species of Plectrohyla, the cristae on the humerus are expanded, although not to the extent that they are in H. armata (Duellman, 1970; Duellman and Campbell, 1992). Until more data, especially on tadpoles and skeletal fea- tures, are available, it is reasonable to assume that the Hyla armata, bogotensis, and larinopygion groups represent mono- phyletic lineages. Hyla armata may have been derived from a lowland lineage or be a highly derived species related to the H. pulchella Group. The H. bogotensis Group might be the sister group of Hyla granosa, with which it shares a mental gland in males. There is no obvious sister group to the H. larinopygion Group. RELATIONSHIPS OF THE HYLA PULCHELLA GROUP Two derived characters of adult males (C, prepollex en- larged with a distal spine projecting medially; G, hyper- trophied forelimbs) of the Hyla pulchella Group are shared with members of the H. circumdata Group in southeastern Brazil. This group contains 16 species: H. alvarengai Bokermann, 1956; H. astartea Bokermann, 1967; H. biobeba Bokermann and Sazima, 1973; H. carvalhoi Peixoto, 1981; H. circumdata Cope, 1867; H. eee Peixoto and Cruz, 1992; Hyla hylax Heyer, 1985; H. ibitiguara Cardoso, 1983; H. ibitipoca Caramaschi and Feio, 1990; H. izecksohni Jim and Caramaschi, 1979; H. langei Bokermann, 1966; H. martinsi Bokermann, 1964; H. nanuzae Bokermann and Sazima, 1973; H. pseudopseudis Miranda-Ribeiro, 1937; H. saxicola Bokermann, 1964 (considered to be a subspecies of H. pseudopseudis by Lutz, 1973, but recognized as a distinct species by Pombal and Caramaschi, 1995); H. sazimai Cardoso and Andrade, 1982. These frogs inhabit the inte- rior highlands to elevations in excess of 2000 m in the states of Goids and Minas Gerais and the coastal ranges from Espirito Santo to Rio Grande do Sul in southeastern Bra- ANDEAN Groups OF HYLA 37 zil, where they breed in streams (Bokermann, 1964; Bokermann and Sazima, 1973; Cardoso, 1983; Cardoso and Andrade, 1982; Heyer, 1985; Lutz, 1973; Peixoto and Cruz, 1992). Most of the species have allopatric distributions, but three species (H. astartea, circumdata and hylax) are sympa- tric at Boracéia at 900 m in the Serra do Mar, Sao Paulo, Brazil, where a member of the H. pulchella group (H. prasina) also occurs (Heyer et al., 1990). Frogs of the Hyla circumdata Group are highly variable in size (adult males from 36 mm in H. ibitipoca and H. sazimai to 76 mm in H. alvarengai and H. biobeba) and are tan to reddish brown dorsally with darker brown spots or transverse bars on the dorsum; the flanks and hidden sur- faces of the thighs are boldly marked with dark spots or vertical bars. (See Heyer et al., 1990, and Lutz, 1973, for color photographs of H. circumdata, hylax, and saxicola). The dentigerous processes of the vomers are relatively small, arched, or oblique between the posterior margins of the choanae. Adult males have distinctly hypertrophied fore- limbs and projecting prepollices. In at least some species (e.g., H. gouveai and H. circumdata) the prepollical spines are simple, curved elements terminating in a sharp point. The prepollex is weakly bifid in H. biobeba (Bokermann and Sazima, 1973) and strongly bifid in H. alvarengai, langei, and martinsi (Bokermann, 1956, 1964, 1966). The tadpoles show only slight adaptations to lotic environments. All known tadpoles of the group (H. biobeba, ibitiguara, nanuzae, and sazimai) are large (60-66 mm in total length), have only slightly enlarged, anteroventral oral discs, and have mod- erately long tails and moderately low fins terminating ina point. Marginal papillae are complete across the upper la- bium in H. ibitiguara and H. nanuzae, whereas a narrow median gap is present in H. biobeba and H. sazimai; the LTRF is 2/4 in H. biobeba and H. sazimai, and 2/5 in H. ibitiguara and H. nanuzae (Bokermann and Sazima, 1973; Cardoso, 1983; Cardoso and Andrade, 1982). Although there are many similarities between members of the H. circumdata and H. pulchella groups in structure, coloration, and larval features, only four of these can be considered synapomorphies; these are the sharp prepollical spines, hypertrophied forelimbs, larvae with moderately long tails with moderately low fins, and increased LTRF in larvae. How meaningful are these characters? The elongation of the prepollex into a curved spinous structure is unequivocally a derived character state in hylid frogs. Prepollical spines are present (and nuptial excres- cences absent) in five groups of Hyla inhabiting the neotropical lowlands—H. albomarginata Group (8 species), H. albopunctata Group (5 species), H. boans Group (8 spe- cies), H. geographica Group (6 species), H. granosa Group (3 species)—and Plectrohyla (16 species) in nuclear Central America. Examination of skeletal preparations of repre- sentative species in these groups revealed that the prepollical spines vary from shallowly sigmoid to crescent shaped in nearly all species examined; therefore, the spines are like the nonbifid spines in the H. circumdata and H. pulchella groups. In H. granosa, the proximal half of the prepollex is a quadrangular plate, from the distal edge of which a sharp spine curves medially. The tips of the spines are not sharply pointed in H. albopunctata, calcarata, and geographica. In Plectrohyla, there is a transformation series from blunt spines to simple pointed spines to bifid pointed spines (Duellman and Campbell, 1992). Although none of the species in these five groups of Hyla has noticeably enlarged forelimbs, elaborations of the cristae on the humerus were noted in the Hyla boans Group. The crista medialis is expanded in H. boans, crepitans, faber, pardalis, and barely so in H. rosenbergi, whereas the crista ventralis also is expanded in H. faber. Male-male combat using prepollical spines has been observed in three of these species—H. faber (Lutz, 1960), H. pardalis (Lutz, 1973), and H. rosenbergi (Kluge, 1981). In most species of Plectrohyla, the cristae are expanded; although male-male combat has not been observed in Plectrohyla, scars on adult males of P. hartwegi and P. teuchestes are consistent with wounds caused by prepollical spines (Duellman and Campbell, 1992). Frogs of the Hyla albomarginata, albopunctata, boans, and geographica groups have distinctive dentigerous processes of the vomers. Each process is angulate with a median transverse section continuous with a much longer poste- rolateral section; the median section generally is between the posterior margins of the choanae, and the posterolat- eral sections extend to, nearly to, or overlap the neopalatines, which have ventral ridges. Similar dentiger- ous processes are characteristic of Osteocephalus, but the two sections of each process are nearly equal in length, and the posterolateral section is widely separated from the neopalatine (Trueb and Duellman, 1971). The dentigerous processes in species in the H. granosa group are relative small, slightly arched structures between the posterior mar- gins of the choanae. The larvae of these groups are typical pond-type tad- poles with the tail no more than twice the length of the body, relatively high caudal fins terminating in a point, and relatively small oral discs directed anteroventrally with a median gap in the marginal papillae on the upper la- bium. Most species have a LTRF of 2/3, but Hyla rufitela in the H. albomarginata Group, and H. faber, pardalis, and rosenbergi in the H. boans Group have LTRFs of 2/4 (Bokermann, 1968; Cei, 1980; Duellman, 1970). The presence of enlarged prepollical spines lacking a quadrangular base is a synapomorphy of the Hyla albomarginata, albopunctata, boans, circumdata, geographica, and pulchella groups. Prepollical spines apparently evolved independently in Plectrohyla from an ancestor having a moderately enlarged, blunt spine (represented by the sis- ter group, the H. bistincta Group) (Duellman and Campbell, 1992). The prepollical spine in the H. granosa Group, which has distinctly different dentigerous processes of the vomers, is structurally different and apparently represents another independent evolution. The combined H. circumdata-pulchella Group is distinguished by a single oS) synapomorphy, hypertrophied limbs in adult males. We are unaware of any synapomorphies that distinguish the H. circumdata Group from the H. pulchella Group, so we combine these into a single group, the H. pulchella Group. This group is distinguished from the other groups with prepollical spines by having hypertrophied forelimbs in adult males. A single derived character (angulate denti- gerous processe of the vomers) is a synapomorphy for the 8 ScIENTIFIC PAPERS, NATURAL History Museum, THE UNIVERSITY OF KANSAS H. albomarginata, albopunctata, boans, and geographica groups. The H. pulchella Group is especially diverse with 20 species in streams in mountainous regions in southeast- ern Brazil, one species (H. pulchella) in the lowlands of southeastern Brazil and the Platine Region, one taxon (H. pulchella cordobae)in the isolated Cordillera de Cordoba, and eight taxa (the seven species recognized here, plus H. pulchella riojana) in the Andes. DISCUSSION Our original purpose was to review the systematic sta- tus of species of Hyla in the Andes of Argentina, Bolivia, and Peru. Our confidence in the taxonomic conclusions is clouded by four factors: (1) the unknown status of H. pulchella riojana in the Andes to the south of the known range of H. andina, (2) the absence of data on the tadpoles and advertisement calls of some of the species, (3) the ex- istence of another, apparently undescribed, species in the H. armata Group now known only from tadpoles, and (4) incomplete knowledge of the skeletal features of many of the species. Nonetheless, we consider our efforts at the al- pha taxonomic level to be an important contribution to the taxonomy of Andean Hyla. It is evident from the section on phylogenetic relation- ships that existing knowledge of South American frogs now placed in the genus Hyla is too meager to permit mean- ingful phylogenetic analyses. Complete data sets—exter- nal morphology of adults, osteology, tadpoles, advertise- ment calls, and karyotypes—exist for probably less than 20% of the species. Although tissues for molecular analy- ses are being acquired, material is insufficent at this time for meaningful systematic analyses of relationships among the diverse phenetic groups of Hyla. However, on the ba- sis of our incomplete data on morphology, we are reason- ably confident that the eight groups of Andean Hyla de- fined herein are natural groups. Obviously, thorough docu- mentation of the extralimital relationships of all of the Andean groups of Hyla is not only beyond the scope of this paper but is not feasible at this time. We concur with previous suggestions that the H. columbiana, garagoensis, and labialis groups are most closely related to groups that have a diploid number of 30 chromosomes and that in- habit the tropical lowlands. Also, we have provided evi- dence that the H. pulchella Group is monophyletic and in- cludes species formerly recognized in the Hyla circumdata Group in southeastern Brazil. Finally, we have noted that the peculiar mental gland that is a synapomorphy of spe- cies in the H. bogotensis Group may be shared with H. granosa in the cis-Andean lowlands; but it is unknown if the gland in the latter is homologous with that in the H. bogotensis group. Hopefully, the suggestions made herein will inspire our colleagues and future workers to investi- gate these fascinating problems; we wish them well in their endeavors. LITERATURE CITED Andersson, L. G. 1938. Batrachians from Burma collected by Dr. R. Mal- aise and from Bolivia and Ecuador collected by Dr. C. Hammerlund. Arkiv for Zoologi 30A:1—24. Altig, R., and G. F. Johnston. 1989. Guilds of anuran larvae: relationships among developmental modes, morphologies, and habitats. Herpeto- logical Monographs 3:81-109. Ardila-Robayo, M. C., P. M. Ruiz-Carranza, and S. H. Roa—Trujillo. 1993. Una nueva especie de Hyla del grupo larinopygion (Amphibia: Anura: Hylidae) del sur de la Cordillera Central de Colombia. Revista de Academia Colombiana de Ciencias 18:559-565. Ayarzagiiena, J., and J. C. Sefiaris. 1993. Dos nuevas especies de Hyla (Anura: Hylidae) para las cumbres tepuyanas de Estado Amazonas, Venezuela. Memorias de la Sociedad de Ciencias Naturales La Salle 53:127-146. Barrio, A. 1965. Las subespecies de Hyla pulchella Dumeril y Bibron (Anura: Hylidae). Physis 25:115-128. Berg, C. 1896. Batracios Argentinos. Enumeracion sistematica, sinonimica y bibliografica de los batracios de la Republica Argentina. Anales del Museo Nacional de Historia Natural de Buenos Aires 5:147-226. Bogart, J. P. 1973. Evolution of anuran karyotypes. Pp. 337-349 in J. L. Vial (ed.), Evolutionary Biology of the Anurans. Columbia: Missouri: Univ. Missouri Press, xii + 470. Bokermann, W.C. A. 1956. Sobre uma nova espécie de Hyla do Estado de Minas Gerais, Brasil. Papéis Avulsos do Deptamento do Zoologia, Secretaria da Agricultura, Sao Paulo 12:357-362. Bokermann, W. C. A. 1964. 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Occasional Papers of the Museum of Natural History, The University of Kansas 1:1-47. Vellard. J. 1970. Contribucion al estudio de los batracios andinos. Revista del Museo Argentino de Ciencias Naturales, Zoologia 10:1—21. Watrous, L. E., and Q. D. Wheeler. 1981. The outgroup comparison method of character analysis. Systematic Zoology 30:1-11. Wiley, E. O. 1981. Phylogenetics. The Theory and Practice of Phylogenetic Sys- tematics. New York: John Wiley and Sons, xv + 439. APPENDIX SPECIMENS EXAMINED Localites are given only for those specimens of the Hyla armata and Hyla pulchella groups from the Andes. Some specimens in the Museo de Historia Natural Noel Kempff Mercado (MNK) have not been catalogued; they are referenced by the field numbers of Michael B. Harvey (MBH) and Eric N. Smith (ENS). Other taxa are listed only by museum codes and numbers; unless specified otherwise, specimens are standard alco- holic preparations; C&S = cleared-and-stained skeletal preparations. Hyla albonigra BOLIVIA: Cochabamba: Cochabamba, 2600 m, ZSM 18/1949 (8); Incachaca, 2500 m, AMNH 39565-67; Rio Julpe, 30 km S Totora, Prov. Mizque, 2000 m, EBD 28925 (C&S),28926-28. Chuquisaca: Puka Khara, NHRM 1873 (7) (syntypes of Hyla ocapia); Rio Tomoyapo, 1 km E Iscayachi, 3416 m, MNK 840-46, 851, 858; Tacopaya, MNHN 4817 (2) (syntypes of Hyla zebra). Hyla andina ARGENTINA: Catamarca: Arroyo del Durazno, 19 km N La Merced, 750 m, KU 160410-21, 160408 (tadpoles), 160409 (young); 3.5 km NE La Merced, KU 160370-406, 160407 (tadpoles); Nacimiento, ZSM 18/1922 (3) (syntypes). Jujuy: 8.3 km SSW Perico del Carmen, 1230 m, KU 160342-— 60, 160361-62 (skeletons), 160363 (young), 160364 (tadpoles); Rio Grande, 10 km N Tilcara, 2670 m, 160337-41. Salta: Finca Jakulika, 560 m, KU 182964-67. Tucumédn: El Cadillal, 500 m, KU 160368, 160369 (young); Hualinchai, 8 km W San Pedro de Colalao, KU 128922-26, 128927 (skel- eton); Tafecillo, 1275 m, KU 182958-59; 21 km SSE Tafi del Valle, 1600 m, KU 160365, 160366 (tadpoles), 160367; Km 38, Tafi del Valle-Amaicha del Valle road, 1390 m, KU 182960, 182961 (C&S), 182962; Km 44. Tafi del Valle—Amaicha del Valle road, 1640 m, KU 182963. BOLIVIA: Cochabamba: Carahuasi, Prov. Carrasco, UTA 39668-69; Cochabamba, 2600 m, KU 125378, 126083, 154614; 62.7 km E Cochabamba on road to Villa Tunari, USNM 257834; 76.8 km from Cochabamba on road to Villa Tunari, USNM 257835—-44; 10 km N Colomi, Prov. Chapare, 3400 m, CET A122-27; 8.6 km N EI] Empalme, road to Cochabamba, UTA 39670—90; 2.9 km S Epizana, 3070 m, KU 183385-93, 183814 (tadpoles), 183815 (eggs); 3 km S Epizana, 2825 m, KU 160143, 160144 (tadpoles); 34.2 km W Epizana, 3260 m, KU 183394-95; Rio Chua Kocha, 9 km N Santa Cruz de la Sierra-Cochabamba road, Prov. Carrasco, 2200 m, CET A144—45; Rio Corani, on road to Tablasmontes, Prov. Chapare, 2400 m, CET A138-143; Serrania de La Siberia, 45 km from Comarapa on road to Pojo, Prov. Carrasco, 2500 m, CET A133-35; near Tiraque, about 78 km from Cochabamaba on road to Santa Cruz de la Sierra, Prov. Arani, 3550 m, CET A136-37; Totora, USNM 93218. La Paz: Cota Cota, Prov. Murillo, USNM 281287, 281289-93, 28129597, 281383-91, 281473. Potost: 6 km W Betanzos, 3330 m, KU 160195; Rio Tupiza, 12.5 km SSE Tupiza, 2920 m, KU 160236—42, 160243 (skeleton). Santa Cruz: Comarapa, Prov. Caballero, 1800 m, CET A117-21; Rio Comarapa, Prov. Caballero, 1750 m, CET A128— 32; 5 km E Samaipata, Prov. Florida, 1650 m, CET A116; Vallegrande, Prov. Vallegrande, 2100 m, CET A146. Hyla armata BOLIVIA: Cochabamba: Carahuasi, Prov. Carrasco, CET A-484-85 (tad- poles); 0.75 km E Khara Huasi, Prov. Carrasco, UTA 40035; Miguelito, 99 km from Cochabamba on road to Villa Tunari Chapare Prov., 1769 m, USNM 257812-17; Paracti, 83 km from Cochabamba on road to Villa Tunari, Chapare Prov., 2044 m, USNM 257818; Rio Chua Kocha, 9 km N Santa Cruz, 2200 m, CET A39-40, MNK 963; 68.8 km SW Villa Tunari, 1860 m, KU 183299-325. La Paz: 10 km down from Chuspipata, 2350 m, MNHNB B140; La Paz, BMNH 1947.2.13.59-60 (syntypes); Serrania Bellavista, 1450 m, CET A41-42, CET A483 (tadpole). Santa Cruz: El Chapé, Prov. Florida, MNK (MBH 1280, 1561, ENS 3590, 3593-94, 3611, 3621), UTA 40032-34, 40036-37; Quebrada Huertillas, Prov. Florida, MNK (ENS 3630), UTA 40038-39. PERU: Ayacucho; Tutumbaro, Rio Piene, 1840 m, KU 163345, Cuzco: 1 km W Amaybamba, tributary of Rio Santa Maria, FMNH 235542 (tad- poles). Buenos Aires, 2400 m, KU 173219-22; Hacienda Huyro, between Huayopata and Quillabamba, 1720 m, KU 196832, LSU 32061-62; Rio Cosnipata, 4 km SW Santa Isabel, 1700 m, KU 163322-33, 163334 (C&S), 163335-44, 164085 (skeleton), 164086-87 (C&S). Puno: Rio Quiquira, 1 km N Abra Marrancunca, 2000 m, USNM 299310-38; MHNSM 10348, 10350. Hyla balzant BOLIVIA: Cochabamba: Charuplaya, 1350,. BMNH 1947.2.13.64-73 (syntypes of H. callipleura), NHMW 6223 (2); Chicotal, Prov. Chapare, KU 148602-03 (skeletons); Miguelito, 99 km from Cochabamba on road ANDEAN GROUPS OF HYLA 4] to Villa Tunari, Prov. Chapare, USNM 257819-21; 2.5 km NW of Miguelito, Prov. Chapare, 1769 m, USNM 257822-26; Paracti, 83 km from Cochabamba on road to Villa Tunari, Prov. Chapare, 2044 m, Puente de la Via, Yungas Valley, BMNH 1940.4.9.99; USNM 257827-33; San Ernesto, 500 m, BMNH 1947.2.13.74 (syntype of H. callipleura); Serrania Bellavista, Prov. Noryungas, 1450 m, CET A99-100; Tablasmontes, Prov. Chapare, 2,400 m, CET A93, A96-98, A107, A110-15; 58.1 km W Villa Tunari, 1980 m, KU 183326-29; no specific locality, BMNH 1919.11.12.16, UMMZ 68173 (10), 68174 68175 (21), USNM 93222. La Paz: 4.4 km E Chulumani, 1760 m, KU 183330-34, 183425; Coroico, 1750 m, 18333746, 183320-21 (tad- poles); 33.9 km E Unduavi, 2160 m, KU 183335-36; Yungas, between Coroica and Chulumani, MSNG 28872 (holotype). Santa Cruz: 10 km NW Masicuri, CET A1453. PERU: “Kalinourhin,” NHMW 18448. Cuzco: 72 km (by road) N Paucartambo, USNM 298210; 73 km (by road) N Paucartambo, USNM 298211-13; 74 km (by road) N Paucartambo, USNM 298214-18; 75 km (by road) N Paucartambo, USNM 298219-21; 4 km WSW Santa Isabel, KU 139212, 163315-18, 164052-53 (tadpoles), 164054 (young). Junin: Chanchamayo, BMNH 1911.12.13.81-82. Pasco: Huancabamba, 925 m, BMNH 1911.12.13.63, 1912.11.1.25. Puno: Abra Marracunca, 14 km W Yanahuaya, 2210 m, USNM 299399-49; Cerros de Tavara, 900 m, USNM 332412-14; Juliaca, AMNH 6121-28; La Union, Rio Huacamayo, Carabaya, 620 m, BMNH 1907.5.7.33; 11 km (airline) NNE Ollachea, 1880 m, USNM 298953-58; Santo Domingo, Carabaya, 1850 m, BMNH 1902.11.28.17-21, 1903.6.30.16-17, 1907.5.7.34-35. Hyla marianitae ARGENTINA: Salta: Rio Pescado, Serrania de las Pavas, SW part of Parque Nacional Baritu, Depto. Oran, MACN 32330 (holotype). BOLIVIA: Cochabamba: Carahuasi, Serrania de La Siberia, 18 Km N Santa Cruz de la Sierra on road to Cochabamba, 1700 m, CET A85-88, A90, A95, A101—-06, A108, A838, MNK (MBH 1559, 1680, ENS 3874), UTA 40027-29; 2 km from Carahuasi on road to Rio Grande, Prov. Carrasco, UTA 40026. Santa Cruz: El Chapé, Prov. Florida, MNK (MBH 2671-72); Arroyo Las Juntas, 8 km NW Samaipata, Prov. Florida, 1400 m, CET A89; El Chape, Prov. Florida, UTA 40025; Quebrada El Fuerte, 5.1 km E Samaipata, UTA 40030-31; Quebrada Huertillas, Prov. Florida, MNK (ENS 3637), UTA 40024; Rio Cheyo, Prov. Ichilo: Parque Nacional Ambor6, 700 m, CET A109; stream between La Angostura and Bermejo, Prov. Florida, 800 m, CET A91-92, A94. Hyla melanopleura PERU: Pasco: Huancabamba, 900 m, BMNH 1947.2.13.45-58 (syntypes). Hyla palaestes PERU: Ayacucho: San José, Rio Santa Rosa, 1005 m, KU 196844—53, LSU 26002-15; Tutumbaro, Rio Piene, 1840 m, KU 163305 (holotype), 163306-14, 164051 (tadpoles), 164092 (skeleton). OTHER MATERIAL EXAMINED Hyla albomarginata: KU 91925, 91926 (skeleton), 91927, 211294 (larvae); H. albopunctata: KU 91931-33, 91936 (skeleton). 146854 (larvae); H. albosignata: KU 9193740, 91942 (skeleton), 211297 (larvae); H. alytolylax: KU 117982-84, 111904 (C&S), 120851-60, 166310 (C&S), 180351 (larvae); H. boans: KU 104408-09 (skeletons), 109428 (larvae), 150020-24; H. bogotensis: KU 169434-60, 170109 (C&S); H. calcarata KU 105396 (skel- eton), 112355-56 (larvae), 122658-65; H. callipeza: KU 169566-70; H. carnifex: KU 98142 (C&S), 111867-68 (C&S), 166311-14 (C&S); 111838—66, 112360 (larvae); H. caucana: KU 189598; H. chlorostea: USNM 257811 (ho- lotype); H. circumdata: KU 71796-97, 91987-89, 152233; H. columbiana: KU 139524 (larvae), 148483 (larvae), 169461-98, 170110-12 (C&S); H. colymba: KU 31864, 31865 (C&S) 77414, 95979, 116357, 116779-81 (larvae); H. crepitans: KU 101550-52, 167663-64 (skeletons), 169499-501; 1702046 (lar- vae); H. denticulenta: KU 133450-52, 139531 (larvae); H. faber: KU 92310 (skeleton), 92311-12; H. fasciata: KU 205461-75, 205476 (C&S), 215797-98 (larvae); H. geographica: KU 92040 (skeleton), 146789 (larvae), 150229-45; H. granosa: KU 109495 (C&S), 122735-38; H. labialis: KU 124888 (larvae), 16950246, 169547 (C&S), 179178 (C&S), 187800 (C&S); H. lanciformis: KU 104399-400 (skeletons), 112346-48 (larvae), 122739-60; H. larinopygion: KU 144127-28; H. lascinia: KU 181079-88, 181145 (larvae); H. lindae: KU 155475, 155477 (skeleton), 155523 (larvae), 202728-31, 202966 (larvae); H. multifasciata: KU 127811-15, 128478 (skeleton); H. pacha: KU 202760-63; H. palmeri: KU 148703-04, 169584-85, 170209, 170210 (larvae), 178800; H. pantosticta: KU 140385, 190000-01, 202732-33, 202965 (larvae); H. pardalis: KU 92109, 92111-12 (skeletons); H. pelidna: KU 181108-10; H. pellucens: KU 109537-53, 109556 (skeleton); H. phyllognatha: KU 143197-205, 146793— 94 (larvae), 16433447, 166315-16 (C&S); H. platydactyla: KU 139521-22 (larvae), 166990-97, 180195-97, 167766 (C&S); H. praestans: KU 169574— 80, 170113 (C&S). H. psarolaima: KU 164313-15, 164316 (skeleton), 169582— 85, 202727; H. ptychodactyla: KU 132553-54 (larvae), 180360 (larvae), 209780-81; H. pugnax: KU 101554-55; H. pulchella: BMNH 1947.2.23.35— 36 (syntypes of Hyla agrestis), 1947.2.23.45 (holotype of Hyla vauterti), KU 71822-24, 92126-29, 92130-31 (skeletons), 197291-94,, MNHN 163, 4836 (syntypes), 176, 751 (syntypes of Hyla leucomeias); H. raniceps KU 92143 (skeleton), 130070-72, 130074, 130077-78; H. rosenbergi: KU 77291-98, 84990-91 (skeletons), 180356 (larvae); H. rufitela: KU 77306-13, 77662 (skel- eton), 104300 (larvae); H. simmonsi: KU 169554-65; H staufferorum: KU 217694—-96, 218315 (larvae); H. torrenticola: KU 169571-73, 171187 (larvae); H. sp.: KU 181882-83 (larvae). Plectrohyla avia: KU 106295 (skeleton); P. hartwegi: KU 192443 (skeleton); P. teuchestes: KU 59831 (skeleton). 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