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diff --git a/32505-8.txt b/32505-8.txt new file mode 100644 index 0000000..e43d444 --- /dev/null +++ b/32505-8.txt @@ -0,0 +1,2526 @@ +The Project Gutenberg EBook of The Systematics of the Frogs of the Hyla +Rubra Group in Middle America, by Juan R. León + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: The Systematics of the Frogs of the Hyla Rubra Group in Middle America + +Author: Juan R. León + +Release Date: May 24, 2010 [EBook #32505] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK FROGS OF HYLA RUBRA GROUP *** + + + + +Produced by Chris Curnow, Joseph Cooper and the Online +Distributed Proofreading Team at https://www.pgdp.net + + + + + + +UNIVERSITY OF KANSAS PUBLICATIONS + +MUSEUM OF NATURAL HISTORY + + +Volume 18, No. 6, pp. 505-545, 7 figs., 4 pls. + +December 2, 1969 + + +The Systematics of the Frogs of the +_Hyla rubra_ Group in Middle America + + + +BY + +JUAN R. LEÓN + + + +University of Kansas +Lawrence +1969 + + + +UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + +Editors: Frank B. Cross, Philip S. Humphrey, Robert M. Mengel. + + +Volume 18, No. 6, pp. 505-545, 7 figs., 4 pls. +Published December 2, 1969 + +UNIVERSITY OF KANSAS +Lawrence, Kansas + + +PRINTED BY +ROBERT R. (BOB) SANDERS, STATE PRINTER +TOPEKA, KANSAS +1969 + + + + +The Systematics of the Frogs of the _Hyla rubra_ Group in Middle +America + +BY + +JUAN R. LEÓN + + + + +CONTENTS + + + PAGE + +INTRODUCTION 508 + Acknowledgments 508 + Materials and Methods 509 + +THE HYLA RUBRA GROUP 509 + Key to Species and Subspecies 510 + Key to Known Tadpoles 511 + +ACCOUNTS OF SPECIES AND SUBSPECIES 511 + _Hyla boulengeri_ (Cope) 511 + _Hyla foliamorta_ Fouquette 520 + _Hyla rubra_ Laurenti 524 + _Hyla elaeochroa_ Cope 525 + _Hyla staufferi_ Cope 532 + _Hyla staufferi staufferi_ Cope 537 + _Hyla staufferi altae_ Dunn 540 + +EVOLUTIONARY HISTORY 540 + +LITERATURE CITED 543 + + + + +INTRODUCTION + + +The tree frogs of the _Hyla rubra_ group are abundant and form a +conspicuous element of the Neotropical frog fauna. Representatives of +the group occur from lowland México to Argentina; the greatest +diversity is reached in the lowlands of southeastern Brazil (Cochran, +1955). The group apparently originated in South America; the endemic +Central American species evolved from stocks that invaded Middle +America after the closure of the Colombian Portal in the late Pliocene. + +Dunn (1933) partially defined the _rubra_ group as it occurs in Central +America. Cope (1865, 1876, 1887), Brocchi (1881), Boulenger (1882), +Günther (1901), Noble (1918), Kellogg (1932), Dunn and Emlen (1932), +Stuart (1935), and Gaige (1936) dealt with the Middle American species +now considered to make up the _rubra_ group. More recently, Taylor +(1952, 1958), Fouquette (1958), Starrett (1960), and Duellman (1960, +1963, 1966a) studied aspects of the taxonomy and biology of the species +of this group. The five species of the _rubra_ group in Central America +have received ten different names. One species, _Hyla staufferi_, has +received five names (two subspecies are recognized herein). _Hyla +boulengeri_ was named in the genus _Scytopis_, but the type species of +_Scytopis_ is a member of the genus _Phrynohyas_ Fitzinger, 1843 +(Duellman, 1956.) + +Little has been published concerning the ecology, life history, +osteology, and mating calls of the Middle American species of this +group. The purpose of the present report is to describe the species +occurring in Middle America and to comment on their distributions, +ecology, cranial osteology, and mating calls, and in so doing provide +evidence for the evolutionary history of the species inhabiting Middle +America. + + +Acknowledgments + +For permission to examine specimens in their care, I am grateful to +Drs. Richard G. Zweifel, American Museum of Natural History (AMNH); +Robert F. Inger, Field Museum of Natural History (FMNH); Ernest E. +Williams, Museum of Comparative Zoology (MCZ); Hobart M. Smith, +University of Illinois Museum of Natural History (UIMNH); Charles F. +Walker, University of Michigan Museum of Zoology (UMMZ); Jay M. Savage, +University of Southern California (USC); James A. Peters, United States +National Museum (USNM); Richard J. Baldauf, Texas Cooperative Wildlife +Collection (TCWC); and W. Frank Blair, Texas Natural History Collection +(TNHC). KU refers to specimens in the Museum of Natural History, +University of Kansas. For the loan of tape-recordings of mating calls I +thank Drs. W. Frank Blair, University of Texas, and Richard G. Zweifel, +American Museum of Natural History. + +I am indebted to the Ford Foundation-Universidad de Oriente (Venezuela) +Science Project for a scholarship which enabled me to study for two +years at The University of Kansas, foster institution of the project. I +have benefited by being able to work in the Museum of Natural History +at The University of Kansas and I am grateful to Dr. E. Raymond Hall, +Director, for providing space and equipment. + +I gratefully acknowledge the assistance and advice of Dr. William E. +Duellman, who suggested and directed this work, made available +specimens under his care and gave much of his time in reading the +manuscript and suggesting improvements. I am grateful to Dr. Frank B. +Cross who critically read the manuscript and made many editorial +suggestions. I am indebted to Linda Trueb for assistance with the +osteological aspects of this study; she helped to clarify many +confusing points. I am grateful to Charles W. Myers for making +available his field notes on these frogs in Panamá, to Arthur C. +Echternacht for reading part of the manuscript, and to John D. Lynch +for many suggestions and helpful criticisms. The illustrations were +executed by David M. Dennis. + + +Materials and Methods + +For the purposes of the present study I examined 1383 preserved +specimens, 50 skeletons, and 9 lots of tadpoles. External +characteristics used in the analysis of variation are those currently +employed in the study of anuran systematics. Twelve measurements +and six proportions were taken in the manner described by Duellman +(1956). Only the most important references are given in the synonymies, +except those of the two subspecies of _Hyla staufferi,_ which are +more nearly complete. The taxonomic history of each frog is discussed +under _Remarks_ in each account. The cranial osteology was studied +by using skeletons and cleared and stained specimens of all species. +Developmental stages of tadpoles were determined from Gosner's (1960) +table. Personal field work in Central America in the summer of 1966 +provided an opportunity to make observations on the ecology, calling +sites, and color in life; these data were supplemented by field notes +from, and discussions with, Dr. William E. Duellman and Charles W. +Myers. + +The mating calls of the frogs were recorded in the field on Magnemite +and Uher Tape Recorders by Dr. Duellman in the course of his work on +the hylid frogs of Middle America--supported by grants from the +National Science Foundation (G-9827 and GB-1441). These recordings, +plus those borrowed from other institutions, provided 50 tapes for +analysis of the mating calls. The calls were analyzed on a Vibralyzer +(Kay Electric Company). + + + + +THE HYLA RUBRA GROUP + + +_Definition._--The species forming the group are small to moderate-sized +tree frogs (maximum snout-vent length of males of various species 20-49 +mm.), distinguished from other groups in the genus _Hyla_ as follows: +Brown, grayish brown, or yellowish tan above; thighs plain, marbled +with dark brown, or having vertical bands; vocal sac single, median, +subgular; snout flat, protruding, rounded or pointed; webbing between +fingers reduced or absent; web between first and second toes reduced to +fringe on second toe, rest of toes about half webbed; tarsal fold +reduced or absent; shanks robust; inner metatarsal tubercle larger than +outer; prevomerine teeth on transverse ridges between small to large +sized choanae; skull generally longer than wide; nasals large (length +more than 40 per cent total length of skull) and having pointed +maxillary processes; maxillary bearing small ventromedial palatine +process; quadratojugal slender, always joined to maxillary by bony +suture; auditory region of proötic slender and short; delicate +spatulate columella ventral to crista parotica, broad basally, +compressed anterolaterally, slightly rounded distally; anterior arm of +squamosal extending about half distance to maxillary; sphenethmoid +wider than long; frontoparietal fontanelle present or absent; +prevomerine, premaxillary, and maxillary teeth present; prevomer with +two lateral processes forming incomplete bony margin to internal nares; +tadpoles having pointed xiphicercal tail, snout short, rounded; 2/3 +tooth rows; dorsal fin deeper than ventral fin; sinistral spiracle; +short dextral anal tube not reaching edge of ventral fin; mating calls +consisting of single long note or series of short notes. + +_Composition._--This group contains about 24 currently recognized +species, most of which occur in Brazil. Only five species--_boulengeri,_ +_elaeochroa_, _foliamorta_, _rubra_, and _staufferi_ with two +subspecies--occur in Central America. _Hyla boulengeri_ and _rubra_ are +widespread in South America, and _foliamorta_ occurs in Colombia, +whereas the other species are known only from Middle America. + +_Distribution._--The species of the _Hyla rubra_ group range from the +lowlands of northern Argentina and Bolivia to southern Tamaulipas and +Guerrero, México. + +_Comments._--In Central America two subgroups of species can be +recognized. _Hyla boulengeri_ and _H. foliamorta_ are distinctive in +the large size of adults (snout-vent lengths 41-49 mm.); both have +prominent bars on the thighs, a well-defined interorbital triangular +mark, blotches or spots dorsally, and large choanae. _Hyla elaeochroa,_ +_H. rubra,_ and _H. staufferi_ are smaller (snout-vent lengths 29-40 +mm.); they have the thighs weakly barred or vermiculate anteriorly and +posteriorly or unmarked, an ill-defined interorbital triangular mark, +linear markings dorsally, and small choanae. + + +Key to Species and Subspecies + + 1. Larger frogs (males to 49 mm. snout-vent length); thighs + strongly barred; supratympanic fold black; dorsum blotched or + spotted 2 + + Smaller frogs (males to 40 mm. snout-vent length); thighs weakly + barred or plain; supratympanic fold pale brown; dorsum usually + having linear pattern 3 + + 2. Dorsum tuberculate; snout subacuminate; vocal sac flecked with + brown; tarsal fold rudimentary; web absent between fingers; black + spots absent in scapular region _H. boulengeri_ + + Dorsum smooth; snout pointed; vocal sac dark gray; tarsal fold + absent; trace of web between fingers; two or more elongate dark + spots in scapular region _H. foliamorta_ + + 3. Snout-vent length more than 30 mm.; tympanum 2/3 to 3/4 diameter + of eye; prevomerine elevations about size of choanae 4 + + Snout-vent length less than 30 mm.; tympanum less than 1/2 diameter + of eye; prevomerine elevations smaller than choanae 5 + + 4. Thighs mottled posteriorly; discs on fingers about 1/2 size of + tympanum; faint dark line from nostril to eye _H. rubra_ + + Thighs faintly barred or plain posteriorly; discs on fingers + about size of tympanum; distinct dark line from nostril to eye + _H. elaeochroa_ + + 5. Dorsum brown with irregular dorsolateral stripes and + interrupted paravertebral stripes; two transverse bars on shanks; + interorbital bar present _H. staufferi staufferi_ + + Dorsum gray with complete dorsolateral and paravertebral stripes; + longitudinal stripe on shank; interorbital bar absent + _H. staufferi altae_ + + +Key to Known Tadpoles + + 1. Entire lower beak black; beaks moderate-sized, serrate; dorsal + fin high, extending to middle of back 2 + + No more than half of lower beak black; beaks small, finely serrate; + dorsal fin lower, barely extending onto body 3 + + 2. Papillae present only laterally _H. boulengeri_ + Papillae present laterally and ventrally _H. foliamorta_ + + 3. Distinct brown stripe from nostril to eye; two stripes below + eye, _H. elaeochroa_ + + Faint stripe from nostril to eye; no stripe below eye + _H. staufferi_ + + + + +ACCOUNTS OF SPECIES AND SUBSPECIES + + +_Hyla boulengeri_ (Cope) + + _Scytopis boulengeri_ Cope, Bull. U.S. Natl. Mus., 32:12, December + 1, 1887 [Holotype.--USNM 13974, from "Nicaragua"; J. A. McNiel, + collector]. + + _Hyla boulengeri:_ Günther, Biologia Centrali-Americana, Reptilia + and Batrachia, p. 267, June 1901. Noble, Bull. Amer. Mus. Nat. + Hist, 38:339, June 1918. Taylor, Univ. Kansas Sci. Bull., 35:856, + July 1, 1952. + +_Diagnosis._--Size large (Male to 49 mm., Female to 53 mm.); skull as +long as wide; frontoparietal fontanelle present; snout subacuminate; +canthus not pronounced; choanae large; tongue cordiform, slightly +longer than broad; interorbital triangle tubercular; skin on dorsum +tuberculate; tarsal fold reduced or absent; thighs, shanks, and tarsi +boldly barred with dark brown and pale yellow-green in life. + +_Description._--Head flattened, longer than wide; snout projecting +beyond lower lip; loreal region oblique; canthus not pronounced; length +of eye less than interorbital distance; tympanum large, about 70 per +cent of diameter of eye; interorbital triangle distinct; arms short; +fingers lacking web; palmar tubercle tripartite; subarticular tubercles +distinct; long tubercle on base of first finger; discs truncate; legs +long; tarsal fold reduced or absent; inner metatarsal tubercle rounded, +larger than outer, both elevated; subarticular tubercles distinct; one +phalanx free of web on second, third, and fifth toes, three free on +fourth toe (Fig. 1A and B); skin tuberculate on dorsum, less so on +flanks; skin of belly granular, that on chest and throat weakly +granular; tongue cordiform, longer than wide, free and notched behind; +vocal slits large, lateral to tongue. + + [Illustration: Fig. 1. A and B.--Hand and foot of _Hyla boulengeri_ + (KU 102173), × 3. C and D.--Hand and foot of _Hyla s. staufferi_× (KU + 57790), × 6] + +In life, dorsum tan or brown with dark spots on snout, head, and +scapular region; interorbital triangle and blotch posteriorly on dorsum +dark brown; flanks pale green; groin pale green or orange, mottled with +dark brown; thighs tan or brown above with dark transverse bars on +anterior and posterior surfaces; spaces between bars green or orange; +inner surfaces of shanks and outer surfaces of tarsi brown and orange; +foot brown above; forelimbs brown and pale green above, weakly barred; +belly creamy white with scattered brown spots; vocal sac creamy white +flecked with brown; lower jaw brown with white spots on lips (Pl. 1A). + +In preservative, head and dorsum dark brown with triangular spot +between eyes; dark spots on head and scapular region and dark brown +blotch posteriorly on dorsum; flanks creamy white with brown spots; +groin creamy white mottled with dark brown; thighs brown above with +dark brown transverse bars on anterior and posterior surfaces; inner +surfaces of shanks and outer surfaces of tarsi barred with pale brown; +dorsal surface of foot mottled brown and creamy white; ventral surface +of foot and toes pale brown; forelimbs faintly barred with pale brown; +belly white with a few pale brown spots; vocal sac flecked with pale +brown; lower jaw marked with small white spots on lips. + +_Variation._--Geographic variation is evident in the snout-vent length, +tibia length, and foot length, all in relation to snout-vent length, +and the relative size of the tympanum to the eye (Table 1). The largest +specimens are from the humid Pacific lowlands of Costa Rica; +individuals from the Caribbean lowlands of Costa Rica, Canal Zone, and +South America are smaller. A general trend for increase in size extends +from South America to the Pacific lowlands of Costa Rica. + +Most variation in color does not seem to be correlated with geography; +color variation is nearly as great within a given population as between +separated populations. However, most specimens from Rincón de Osa, +Puntarenas Province, Costa Rica, are dusky brown, but a few are darker. +In comparison with specimens from the Caribbean lowlands of Central +America, specimens from the Pacific slopes in Costa Rica have a darker +interorbital triangle. In some specimens from the latter area +rugosities are present on the borders of the interorbital triangle, on +the snout, on the upper eyelid, and on the heel. Specimens from the +Caribbean lowlands are less tuberculate, and most individuals from +there lack rugosities on the tarsus. Living individuals from Puerto +Viejo, Heredia Province, Costa Rica, and from the Canal Zone, Panamá, +are brown above with a metallic green tint. Rugosities are present on +the posterior edges of the forelimbs in some specimens from throughout +the range. In most respects, specimens from the Canal Zone resemble +those from the Caribbean lowlands of Costa Rica more than they resemble +those from the Pacific lowlands of Costa Rica, but some individuals +from the Canal Zone are less metallic above and have small white spots +dorsally on the body, head, and limbs. + +A moderate amount of color change from night to day has been noted. At +night, a male from Puerto Viejo, Heredia Province, Costa Rica, was +grayish tan above with slightly darker markings; the flanks were pale +yellowish green. By day, the dorsum was brown with darker markings, and +the throat was pale gray with white flecks; the rest of the venter was +creamy white. The groin was pale green with black mottling; the +anterior and posterior surfaces of the thighs and inner edges of the +tarsi were greenish yellow with black bars. + +TABLE 1.--Geographic Variation in Size and Proportions in Males of +_Hyla boulengeri_. (Means in parentheses below observed ranges.) + +========================================================================= + | | Snout-vent| Tibia | | + | | length | length/ |Tympanum/|Foot length/ +Locality | N | (mm.) | snout-vent| eye | snout-vent +--------------------+----+-----------+-----------+---------+------------- +Costa Rica: Tilarán | 23 | 37.4-48.7 | 0.52-0.58 |0.62-0.76| 0.39-0.45 + | | (43.8) | (0.55) | (0.71) | (0.42) + | | | | | +Costa Rica: Rincón | 10 | 41.4-46.1 | 0.54-0.60 |0.68-0.80| 0.40-0.45 + de Osa | | (44.0) | (0.57) | (0.74) | (0.43) + | | | | | +Costa Rica: Alajuela| 13 | 35.6-43.1 | 0.55-0.60 |0.63-0.78| 0.41-0.46 + Province | | (39.8) | (0.57) | (0.69) | (0.44) + | | | | | +Costa Rica: Puerto | 25 | 37.5-42.9 | 0.51-0.62 |0.63-0.79| 0.38-0.46 + Viejo | | (41.6) | (0.55) | (0.71) | (0.43) + | | | | | +Costa Rica: Suretka | 9 | 38.7-42.0 | 0.56-0.58 |0.53-0.72| 0.35-0.45 + | | (41.0) | (0.56) | (0.62) | (0.42) + | | | | | +Panamá: Canal Zone | 16 | 36.7-42.9 | 0.52-0.58 |0.70-0.78| 0.40-0.44 + | | (39.0) | (0.54) | (0.74) | (0.42) + | | | | | +Venezuela: Santomé | 4 | 35.5-40.9 | 0.45-0.48 |0.63-0.67| 0.36-0.40 + | | (38.5) | (0.46) | (0.65) | (0.38) + + +TABLE 2.--Comparison of Mating Calls in the _Hyla rubra_ Group. (Means +in parentheses below observed ranges.) + +============================================================================ + | |Notes| | | | Major frequencies + | | per |Duration| Pulses|Fundamental| (cps) + | |call | of note| per | frequency |--------------------- +Species | N |group| (sec.) | second| (cps) | Lower | Upper +---------------+---+----+---------+-------+-----------+---------+----------- +H. boulengeri | 8 | 1 | 0.24- | 80-120| 70-74 |1400-1820|2520-3182 + | | | 0.47 | (101)| (71) | (1611) | (2840) + | | | (0.35) | | | | + | | | | | | | +H. foliamorta | 7 | 1 | 0.23- | 50-60 | 52-61 | 912-1026|2736-3477 + | | | 1.86 | (51) | (56) | (918) | (3055) + | | | (0.69) | | | | + | | | | | | | +H. elaeochroa |15 | 2-95| 0.12- | 40-50 | 48-65 |1254-1586|2562-3477 + | | (19)| 0.24 | (42) | (57) | (1499) | (2911) + | | | (0.17) | | | | + | | | | | | | +H. s. staufferi|18 | 2-77| 0.13- |100-130| 96-130 |1582-1872|1962-3744 + | | (23)| 0.23 | (120) | (106) | (1743) | (3056) + | | | (0.18) | | | | + | | | | | | | +H. s. altae | 7 | 2-22| 0.14- |110-130| 104-117 |1853-2106|3379-4056 + | | (11)| 0.18 | (120) | (112) | (2008) | (3775) + | | | (0.15) | | | | + +_Cranial Osteology._--The skull of _Hyla boulengeri_ is as long as it +is wide, and is flat; the premaxillary is small and bears 13 to 17 +teeth (mean for 6 specimens, 14.9). The alary processes of the +premaxillaries are widely separated, concave posteriorly, and vertical. +Ventrally, the premaxillary is connected to the prevomer by bony +tissues. The maxillary is slender and bears 70 to 91 teeth (mean for 6 +specimens 78.1). The pars facialis of the maxillary is laterally convex +and about four times as high as the pars dentalis. + +The nasal is large (its length about 40 per cent of total length of +skull), and pointed anteriorly and posteriorly in dorsal view. The +nasals are separated anteromedially by the cartilaginous septum nasi. +One or two protuberances are present on the midlateral concavity of the +nasal. Posteriorly, the nasal overlaps the sphenethmoid and articulates +with the palatine. Dorsally the sphenethmoid is large, pentagonal, and +completely ossified. The frontoparietal is elongate, smooth, and bears +a small supraorbital process on the anterior edge of the orbit. A +keyhole-shaped frontoparietal fontanelle is present; the fontanelle is +narrow anteriorly and wide posteriorly. + +The bony part of the proötic is separated dorsally from the squamosal +by the cartilaginous crista parotica. The squamosal is small, its +anterior arm slender and pointed. The posterior arm of the squamosal is +pointed terminally and articulates with the proötic medially. + +The prevomer is large and elongate. Anteriorly the prevomer is +connected to the maxillary-premaxillary articulation; posteriorly, the +prevomer is separated from the sphenethmoid by cartilage. Each prevomer +bears six to nine teeth. The palatine is present and edentate. The +anterior end of the parasphenoid is broad (less pointed than in _Hyla +foliamorta_). The pterygoid is slender and well developed. + +_Natural History._--_Hyla boulengeri_ inhabits humid lowland tropical +forests and breeds in temporary ponds. Clasping pairs and gravid +females were observed at Puerto Viejo, Heredia Province, Costa Rica, on +June 21, 1966. Males were calling from depressions in decaying logs and +stumps, in forked stems, and from leaves of broad-leafed plants near +the pond. Males were observed in late July and early August calling +from _Calathea_ and _Heliconia_ leaves at the edge of a pond in the wet +forest of the Osa Peninsula. William E. Duellman informed me that he +collected calling males in January at El Real, Darién, and in March at +Almirante, Bocas del Toro, Panamá. Taylor (1952) found calling males in +June at Turrialba, Cartago Province, Costa Rica, and Dunn (1931a) +observed males calling in July, November, and December in Panamá. +Gravid females have been found from April to August. Breeding +activities of _Hyla boulengeri_ always seem to be associated with +temporary ponds; in Central America breeding apparently takes place +throughout most of the year. + +The mating call of _Hyla boulengeri_ consists of one short, moderately +low-pitched note. Each note has a duration of 0.24 to 0.47 second and +is repeated at intervals of one second to several minutes. The notes +have 80 to 120 pulses per second, a fundamental frequency of about 70 +cycles per second, and a dominant frequency of 2,840 cycles per second +(Table 2, Pl. 3A). + +The eggs are deposited in a mass in the water. No information is +available concerning early development. Tadpoles in advanced stages of +development were found in a temporary pond at Rincón de Osa, Puntarenas +Province, Costa Rica. The pond was about 10 cm. deep, had a muddy +bottom and lacked vegetation. Three recently metamorphosed young were +found in mid-August, 1966, on grass at the edge of another temporary +pond in the forest. + +_Tadpoles_--Twelve tadpoles are available. These were collected at +Rincón de Osa, Puntarenas Province, Costa Rica. The maximum size +represented is 34.0 mm., total length (stage 42 of development). + +A typical tadpole in stage 36 of development (KU 104295) has a body +length of 12.0 mm., tail length of 20.0 mm., and total length of 32.0 +mm. Other characters are as follows: depth of tail equal to length of +body; body deeper than wide; distance between eye and nostril equal to +that between nostril and tip of snout; mouth anteroventral, upper and +lower lips bare; papillae present laterally; tooth rows 2/3; upper rows +about equal in length; first upper row slightly, and second upper row +widely, interrupted medially; lower rows about equal in length, shorter +than upper rows; third lower row containing 5-10 large teeth; beak +strong, serrate; spiracle nearer anus than eye; anal aperture not +extending to border of ventral fin; caudal musculature slender +posteriorly, extending to tip of pointed tail; dorsal fin extending to +middle of body, slightly deeper than ventral fin; posterior three +fourths of tail spotted; rest of tail and body gray-brown or +transparent; hindlimbs flecked or spotted with brown (Table 3, Fig. 2A +and 3A). + +TABLE 3.--Sizes of Tadpoles of _Hyla boulengeri_ in Relation to +Developmental Stages. (Means in parentheses below observed ranges; +measurements in mm.) + + ======================================================= + Stage | N | Body length | Tail length | Total length + --------+---+-------------+-------------+-------------- + 30 | 1 | 11.0 | 22.2 | 33.2 + | | | | + 35 | 1 | 11.0 | 12.0 | 23.0 + | | | | + 36 | 3 | 9.5-12.0 | 20.0-21.5 | 31.0-32.0 + | | (11.2) | (20.5) | (31.7) + | | | | + 38 | 2 | 11.5 | 22.0 | 33.5 + | | | | + 42 | 2 | 10.5-13.0 | 21.0-22.0 | 32.5-34.0 + | | (11.8) | (21.5) | (33.3) + | | | | + 44 | 2 | 14.0-15.0 | 8.0-15.0 | 22.0-30.0 + | | (14.5) | (12.5) | (26.0) + | | | | + 46 | 1 | 15.0 | 15.0 | + +A recently metamorphosed young has a snout-vent length of 15 mm.; the +head is as long as wide, the eyes are prominent; the limbs are weakly +barred; the skin is rugose above and granular below. The venter is +immaculate; the dorsum and limbs are gray-brown in preservative (pale +green in life). The interorbital space, supratympanic fold, and +scapular region are darker than the rest of the body; the fingers lack +webbing; the webbing on the foot is the same as in adults; small +metatarsal tubercles are present, but the tarsal fold is absent. + + [Illustration: Fig. 2. Tadpoles of (A) _Hyla boulengeri_ (KU 104295) + and (B) _Hyla elaeochroa_ (KU 104134), × 3.] + + [Illustration: Fig. 3. Mouthparts of tadpoles of (A) _Hyla + boulengeri_ (KU 104295) and (B) _Hyla elaeochroa_ (KU 104134), × 25.] + +_Remarks._--Cope (1887:12) described _Scytopis boulengeri_ from +Nicaragua. Günther (1901:267) placed _boulengeri_ in the genus _Hyla_, +and stated that Cope possibly placed _boulengeri_ in the genus +_Scytopis_ on the supposition that it had an accumulation of +"sebaceous glands" above the tympanum. Noble (1918:339) redescribed +_Hyla boulengeri_ on the basis of three specimens from Zelaya +Province, Nicaragua, and noted that the glands were not prominent in +any of the specimens. Duellman (1956:8) showed that _Scytopis hebes_ +(generotype of _Scytopis_ by monotypy) is a Phrynohyas, and thus +placed _Scytopis_ Cope, 1862, in the synonymy of _Phrynohyas_ +Fitzinger, 1843. + +Dunn and Emlen (1932:25) placed _Hyla lancasteri_ Barbour in the +synonymy of _Hyla boulengeri_; the former was known solely from one +juvenile. They made no qualifying statements, but probably they were +impressed by the strongly barred thighs, a coloration known among +Central American hylids at that time only in _Hyla boulengeri_ +(Duellman, 1966a:271). Taylor (1952:856) followed Dunn and Emlen with +reservation and noted some differences. Duellman (1966a:271) showed +that the holotype of _lancasteri_ was a juvenile of a species +subsequently named as _Hyla moraviaensis_ by Taylor (1952:865). + +In Central America, _Hyla boulengeri_ can be confused only with _Hyla +foliamorta;_ the latter is restricted to central and eastern Panamá +and northern Colombia. The snout of _foliamorta_ is more pointed and +protruding, and the vocal sac is darker than in _boulengeri_; the +groin of _foliamorta_ usually is creamy white, whereas _boulengeri_ +usually has a dark spot. The skulls differ in that _boulengeri_ has a +frontoparietal fontanelle, the prevomer is larger and elongate, +anteriorly connected to the premaxillary, and posteriorly separated +from the sphenethmoid by cartilage; _foliamorta_ lacks a fontanelle, +the prevomer is smaller, anteriorly separated from the premaxillary by +cartilage, but connected by a bony suture to the sphenethmoid. The +mating call of _boulengeri_ differs by having shorter notes, twice as +many pulses per second, a higher fundamental frequency, and more +closely approximated major frequencies than does that of _foliamorta_. + +_Hyla boulengeri_ need not be compared in detail with the other Central +American members of the _Hyla rubra_ group, because all of them are +smaller and have shorter snouts, smoother skin, and dissimilar color +patterns. + +_Distribution._--In Central America _Hyla boulengeri_ inhabits the +forested lowlands in locally humid areas in Guanacaste Province, Costa +Rica, and in the humid Golfo Dulce region of Costa Rica; it occurs on +the Carribbean lowlands from central Nicaragua to South America, where +it ranges to Guyana and Ecuador. The highest elevations where _H. +boulengeri_ has been found are 620 meters at Turrialba, Cartago +Province, and 700 meters at Tilarán, Guanacaste Province, Costa Rica +(Fig. 4). + +_Specimens Examined._--Costa Rica: _Alajuela_: 9 km N Ciudad Quesada, +near La Florencia, USC 8059 (4); 18 km N Florencia, USC 2624; Laguna +Monte Alegre, KU 64334; Las Playuelas, 11 km S Los Chiles, USC 7216, +7217 (2), 7219; 3 km NE Muelle del Arenal, USC 2644 (5). _Cartago_: +Turrialba, KU 24741. _Guanacaste_: 7 km N Liberia, USC 8096 (2), 8138 +(6); 13.6 km N Liberia, USC 8151, 8171 (2); 20.5 km S Liberia, USC +8205; Taboga, 20 km SE Las Cañas, KU 102170, USC 7166; 4 km NE Tilarán, +USC 8023; 6 km NE Tilarán, USC 523 (3), 6262, 7019. _Heredia_: Puerto +Viejo, KU 64323-7 (skeletons), 104351-3 (skeletons), 64330-3, +103592-620; 1 km NE Puerto Viejo, UMMZ 126042; 1 km S Puerto Viejo, KU +84983-4 (skeletons), 86317-22, 87774 (skeleton); 4.2 km W Puerto Viejo, +KU 64329, 64328 (skeleton). _Limón_: Mountain Cow Creek, near Banano, +KU 37031, 41067 (skeleton); 3 km S Río Tortuguero, AMNH 69057; Suretka, +KU 36482-8, 36699. _Puntarenas_: 4.8 km S Bahía Rincón on NW side Río +Rincón, USC 705; Parrita, USC 6163; 4.5 km W Rincón de Osa, KU +102177-9, 104295-6 (tadpoles); 6 km SW Rincón de Osa, KU 102171-6; 4.4 +km NW Villa Neilly, USC 8003; 10.5 km WNW Villa Neilly, KU 64321. _San +José_: 21 km WSW San Isidro el General, KU 34104-6. + + [Illustration: Fig. 4. Map showing locality records for _Hyla + boulengeri_ (circles) and _H. foliamorta_ (dots).] + +Panamá: _Bocas del Toro_: 3.2 km W Almirante, KU 95978. _Canal Zone_: +Barro Colorado Island, FMNH 13379; near Clayton Reservation, UIMNH +42000; 2.6 km SW Fort Kobbe, KU 95977; Miraflores Locks, AMNH 69764-5; +Summit, AMNH 73445, KU 97777, 101540-9, 104350 (skeleton). _Colón_: Río +Gatuncillo, near Nuevo San Juan, KU 95976. _Darién_: El Real, KU +80451-3. + + +_Hyla foliamorta_ Fouquette + + _Hyla foliamorta_ Fouquette, Herpetologica, 14:125, April 25, + 1958 [Holotype.--TNHC 23109, 11 km. NW Miraflores Locks, Canal + Zone, Panamá; M. J. Fouquette, Jr. collector]. + +_Diagnosis._--Size medium (Male to 43 mm., Female to 41 mm.); skull +longer than wide; frontoparietal fontanelle absent; snout acuminate, +projecting; interorbital triangle bordered by white lines; scapular +region having two or more elongate spots; dorsum smooth; vocal sac +dark gray; groin creamy white; traces of web between fingers. + +_Description._--Head flattened, longer than wide; snout flat, pointed, +protruding beyond lower lip; loreal region slightly concave; canthus +moderately prominent; eyes smaller than interorbital space; tympanum +distinct, 55 to 75 per cent of diameter of eye, smaller than +internarial space; arms short; fingers having rudimentary webs; median +palmar tubercle tripartite; inner palmar tubercle on base of first +finger flat; subarticular tubercles distinct; discs of fingers smaller +than diameter of tympanum; legs long; tarsal fold lacking; inner +metatarsal tubercle larger than outer; one phalanx free on second, +third, and fifth toes, two and one half phalanges free on fourth toe; +narrow fringe continuing from web to discs of toes; discs of toes +about the size of those on fingers; skin smooth on dorsum and flanks, +that on belly and posterior part of thighs granular; tongue oval, +longer than wide; vocal slits oblique, about one half length of +tongue. + +In life, dorsum pale tan to pale reddish brown with irregular reddish +brown markings; small dark spots on head; distinct dark brown +triangular mark between eyes, bordered by thin white lines; apex of +triangle always directed backward; supratympanic fold with black edge; +scapular region having two to five small, elongate black spots; belly +creamy tan with small brown spots; vocal sac uniformly dark brown with +scattered creamy tan flecks; upper jaw dark brown; limbs creamy white +below with scattered brown spots; groin marked with small brown spots +in some specimens; anterior and posterior surfaces of thighs +yellow-orange with three distinct black blotches; two dark bands on +upper surface of shanks; webbing of feet yellowish tan with brown +mottlings (Pl. 1B). + +In preservative, dorsum brown or gray with darker markings; interorbital +triangle distinct, bordered by white lines; supratympanic fold with +black edge; two or more small elongate black spots in scapular region; +belly white with numerous brown flecks; edge of upper lip dark brown; +vocal sac dark gray; undersides of limbs creamy white; groin creamy +white with or without brown spots; anterior and posterior surfaces of +thighs having three black blotches separated by creamy white spaces; +shanks having two brown bands; webbing of feet mottled with brown. + +_Variation._--Twenty-eight breeding males from the area between Chepo +and Tocumen, Panamá, have snout-vent lengths of 39.0 mm. to 46.0 mm. +(mean 42.5 mm.). In these specimens, the ratio of the tibia length +to the snout-vent length is 0.54 to 0.61 (mean, 0.57); the ratio +of the diameter of the tympanum to that of the eye is 0.55 to 0.75 +(mean, 0.67). One female has a snout-vent length of 41.0 mm., +tibia/snout-vent length ratio of 0.57, and tympanum/eye ratio of 0.76. +Two to five (usually three) elongate black spots are present in the +scapular region in different individuals. The flanks in some are +spotted with brown; in others they are creamy white. A small black +spot is present in the groin of some specimens. Usually two to four +blotches are present on the anterior and posterior surfaces of the +thighs; in some specimens the blotches are reduced to small spots. One +or two brown spots are present proximally on the shanks in most +specimens. In some individuals tuberculations are scattered on the +head and in the tympanic and scapular regions, but the dorsum is +smooth in most specimens; the belly is creamy white flecked with +brown. + +_Cranial Osteology._--The skull of _Hyla foliamorta_ is flat and +longer than it is wide. The premaxillary is small and bears 13 to 16 +teeth (mean for 2 specimens, 14.8). The alary process of the +premaxillary is vertical and concave posteriorly. Ventrally, the +premaxillary is completely separated from the prevomer by cartilage. +The maxillary is slender; each bears 77 to 84 teeth (mean for 2 +specimens, 81). The pars facialis of the maxillary is laterally convex +and less than three times the height of the pars dentalis. + +The nasal is large and pointed anteriorly and posteriorly in dorsal +view. The length of the nasal comprises about 40 per cent of the total +length of the skull. The nasals are separated anteromedially by the +cartilaginous septum nasi. One protuberance is present on the +midlateral concavity of the nasal. Posteriorly, the nasal overlaps the +sphenethmoid; posterolaterally the nasal articulates with the +palatine. The sphenethmoid is completely ossified and pentagonal in +dorsal view. The frontoparietal is elongate, without a pronounced +anterior supraorbital process. The frontoparientals are sutured +medially throughout their lengths; the frontoparietal fontanelle is +absent. + +The bony part of the proötic is narrowly separated dorsolaterally from +the squamosal by the cartilaginous crista parotica. The squamosal is +large; the anterior arm is pointed. The posterior arm of the squamosal +is broad, rounded terminally, and articulates with the proötic +medially. + +The prevomer is short and separated anteriorly from the premaxillary +and maxillary by cartilage. The posterior margin of the prevomer has a +bony articulation with the sphenethmoid. Each prevomer bears five to +seven teeth. The palatine is small and edentate. The anterior end of +the parasphenoid is narrow (more pointed than in _Hyla boulengeri_). +The pterygoid is slender and well developed (Fig. 5A). + + [Illustration: Fig. 5. Dorsal views of the skulls of (A) _Hyla + foliamorta_ (KU 77687) and (B) _H. elaeochroa_ (KU 68289), × 3.] + +_Natural History._--_Hyla foliamorta_ inhabits lowland forests in +eastern Panamá and breeds in temporary ponds. Males have been observed +calling from grasses, bushes, and emergent vegetation near temporary +ponds and ditches along roads. William E. Duellman informed me that he +found a breeding congregation of this species in June near Chepo, +Panamá, where males were calling from spiny palms at the edge of a +woodland pond. Fouquette (1958) found calling males in May, August, +and September near Miraflores Locks, Canal Zone. Calling stations vary +from one to two meters above ground. No clasping pairs have been +found; only one female is known (KU 101589, from 8 km NE Tocumen, +Panamá); this gravid individual was collected in early June. + +The mating call of _Hyla foliamorta_ consists of one pulsed, +low-pitched, moderate trill of about O.5 second duration. Each note is +repeated at intervals of 5 seconds to a few minutes. The notes have +about 50 pulses per second, a fundamental frequency of 56 cycles per +second and a dominant frequency of about 3000 cycles per second (Table +2, Pl. 3B). + +Egg deposition sites are unknown. No information is available +concerning early development, and little is known about the breeding +season of _Hyla foliamorta_. Probably its breeding activities are +restricted to the rainy months. + +_Tadpoles._--Eight tadpoles were collected from a weedy temporary pond +near Chepo, Panamá, in early June. + +A typical tadpole in stage 35 of development (KU 104244) has a body +length of 9.5 mm., tail length of 25.0 mm., and a total length of 34.5 +mm. Other characters are as follows: depth of tail equal to length of +body; body deeper than wide; distance between eye and nostril equal to +distance between eye and spiracle; mouth anteroventral; median part of +upper lip bare; rest of lip having one row of papillae; a few other +rows of small papillae at corners of mouth; tooth rows 2/3; first +upper row entire, second upper row interrupted medially, shorter than +first; lower rows shorter than upper rows, third shortest; beak +moderately robust; spiracle nearer eye than anus; anal tube short, +aperture not extending to border of ventral fin; caudal musculature +slender, extending to tip of pointed tail; dorsal fin extending onto +body (Table 4). + +TABLE 4.--Sizes of Tadpoles of _Hyla foliamorta_ in Relation to +Developmental Stages. (Means in parentheses below observed ranges; +measurements in mm.) + + ======================================================= + Stage | N | Body length | Tail length | Total length + --------+---+-------------+-------------+-------------- + 25 | 2 | 5.0-5.2 | 8.0-8.5 | 13.0-13.7 + | | (5.1) | (8.3) | (13.4) + | | | | + 26 | 3 | 7.0-7.5 | 12.0-12.4 | 17.0-19.5 + | | (7.2) | (12.1) | (18.6) + | | | | + 28 | 2 | 6.5-7.0 | 18.0 | 25.0 + | | (6.8) | | + | | | | + 35 | 1 | 9.5 | 25.0 | 34.5 + +In life, yellow above, white below; caudal fin greenish yellow with +black or gray reticulations; dark line from snout to eye; dark spot +behind eye; tail unpigmented except for fine dark reticulations. In +preservative body creamy white, transparent below with dark pigment +above in some specimens. + +_Remarks._--_Hyla foliamorta_ can be confused only with _Hyla +boulengeri_. The differences between adults of these species were +discussed in _Remarks_ on _H. boulengeri_. The tadpoles of +_foliamorta_ have labial papillae on the lower lip and a stripe +between the eye and the tip of the snout. By comparison the tadpoles +of _boulengeri_ have a bare lower lip and no stripe between the eye +and the tip of the snout. + +_Distribution._--_Hyla foliamorta_ inhabits the subhumid Pacific +lowlands (elevations of less than 100 meters) of Central Panamá and +Caribbean lowlands of northern Colombia (Fig. 4). + +_Specimens Examined._--Panamá: _Panamá_: 3 km WSW Chepo, KU 77164-9, +101573-5, 104243-4 (tadpoles); 6 km WSW Chepo, KU 77170, 101576-8; 1.5 +km SW Naranjal, KU 77171, 77687 (skeleton); 2 km N Tocumen, KU +101579-83, 104349 (skeleton); 8 km NE Tocumen, KU 101584-92. + +No specific locality: TNHC 24401. + + +_Hyla rubra_ Laurenti + + _Hyla rubra_ Laurenti, Synopsis Reptilium Emendatum, p. 35, 1768. + Daudin, Hist. Nat. Rainettes Grenouilles Crapauds, II:26, 1802. + Daudin, Hist. Nat. Particuliere Reptiles, 8:53, 1803. Günther, + Catalogue Batrachia Salientia Brit. Mus., p. 110, 1859. Boulenger, + Catalogue Batrachia Salientia s. Ecaudata, p. 403, February 1, + 1882. Dunn, Occas. Papers, Boston Soc. Nat. Hist., 5:413, October + 10, 1931. + + _Hyla elaeochroa_ (part): Dunn and Emlen, Proc. Acad. Nat. Sci. + Philadelphia, 84:25, March 22, 1932. + +_Diagnosis._--Size medium; skull longer than wide; frontoparietal +fontanelle absent in adults; snout subovoid; choanae rounded; dorsal +stripes present; black vermiculations on posterior surfaces of thighs. + +_Description._--Head flat, longer than wide; snout long, subovoid, +slightly protruding beyond lower lip; loreal oblique, concave; canthus +rounded, indistinct; diameter of eye about equal to interorbital +space; tympanum large, about three fifths diameter of eye, smaller +than internarial distance; supratympanic fold indistinct; arms short; +fingers free of webs; subarticular tubercles distinct; median palmar +tubercle large, bifid; inner palmar tubercle on base of first finger +flat, elongate; disc of third finger about one half diameter of +tympanum; legs moderately long; tarsal fold absent; inner metatarsal +tubercle distinct, oval; toes about half webbed; web on fourth toe +extending to disc; discs of toes about size of those on fingers; skin +smooth above with small granules on head and in scapular region in +some specimens; skin on flanks, throat, belly, and lower surfaces of +thighs granular; tongue oval, longer than wide, not free behind; +choanae small, oval; vocal slits long, lateral to tongue. + +In preservative, dorsum pale brown with darker dorsolateral stripes; +narrow dark brown line from nostril to eye; groin, anterior surface of +thighs, and posteroventral surfaces of shanks creamy tan with dark +brown vermiculations; white spots present on thighs in some specimens; +throat flecked with brown; belly creamy white or gray. + +_Remarks._--The taxonomic history of _Hyla rubra_ Laurenti is +confused. Seba (1734:70) illustrated and diagnosed a frog for which +he used the name _Ranula, Americana, Rubra_. Linnaeus (1758:213) +considered Seba's frog to be a variety of _Hyla arborea_. Laurenti +(1768:35) apparently examined the same individual that Seba called +_Ranula, Americana, Rubra_. For this specimen, Laurenti used the +binominal _Hyla rubra_ and provided a brief diagnosis. The type +locality was given as America. + +Daudin (1802:26) redescribed the same specimen(s?) treated by Seba and +Laurenti and provided a fairly good description and figures. Daudin +restricted the type locality to Surinam and indicated that Marin de +Baize was the probable collector. Daudin (1802:26 and 1803:53) +neglected to consider Laurenti's work, but he applied the same name +used by Laurenti. Most authors have credited _Hyla rubra_ to Daudin, +but Rivero (1961:120) noted that _Hyla rubra_ Laurenti, 1768, has +priority over _Hyla rubra_ Daudin, 1802. Since both Laurenti and +Daudin worked on Seba's material, it is reasonable to assume that +Daudin redescribed the same frog that was named by Laurenti; this was +not an uncommon practice in the early nineteenth century. Thus I +conclude that _Hyla rubra_ Daudin, 1802, is a junior synonym of _Hyla +rubra_ Laurenti, 1768. + +Dunn (1931a:413) first reported _Hyla rubra_ from Central America; he +recorded the species from the Canal Zone and San Pablo, Panamá. I have +examined the material of _Hyla rubra_ from Panamá deposited in various +museums. Most of the specimens are faded, discolored, and do not have +distinct brown vermiculations on the thighs. The specimens seem to be +more like _Hyla rubra_ than any of the other species in the _rubra_ +group. The presence of oval choanae and a tympanum larger than the +largest finger disc separate these specimens from _Hyla elaeochroa_, a +species with which _rubra_ has been confused. _Hyla elaeochroa_ does +not occur in the Canal Zone or eastern Panamá. All museum specimens +from Nicaragua, Costa Rica, and western Panamá that have been called +_Hyla rubra_, plus those mentioned by Dunn and Emlen (1932:25) and +Dunn (1933:61) are _Hyla elaeochroa_. + +The taxonomic status of the many South American populations referred +to _Hyla rubra_ and of other populations now recognized as different +species is not clear at the present time. Considerable variation in +external characters and in cranial features has been observed in South +American _rubra_. A review of the taxonomy of these populations is +beyond the scope of this paper. Possibly the Central American +specimens herein referred to _rubra_ will ultimately be found to be +specifically distinct from those in Surinam. Since I have no +osteological material from Central America, I have been unable to +describe the cranium in this account. Furthermore, I have no data on +the ecology and life history of _rubra_ in Central America. + +_Distribution._--_Hyla rubra_ inhabits lowland tropical forests from +central-eastern Panamá to northern South America and thence through +lowlands east of the Andes to northern Argentina (Fig. 6). + +_Specimens Examined._--Panamá: _Canal Zone_: Gatun, UMMZ 52720 (2); +Madden Dam, FMNH 67820; no specific locality, UMMZ 56517 (3), USNM +37863. _Colón_: Cerro Bruja, MCZ 13248. _Darién_: El Real, USNM +140569-70, 140573. _Panamá_: Juan Díaz, MCZ 17973; Las Sabanas, MCZ +17581; Río Trinidad, UMMZ 64003; San Pablo, MCZ 1398-9. + + +_Hyla elaeochroa_ Cope + + _Hyla elaeochroa_ Cope, Jour. Acad. Nat. Sci. Philadelphia, 8:105, + 1876 [Holotype.--USNM 30689, Sipurio, Limón Province, Costa Rica; + William M. Gabb collector]. Günther, Biologia Centrali-Americana, + Reptilia and Batrachia, p. 265, June 1901. Taylor, Univ. Kansas + Sci. Bull., 35:859, July 1, 1952. Duellman, Univ. Kansas Publ., + Mus. Nat. Hist., 17:270, June 17, 1966. + + _Hyla quinquevittata_ Cope, Proc. Amer. Philos. Soc., 23:273, + April 1887 [Holotype.--USNM 14187, Nicaragua; J. F. Bransford + collector]. Günther, Biologia Centrali-Americana, Reptilia and + Batrachia, p. 268, June 1901. Noble, Bull. Amer. Mus. Nat. Hist., + 38:340, June 1918. + + _Hyla rubra_ (part): Dunn and Emlen, Proc. Acad. Nat. Sci. + Philadelphia, 84:25, March 22, 1932. + + _Hyla dulcensis_ Taylor, Univ. Kansas Sci. Bull., 39:37, November + 18, 1958 [Holotype.--KU 32168, Golfito, Puntarenas Province, Costa + Rica; Edward H. Taylor collector]. + +_Diagnosis._--Size medium (Male to 38 mm., Female to 40 mm.); skull +wider than long; nasals truncate anteriorly; frontoparietal fontanelle +moderate in size; snout slightly protruding; tympanum about size of +largest discs on fingers; dorsum marked by longitudinal stripes; dark +stripe between eye and nostril; in life tan to olive-green with or +without dark mark between eyes; bones greenish blue. + +_Description._--Head flat, longer than wide; snout long, rounded, +protruding beyond mouth; canthus indistinct; length of eye equal to +interorbital distance; loreal region not pronounced; tympanum distinct +and about two-fifths diameter of eye; interorbital triangle present or +absent; arms short; trace of web between fingers, extending as fringe +along sides of fingers; first finger very short with small disc; other +discs about size of those on toes; discs on third finger and fourth +toe as large as tympanum; outer palmar tubercle moderate in size, +partly bifid; inner palmar tubercle large, elongate, flat; +subarticular tubercles distinct; legs moderately long; tarsal fold +absent; inner metatarsal tubercle flat; outer metatarsal tubercle +smaller, indistinct; subarticular tubercles moderate in size; fringe +on toes to tip of disc of second toe; rest of toes about two-thirds +webbed; foot length about two fifths snout-vent length; tibia length +about one half snout-vent length; skin above smooth or with minute +pustules; belly finely granular; ventral surfaces of thighs and areas +below anus granular; skin on ventral surfaces of limbs smooth; tongue +relatively large, longer than wide, barely notched behind; vocal slits +elongate, lateral to tongue; choanae medium in size. In life, dorsum +yellowish brown, olive green, or grayish brown with dark brown spots +on snout, dark brown stripe from nostril to eye, dark yellow-brown +interorbital triangle, and dark supratympanic region; generally five +interrupted longitudinal dark brown stripes on dorsum (one on each +flank, pair of paravertebral and one vertebral); flanks pale yellow; +groin yellowish brown; thighs marked with one or two transverse +yellow-brown blotches; shanks with two or three yellow-brown blotches +above; spaces between blotches on thighs, shanks, tarsi, and feet +yellow; brown spots on tarsi and in some specimens on feet; arm pale +yellow with pale brown spots; belly creamy white having slight +blue-green tint; vocal sac and chin yellow; axillary region yellow, +blue-green in some specimens (Pl. 2A). + +In preservative, head and dorsum yellowish brown; dark brown stripe +from nostril to eye; dark brown spots on snout; a dark brown +interorbital triangle with apex directed backward; dark brown +supratympanic region; dorsal stripes same as in living individuals; +flanks pale yellow with brown spots in some specimens; groin creamy +white; thighs and shanks having or lacking transverse dark brown +blotches; spaces between blotches creamy white or yellow-brown; arms +pale yellowish brown; belly and vocal sac creamy white. + +_Variation._--Geographic variation in size and some proportions, such +as the ratio of tibia length to snout-vent length and the ratio of the +diameter of the tympanum to that of the eye, have been observed in +this species. The largest individuals are from the Golfo Dulce region +(samples from Piedras Blancas and Rincón de Osa), Puntarenas Province, +Costa Rica. The smallest individuals are from El Recreo, Zelaya +Province, Nicaragua, and from the Caribbean lowlands of Costa Rica. + +The diameter of the tympanum is proportionately larger (relative to +the size of the eye) in males from Tilarán, Guanacaste Province; the +tympanum is nearly as large in males from Piedras Blancas, Puntarenas +Province, and Puerto Viejo, Heredia Province, Costa Rica. The lowest +ratios occur in individuals from Almirante, Bocas del Toro, Panamá, in +specimens from the Caribbean lowlands of Costa Rica (except Puerto +Viejo), and in those from El Recreo, Zelaya Province, Nicaragua. In +general, the tympanum is proportionately larger in females than in +males; the tympanum is largest in females from the Pacific lowlands +of Costa Rica (Table 5). + +Color variation has been observed in individuals from the same +population, as well as in individuals from different localities, +between males and females, and from night to day. In life, most +individuals from the Pacific lowlands of Costa Rica are dark tan to +greenish gray above with dark brown longitudinal stripes that are +entire or broken, but some specimens (mostly males) are dusky brown +and lack longitudinal stripes or an interorbital triangle; females +usually have the dark interorbital triangle and the stripes on the +dorsum. Individuals from Turrialba, Cartago Province, Costa Rica, are +pale olive-tan with olive-brown markings. Individuals from Puerto +Viejo, Heredia Province, Costa Rica, are uniformly yellowish brown +with or without dark longitudinal stripes. Specimens from El Recreo, +Zelaya Province, Nicaragua, are like those from Puerto Viejo. Males +from Almirante, Bocas del Toro, Panamá, are pale brown with dark brown +longitudinal stripes and an indistinct interorbital triangle. Females +have a distinct interorbital triangle and dark brown blotches on the +thighs and shanks. + +By night, the dorsum usually is pale yellow, and the belly is creamy +white. By day, the dorsum is dark tan; the stripes and spots are +darker, and the belly is yellowish white. Taylor (1952) noticed that +considerable variation in color pattern occurred from night to day in +individuals from Turrialba, Cartago Province, Costa Rica. At night +some individuals lacked a dorsal pattern, but by day many of these +individuals developed dorsal stripes. + +_Cranial Osteology._--The skull of _Hyla elaeochroa_ is slightly wider +than it is long, and flat. The premaxillary is small and bears 10 to +15 teeth (mean for 9 specimens, 12.3). The alary process of the +premaxillary is small, vertical, and slightly concave posteriorly. +Ventrally, the premaxillary is partially united to the prevomer by +ossification. The maxillary is slender and bears 70 to 82 teeth (mean +for 9 specimens, 74.3). The pars facialis of the maxillary is +laterally convex and is about twice as high as the pars dentalis. + +The nasal is large, robust, anteriorly truncate, but pointed +posteriorly in dorsal view. The nasal comprises about 45 per cent of +the total length of the skull. There is an anterior cartilaginous +septum nasi separating the two nasals; the latter overlap the +sphenethmoid posteriorly. Each nasal bears a shallow concavity in the +midlateral side and lacks a maxillary process. Dorsally, the +sphenethmoid is wider than long, roughly pentagonal in shape; the +frontoparietal is elongate, smooth, and bears a small anterior +supraorbital process. The sphenethmoid and frontoparietal form the +anterior margin of the frontoparietal fontanelle; the fontanelle is +narrow anteriorly and wider posteriorly (Fig. 5B). + +The entire distal surface of the proötic is in contact with the +posterior arm of the squamosal. A narrow cartilaginous crista parotica +is visible dorsally in some specimens. The squamosal is broad +posteriorly but its anterior arm is slender and not in contact with +the maxillary. + +TABLE 5.--Geographic Variation in Size and Proportions in Males of +_Hyla elaeochroa_. (Means in parentheses below observed ranges.) + +========================================================================== + | |Snout-vent| Tibia | | + | | length | length/ |Tympanum/|Foot length/ +Locality | N | (mm.) | snout-vent | eye | snout-vent +---------------------+----+----------+------------+---------+------------- +Nicaragua: El Recreo | 9 | 28.0-30.3| 0.51-0.57 |0.47-0.59| 0.39-0.54 + | | (29.3) | (0.55) | (0.51) | (0.41) + | | | | | +Costa Rica: Tilarán | 21 | 28.8-33.6| 0.47-0.57 |0.48-0.65| 0.40-0.46 + | | (30.6) | (0.52) | (0.59) | (0.41) + | | | | | +Costa Rica: Puerto | 22 | 26.3-32.4| 0.49-0.54 |0.48-0.65| 0.38-0.45 + Viejo | | (29.7) | (0.52) | (0.57) | (0.42) + | | | | | +Costa Rica: Turrialba| 95 | 28.1-35.0| 0.47-0.56 |0.47-0.68| 0.37-0.46 + | | (30.6) | (0.51) | (0.56) | (0.41) + | | | | | +Costa Rica: Bataán, | 26 | 26.3-32.7| 0.47-0.54 |0.45-0.66| 0.36-0.44 + Limón, and Suretka | | (30.0) | (0.51) | (0.50) | (0.41) + | | | | | +Costa Rica: Piedras | 21 | 33.3-37.7| 0.50-0.54 |0.48-0.64| 0.40-0.46 + Blancas | | (35.2) | (0.51) | (0.57) | (0.43) + | | | | | +Costa Rica: Rincón de| 24 | 31.4-35.9| 0.50-0.56 |0.45-0.61| 0.40-0.46 + Osa | | (34.1) | (0.53) | (0.54) | (0.43) + | | | | | +Panamá: Bocas del | 6 | 31.0-33.5| 0.49-0.54 |0.47-0.50| 0.41-0.43 + Toro | | (32.1) | (0.51) | (0.48) | (0.42) + + + [Illustration: PLATE 1 + + Living _Hyla_: (A) _H. boulengeri_ (KU 86322) and (B) _H. + foliamorta_ (KU 101576), × 2.] + + [Illustration: PLATE 2 + + Living _Hyla:_ (A) _H. elaeochroa_ (KU 91688), (B) _H. staufferi + staufferi_ (KU 57791), and (C) _H. staufferi altae_ (KU 101688), × + 2.] + + [Illustration: PLATE 3 + + Audiospectrograms and sections of mating calls of (A) _Hyla + boulengeri_ (KU Tape No. 511) and (B) _H. foliamorta_ (KU Tape No. + 511) and (B) _H. foliamorta_ (KU Tape No. 288).] + + [Illustration: PLATE 4 + + Audiospectrograms and sections of mating calls of (A) _Hyla + elaeochroa_ (KU Tape No. 97), (B) _H. s. staufferi_ (KU Tape No. + 93), and (C) _H. staufferi altae_ (KU Tape No. 502).] + +The prevomer is short, and broadest anteriorly. The prevomer is joined +to the premaxillary by ossification. The posterior margin of the +prevomer bears a narrow cartilaginous articulation with the +sphenethmoid. The anterolateral and posterolateral processes of the +prevomer form an incomplete bony margin to the small choanae; each +prevomer bears four to seven teeth. The palatine is small, curved +anteriorly and edentate. The anterior part of the parasphenoid is +robust and ends in a point. The pterygoid is slender and weakly +developed. + +_Natural History._--_Hyla elaeochroa_ inhabits humid lowland tropical +forests in lower Central America and breeds in temporary ponds. +Clasping pairs, gravid females, and calling males have been found +mostly in June, July, and August. William E. Duellman informed me that +he also found males calling in mid-February, late April, and May. +Duellman (1967) reported detailed observations of the social +organization in the mating call of _Hyla elaeochroa_. The choruses in +this species are initially organized, but when many individuals call, +the chorus loses organization. I observed this species breeding in a +temporary pond at Puerto Viejo, Heredia Province, Cost Rica, in late +June. Calling males and clasping pairs were extremely abundant within a +few hours after a heavy rain. Males were mostly found calling from low +emergent herbs in the pond and less commonly from bushes and trees to +heights of six meters above the water. Calling males were also observed +at Ricón de Osa, Puntarenas Province, Costa Rica, in late July. These +breeding individuals were found in a shallow pond at the edge of a wet +forest. Calling stations were less than two meters in height. John D. +Lynch informed me that after a heavy rain in early August, he found +several hundred individuals congregated in a small grassy pond less +than a foot deep, at Rincón de Osa. Males were calling from sites on +grass stems a few centimeters above the water. + +The mating call of _Hyla elaeochroa_ consists of short notes, repeated +at intervals of about 0.40 second. Each note has a duration of 0.12 to +0.24 second. The fundamental frequency varies from 48 to 65 cycles per +second, and the notes have 40-50 pulses per second; the dominant +frequency is at about 2,900 cycles per second (Table 2, Pl. 4A). + +The eggs are deposited in a mass in the water near floating vegetation. +William E. Duellman informed me that he observed hatchlings oriented +vertically with the tip of the mouth at the surface of the water. They +gradually sank to bottom, but swam back to surface again. No additional +information is available concerning early development. Tadpoles have +been found in shallow grassy ponds in clearings and in temporary +woodland ponds. + +_Tadpoles._--Three hundred and thirty-one tadpoles in various stages of +development are available. Thirty-five tadpoles in stage 35 have a mean +body length of 8.1 mm. (8.0-9.0 mm.), tail length of 17.7 mm. +(15.0-19.5 mm.), and total length of 25.9 mm. (23.0-27.5 mm.). The +largest tadpole examined is in stage 40 and has a total length of 34.5 +mm. (Table 6). + +A typical tadpole, stage 35 of development (KU 104134, from Puerto +Viejo, Heredia Province, Costa Rica), has a body length of 9.1 mm., +tail length of 17.7 mm., and a total length of 26.8 mm. Other +characters are as follows: body depressed anteriorly; body length +greater than depth of tail; internarial space as broad as interorbital +distance; nostril equidistant between eye and tip of snout; eyes +moderately large; mouth anteroventral and triangular; median fourth of +upper lip bare; rest of lip bordered by one row of papillae; clumps of +small papillae at corners of mouth; tooth rows 2/3; upper rows equal in +length; second row interrupted medially; lower rows shorter than upper +rows, diminishing in length; beak rather weak with small serrations; +spiracle short and nearer eyes than anus; anal opening not reaching +edge of ventral fin; caudal musculature attenuated distally (Figs. 2B +and 3B). + +TABLE 6.--Sizes of Tadpoles of _Hyla elaeochroa_ in Relation to +Developmental Stages. (Means in parentheses below observed ranges; +measurements in mm.) + + ------+---+-------------+-------------+-------------- + Stage | N | Body length | Tail length | Total length + ------+---+-------------+-------------+-------------- + 24 | 2 | 4.0-4.0 | 8.5-9.0 | 12.5-13.0 + | | (4.0) | (8.8) | (12.8) + | | | | + 25 |64 | 5.0-6.5 | 8.5-15.0 | 13.5-21.5 + | | (5.7) | (11.8) | (17.6) + | | | | + 27 |30 | 7.0-7.5 | 13.0-16.0 | 20.0-23.0 + | | (7.1) | (14.2) | (21.3) + | | | | + 30 |15 | 7.0-8.0 | 13.0-16.5 | 20.0-24.0 + | | (7.3) | (15.0) | (22.4) + | | | | + 32 |30 | 7.5-8.5 | 15.0-17.0 | 22.5-25.0 + | | (7.8) | (16.1) | (23.8) + | | | | + 35 |35 | 8.0-9.0 | 15.0-19.5 | 23.0-27.5 + | | (8.1) | (17.7) | (25.9) + | | | | + 37 |22 | 8.5-9.5 | 16.0-22.0 | 25.0-31.0 + | | (9.0) | (18.8) | (27.8) + | | | | + 39 |14 | 9.5-10.5 | 19.0-24.9 | 28.5-33.5 + | | (9.9) | (21.1) | (31.0) + | | | | + 40 |27 | 7.0-11.5 | 15.0-23.0 | 23.0-34.5 + | | (9.1) | (22.0) | (31.2) + | | | | + 43 |10 | 8.0-12.0 | 11.0-17.0 | 20.0-26.0 + | | (10.2) | (13.5) | (23.7) + | | | | + 45 |16 | 10.0-12.0 | 1.0-7.0 | 12.0-17.0 + | | (11.2) | (3.4) | (14.6) + | | | | + 46 |45 | 11.0-13.0 | | + | | (11.8) | | + +In life, dorsum yellowish tan with gray-brown mottling; belly and +ventrolateral surfaces silvery-gold or white; black stripe from tip of +snout to eye; two black blotches below eye, another blotch extending +from eye to base of caudal musculature; caudal musculature and fins +gray-brown. In preservative, yellowish tan and silvery-gold colors +lost; black reticulations present on tail. + +_Remarks._--Cope (1876:105) described _Hyla elaeochroa_ from Sipurio, +Limón Province, Costa Rica. He based his description on a small +specimen, 26.0 mm. in snout-vent length, having a dorsum uniformly +colored and lacking an interorbital triangle and blotches on the +thighs. Cope (1887) described pigmented specimens from Nicaragua as +_Hyla quinquevittata_, which he diagnosed as having dark brown bars on +the hind limbs and five dark brown longitudinal stripes on the dorsum, +the median one of which was expanded anteriorly so as to form a large +triangular spot between the eyes. He thought this species was related +to _Hyla eximia_ Baird and noted that "the hinder legs are much larger; +the muzzle is more acuminate and the color bands are much wider" than +in _eximia_. Cope did not compare _quinquevittata_ with _elaeochroa_, +which he had described ten years before. Günther (1901:268), Noble +(1918:340), and Nieden (1923:251) regarded both _elaeochroa_ and +_quinquevittata_ as valid species. Dunn and Emlen (1932:25) regarded +both as synonyms of _Hyla rubra_, but they made no qualifying +statements. Taylor (1952:859) placed _quinquevittata_ as a synonym of +_elaeochroa_ and indicated that _rubra_ was another species. + +Taylor (1958:37) described _Hyla dulcensis_ from the humid tropical +forests of Golfo Dulce, Puntarenas Province, Costa Rica. He thought +this species was "related to _H. elaeochroa_ but differs in its +somewhat larger size, smaller finger and toe discs, the obsolete +canthus rostralis; the loreal region concave and the choanae larger." +Duellman (1966a:270) compared adults, tadpoles, and mating calls of +_dulcensis_ and _elaeochroa_ and concluded that a single species was +involved. + +_Hyla elaeochroa_ can be easily confused with the closely related _Hyla +staufferi_. Although the durations of the calls are similar, the call +of _elaeochroa_ has only about one third the number of pulses per +second, a much lower fundamental frequency, and a lower dominant +frequency than that of _staufferi_. _Hyla elaeochroa_ is larger and has +a less pointed snout than does _staufferi_. Although the skulls of the +two species are similar, that of _elaeochroa_ differs in having broad +palatines and comparatively larger nasals that are truncate anteriorly. +In _staufferi_ the nasal is rounded anteriorly and the palatine is +absent. + +_Distribution._--_Hyla elaeochroa_ occurs on the Caribbean lowlands +from western Panamá through Costa Rica to eastern Nicaragua, and on the +Pacific lowlands of southeastern Costa Rica and extreme western Panamá. +Most localities where it has been collected are below 800 meters, but +the species has been found at two localities above 1000 meters (El +Silencio and Pacuare, Cartago Province) on the Caribbean slopes of the +Cordillera de Talamanca, Costa Rica (Fig. 6). + +_Specimens Examined._--Nicaragua: _Zelaya_: El Recreo, UMMZ 79721 (9). + +Costa Rica: _Alajuela_: Laguna Monte Alegre, KU 64499. _Cartago_: 2 km E +Chitaría, KU 107058; El Silencio, 14.4 km NE Turrialba, KU 107059-60; +4.6 km ENE Pacuare, KU 64451-75, 64628-37; 4 km S Pavones, KU 64500; +Turrialba (Instituto Interamericano de Ciéncias Agrícolas), KU 30305-26, +24616-57, 30337-54, 31776-91, 31803, 31807-15, 64413-50, 68283-87 +(skeletons), 68390-1 (young), 35042 (eggs), 25207-8 (skeletons), 25221 +(skeleton), 41073-83 (skeletons). _Guanacaste_: 2 km E Tilarán, KU +86356-77, 87667-8 (young). _Heredia_: Puerto Viejo, KU 36696, 46466, +64501-17, 68288-91, 68387, 68388-9 (young), 91803 (young), 91688-9, +104134 (tadpoles), 104135 (young), 104354-6 (skeletons); 1.5 km N Puerto +Viejo, KU 64518-23, 68386 (tadpoles); 1 km S Puerto Viejo, KU 84985-6 +(skeletons), 87669 (young), 87772-3 (skeletons). _Limón_: Bataán, KU +30327-36; La Lola, KU 64478-98, 68281-2 (skeletons); Los Diamantes, KU +31800-02, 64476-7; Peralta, KU 31816-21; Puerto Limón, KU 31792-99; +Suretka, KU 36467-79, 36697, 41084. _Puntarenas_: 5 km NW Buenos Aires, +KU 107057; 10 km E Esparta, KU 87666 (tadpoles); Golfito, KU 32166-8; 8 +km E Palmer Norte KU 93939; 10.7 km SE Palmar Sur, KU 93938 (skeleton), +93940-51, 93952 (eggs), 93953-6 (tadpoles); Piedras Blancas, KU +103646-59; 4.5 km W Rincón de Osa, KU 102208-41, 104298 (tadpoles). + + [Illustration: Fig. 6. Map showing locality records for _Hyla + elaeochroa_ (circles) and _H. rubra_ (dots).] + +Panamá: _Bocas del Toro_: Almirante, KU 80079; Isla Bastimentos, KU +96008-11; Río Cricamola, 3.7 km from coast, KU 96012. _Chiriquí_: Río +Gariché, 8.3 km ESE Paso de Canoas, KU 101571-2. + + +_Hyla staufferi_ Cope + + _Hyla staufferi_ Cope, Proc. Acad. Nat. Sci. Philadelphia, 17:165, + October 1, 1865 [Holotype.--USNM 15317, Orizaba, Veracruz, México; + Francis Sumichrast collector]. + +_Diagnosis._--Small frogs (Male to 29 mm., Female to 31.6 mm.); skull +longer than wide; palatine absent; large cartilaginous crista parotica +present; snout flat, elongate and protruding; dark interorbital bar and +dorsal stripes usually present. + +_Description._--Head flat, especially in females, longer than wide; +snout long, protruding beyond mouth; loreal region concave; canthus +ill-defined; length of eye greater than internarial distance or width +of eyelid; length of eye less than interorbital space; tympanum +distinct; interorbital spot irregular; supratympanic fold faint; arms +short; fingers free of webs; discs on third and fourth fingers equal to +diameter of tympanum; inner metatarsal tubercle on base of first finger +distinct; first finger shorter than second; palmar tubercle distinct +(Fig. 1C); legs short (usually less than 50 per cent of snout-vent +length); tarsal fold absent; metatarsal tubercles small, outer tubercle +smaller than inner; subarticular tubercles small, simple, distinct; +toes less than half webbed (Fig. 1D); skin smooth above with a few +small pustules on head, scapular region, flanks, and supratympanic +region; arms and legs smooth; skin of belly coarsely granular; +posteroventral surfaces of thighs finely granular; tongue small, +rounded, longer than wide, slightly free and notched posteriorly; vocal +slits small, lateral to tongue; choanae moderate in size. + +_Variation._--The largest males of _Hyla staufferi_ are from Jalapa, +Guatemala, and from San Salvador, El Salvador. In these samples the +average snout-vent length is 27 mm. In Panamanian specimens the average +snout-vent length is 23.6 mm. Slight variation in the ratio of tibia +length to snout-vent length exists throughout the range; more variation +exists in the ratio of the diameter of the tympanum to that of the eye; +the tympanum is proportionately larger in northern populations (Table +7). The primary differences between Panamanian and more northern +populations are in size, color pattern on the dorsum and shanks, amount +of webbing between the toes, and duration of notes in the mating call +(Table 2, Pl. 4). + +The color in Panamanian _staufferi_ is gray or gray-brown with a pair +of distinct, complete, dark brown dorsolateral stripes, a pair of +entire paravertebral stripes, and in some specimens a vertebral stripe. +About five per cent of the individuals have interrupted stripes on the +dorsum, whereas in the more northern populations complete paravertebral +stripes are present in less ten per cent of the specimens; when +complete stripes are present, they are irregular. The dorsal ground +color in non-Panamanian specimens is brown, olive-brown, or dark brown. + +Transverse bars are present on the shanks in _Hyla staufferi_ from +Costa Rica northward to México, whereas in Panamá all the individuals +have a longitudinal stripe on the shank (Table 7, Pl. 2). The +interorbital spot or bar is more noticeable in northern populations +than in specimens from Panamá. Frogs from Costa Rica and northward have +the toes about three fourths webbed, whereas in Panamá the toes are +about two fifths webbed. The mating calls of the northern and +Panamanian populations are similar, but the notes have a longer +duration in the northern populations and a higher dominant frequency in +Panamanian populations. + +_Hyla staufferi_ is the most variable member of the _Hyla rubra_ group +in Central America. The Panamanian populations are geographically +separated from the Costa Rican and more northern populations by an area +of tropical rainforest in the Golfo Dulce region in southeastern Costa +Rica and adjacent Panamá. _Hyla staufferi_ does not occur on the +Caribbean versant of Costa Rica and Panamá. The Golfo Dulce region and +the Caribbean versant are humid and inhabited by _Hyla elaeochroa_. +_Hyla staufferi_ is an inhabitant of subhumid and xeric areas. + +On the basis of the discontinuous variation in several characters which +correlate with the disjunct distribution of the two populations, two +subspecies of _Hyla staufferi_ are recognized. The accounts that follow +apply equally to each. + +_Cranial Osteology._--The skull of _Hyla staufferi_ is flat and longer +than wide. The premaxillary is small and bears 9 to 13 teeth (mean for +5 specimens, 11.3). The alary process of the premaxillary is small, +concave posteriorly and vertical. Ventrally, the premaxillary is united +to the prevomers by partially ossified cartilage. The maxillary is +slender and usually bears 49 to 70 teeth (mean for 5 specimens, 60.7). +The pars facialis of the maxillary is convex and less than twice the +height of the pars dentalis. + +The nasal is large, rounded anteriorly, and pointed posteriorly in +dorsal view. The nasal comprises about 40 per cent of the total length +of the skull. Anteromedially the two nasals converge; posteriorly they +overlap the sphenethmoid. The nasals lack a concavity in the midlateral +surface. Dorsally, the sphenethmoid is wider than long, roughly +pentagonal in shape; the frontoparietal is elongate, narrow, and +smooth, with a small supraorbital process anteriorly. The +frontoparietal fontanelle is narrow anteriorly and wide posteriorly. + +TABLE 7.--Geographic Variation in Size and Color in Males of _Hyla +staufferi_. (Means in parentheses below observed ranges.) + + ================================================================= + | | |Complete dorsal| + | |Snout-vent | stripes |Barred shanks + Locality | N |length (mm.)| (per cent) | (per cent) + ---------------+-----+------------+---------------+-------------- + Veracruz | 47 | 23.0-27.3 | 0.0 | 100 + | | (25.4) | | + | | | | + Campeche | 20 | 24.6-27.5 | 0.0 | 100 + | | (25.5) | | + | | | | + Oaxaca | 75 | 24.0-28.7 | 9.3 | 100 + | | (26.4) | | + | | | | + Chiapas | 20 | 23.2-27.8 | 10.0 | 100 + | | (25.5) | | + | | | | + Guatemala | 22 | 25.0-29.0 | 10.9 | 100 + | | (26.9) | | + | | | | + El Salvador | 21 | 24.7-28.6 | 0.0 | 100 + | | (27.0) | | + | | | | + Honduras | 34 | 20.6-27.0 | 3.3 | 100 + | | (24.9) | | + | | | | + Nicaragua | 67 | 21.5-26.8 | 3.0 | 92.7 + | | (24.9) | | + | | | | + Costa Rica | 54 | 20.7-26.6 | 5.5 | 98.1 + | | (24.2) | | + | | | | + Total | 360 | 20.7-29.0 | 5.4 | 98.3 + Non-Panamanian | | (25.9) | | + | | | | + Panamá | 72 | 21.7-26.0 | 94.5 | 0.0 + | | (23.6) | | + +Only a narrow connection exists between the posterior, pointed arm of +the squamosal and the lateral edge of the proötic. The crista parotica +is visible dorsally along the lateral edge of the bony proötic. The +squamosal is narrow anteriorly and posteriorly. + +The prevomers are short and separated anteriorly by partly ossified +cartilage of the overlying solum nasi. The prevomer is joined to the +premaxillary by cartilage. The posterior margin of the prevomer +articulates directly with the sphenethmoid. The anterolateral and +posterolateral processes of the prevomers form the incomplete bony +internal margin of the choanae. Each prevomer bears three to six teeth. +The palatine is absent. The anterior part of the parasphenoid is narrow +and ends in a point. The pterygoid is slender and weakly developed. + +_Natural History._--Throughout its range _Hyla staufferi_ occurs in +subhumid forests and savannas; consequently, the breeding activities +are limited by the seasonal occurrence of rainfall, which accumulates +in temporary ponds where this species breeds. Clasping pairs and gravid +females have been found mostly from June to August throughout its +range. This species was observed calling at Finca Taboga, Guanacaste +Province, Costa Rica, in mid-July. The males were calling from +temporary grassy and weedy ponds in which _Hyla microcephala_ also was +calling, but the two species had different calling sites. _Hyla +staufferi_ called at stations at heights of five to 80 cm. near the +edge of the pond, whereas _Hyla microcephala_ called from emergent +vegetation in the middle of the pond. Charles W. Myers informed me that +at Penonomé, Coclé, Panamá, he found _staufferi_ calling from grass in +puddles where _microcephala_ was absent, and at El Caño, Coclé, Panamá, +_staufferi_ was calling from higher sites ("several inches to a few +feet above water") than _microcephala_. + +Stuart (1948:34) reported breeding individuals from La Libertad, +Guatemala, after rainfall in late May, and Schmidt and Stuart +(1941:239) reported _staufferi_ breeding in July in the Salamá basin, +Alta Verapaz, Guatemala. Stuart (1935:38) and Duellman (1960:63 and +1963:226) agreed that this species breeds early in the rainy season. +However, Rand (1957:519) stated that in El Salvador "these frogs did +not begin to call until almost a month and a half after the beginning +of the rains." Blair (1960:133) reported that males call in June and +July in Chiapas, Oaxaca, Veracruz, and Tamaulipas, México. + +The mating call of this species is a series of closely spaced notes +having a fundamental frequency of about 100 cycles per second. Each +note has a duration of 0.13 to 0.23 second, repeated at intervals that +are longer than the duration of the call. The notes are moderately +low-pitched and have a dominant frequency of more than 3,000 cycles per +second and about 120 pulses per second (Table 2). + +_Tadpoles._--Measurements of the 33 tadpoles that are available are +given in Table 8. The largest tadpole examined is in stage 38 and has a +total length of 29.5 mm. + +A typical tadpole in stage 38 of development (KU 104162, 5 km ESE +Córdoba, Veracruz, México) has a body length of 10 mm., tail length of +19.5 mm., and a total length of 29.5 mm. Other characters are as +follows: body as deep as wide, depressed anteriorly; body as long as +depth of tail; interorbital space greater than distance between eye and +snout but equal to internarial space; nostril equidistant between eye +and tip of snout; distance between spiracle and eye less than distance +between eye and snout; eyes large, situated dorsolaterally; mouth +anteroventral, approximately triangular in outline; one row of papillae +covering lower lip and all except median fourth of upper lip; scattered +papillae at corners of mouth; tooth rows 2/3; first upper row entire, +second row interrupted medially, shorter than first; lower rows shorter +than upper rows; beak weak; spiracle short and nearer eyes than anus; +anal opening not reaching edge of ventral fin; dorsal fin barely +extending onto body; caudal musculature pointed distally. + +TABLE 8.--Sizes of Tadpoles of _Hyla s. staufferi_ in Relation to +Developmental Stages. (Means in parentheses below observed ranges; +measurements in mm.) + + ====================================================== + Stage | N | Body length| Tail length | Total length + --------+---+------------+-------------+-------------- + 25 | 3 | 6.0-7.0 | 12.0-13.0 | 18.0-20.0 + | | (6.7) | (12.5) | (19.2) + | | | | + 26 | 2 | 7.0-7.5 | 14.0-15.0 | 21.5-22.0 + | | (7.3) | (14.5) | (21.8) + | | | | + 27 | 9 | 7.0-8.0 | 13.0-17.0 | 21.0-25.0 + | | (7.6) | (14.5) | (22.0) + | | | | + 32 | 1 | 8.5 | 15.5 | 24.0 + | | | | + 36 | 2 | 8.0-10.0 | 16.5-17.0 | 25.0-26.5 + | | (9.0) | (16.8) | (25.8) + | | | | + 38 | 6 | 9.0-10.0 | 19.0-20.5 | 28.0-29.5 + | | (9.6) | (19.5) | (29.1) + | | | | + 41 | 1 | 10.0 | 14.0 | 24.0 + | | | | + 42 | 6 | 11.0-14.0 | 10.0-13.0 | 20.0-29.0 + | | (11.8) | (11.9) | (24.8) + | | | | + 45 | 1 | 12.5 | 0.5 | 13.0 + | | | | + 46 | 1 | 13.0 | -- | -- + +In life, body pale olive-tan, belly silvery white with pinkish-orange +reticulations in some specimens; tail creamy white with silvery flecks +and black or brown reticulations. In preservative, tan and +pinkish-orange coloration lost; body transparent, reticulations on tail +present. + +_Remarks._--_Hyla staufferi_ was described by Cope (1865:195) on the +basis of specimens from Orizaba, Veracruz, México. He described the +color pattern as "color above dark olive, with a short black bar over +each scapula, and one from eye to eye, with a trace along the coccyx." +Cope (1887:14) placed _staufferi_ as a subspecies of _Hyla eximia_, but +he did not justify his action. Günther (1901:262) also considered +_staufferi_ to be conspecific with _eximia_ without making any +qualifying statement. Dunn and Emlen (1932:24) named _Hyla culex_ from +Tela, Honduras, on the basis of a male (MCZ 16098) having a snout-vent +length of 25.1 mm., and a female (USNM 20267) from Patuca, Honduras. +They diagnosed the species as having "discs larger than tympanum ... +black interorbital triangle, traces of black dorsal marking; three +black bars on anterior and posterior face of thighs, two black bars on +tibia, on tarsus and on forearm." The holotype now is faded but has +some of the pattern described. Dunn and Emlen did not compare _culex_ +with _staufferi_ but did compare it with _boulengeri_ and _rubra_. + +Dunn (1933:61) named _Hyla altae_ from Summit, Canal Zone. His +description was based on a male (MCZ 17972) having a snout-vent length +of 25.1 mm., the color pattern was described as "gray with four darker +dorsal stripes ... a faint trace of mid-dorsal striping...." Dunn +defined the _Hyla rubra_ group and recognized _boulengeri_, _altae_, +_culex_, and _rubra_ as members. _Hyla elaeochroa_ and _staufferi_ were +omitted from his key to the group in Central America. + +Kellogg (1932:174) compared _staufferi_ with _eximia_ and concluded +that the two were probably distinct species. Stuart (1935:38) +considered _altae_ to be a synonym of _culex_. Gaige (1936:293) +considered _altae_ and _culex_ to be conspecific but regarded +_staufferi_ as a different species. She also suggested that _staufferi_ +was not related to _eximia_ but belonged to the _rubra_ group. Taylor +(1952:865) and Duellman (1966a:274) considered _altae_ and _culex_ to +be synonyms of _staufferi_. + +The only other worker besides Cope and Günther to consider _Hyla +staufferi_ as a member of the _eximia_ group was Blair (1960:129), who +suggested the relationship on the basis of similarities in the +structure of the calls of _eximia_ and _staufferi_. Taylor (1938:421) +and Smith and Taylor (1948:78) excluded _staufferi_ from the _eximia_ +group on the basis of morphological characteristics. I consider _culex_ +to be inseparable from _staufferi_, whereas _altae_ is recognizable as +a Panamanian subspecies of _staufferi_. + + +_Hyla staufferi staufferi_ Cope, New Combination + + _Hyla staufferi_ Cope, Proc. Acad. Nat. Sci. Philadelphia, 17:195, + October 1865 [Holotype.--USNM 15317, Orizaba, Veracruz, México; + Francis Sumichrast collector], Brocchi, Mission Scientifique au + Mexique et dans L'Amerique Centrale, 1881, p. 36. Boulenger, + Catalogue, of the Bratrachia Salientia s. Ecaudata, p. 400, + February 1, 1882. Kellogg, Bull. U.S. Natl. Mus., 160:173, March + 31, 1932. Smith and Taylor, Bull. U.S. Natl. Mus., 194:88, 1948. + Taylor, Univ. Kansas Sci. Bull., 35:862, July 1, 1952. Rand, + Fieldiana Zool. Chicago Nat. Hist. Mus., 34:518, April 18, 1957. + Duellman, Univ. Kansas Publ, Mus. Nat. Hist., 17:274, June 17, + 1966. + + _Hyla eximia staufferi_ Cope, Bull. U.S. Natl. Mus., 32:14, January + 16, 1887. + + _Hyla eximia_ (part): Günther, Biologia Centrali-Americana, + Reptilia and Batrachia, p. 261, June 1901. Nieden, Das Tierreich, + Anura I, p. 245, June 1923. + + _Hyla culex_ Dunn and Emlen, Proc. Acad. Nat. Sci. Philadelphia, + 84:24, March 22, 1932 [Holotype.--MCZ 16098, Tela Honduras; Raymond + A. Stadelman collector]. Stuart, Misc. Publ., Univ. Michigan Mus. + Zool., 29:38, October 1935. Gaige, Carnegie Inst. Washington Publ., + 457:293, 1936. + +_Diagnosis._--Small frogs (Male to 29 mm., Female to 31.6 mm.); +dorsolateral stripes irregular; paravertebral stripes usually broken; +two or three transverse bars on shanks; thighs spotted or not; arms +usually barred; interorbital bar usually present; toes about three +fourths webbed; color brown, tan, or olive-green. + +_Variation._--Three hundred and sixty males chosen at random from +throughout the range have snout-vent lengths of 20.7 to 29 mm. (25.9 +mm.). The smallest individuals are from Costa Rica and Nicaragua (means +24.2 and 24.4 mm., respectively). The largest individuals are from +Guatemala and El Salvador (mean of each 27.0 mm.). The ratio of the +diameter of the tympanum to that of the eye is more than 60 per cent in +most samples, but in those from Costa Rica and British Honduras it is +smaller. The color pattern is highly variable. Some specimens are dark +brown or pale brown in color. Incomplete dorsal stripes are present in +94.6 per cent of the specimens, and transverse bars are present on the +shanks in 98.3 per cent of the specimens. The interorbital spot varies +from transverse to longitudinal in position, and an irregular white +line extends from the upper jaw to the arm in some specimens (Table 7). + +_Distribution._--_Hyla staufferi staufferi_ inhabits savanna and +subhumid and xeric forests in the lowlands and moderate elevations from +southern Tamaulipas southward to Nicaragua on the Caribbean versant and +from Guerrero, México to northwestern Costa Rica on the Pacific +lowlands (Fig. 7). Duellman (1963:226) commented that a specimen from +Chinajá, Guatemala, possibly was transported there in the cargo from +Toocog, because with this one exception the species is unknown in +tropical rainforest in Guatemala. + + [Illustration: Fig. 7. Map showing locality records for _Hyla + staufferi staufferi_ (circles) and _H. staufferi altae_ (dots).] + +_Specimens Examined._--México: _Campeche_: 5 km S Champotón KU 71296-7; +7 km W Escárcega, KU 71298-308; 13 km W, 1 km N Escárcega, KU 71309-10, +75090-4. _Chiapas_: 32 km S Arriaga, KU 57789-92; 4 km N Ixtapa, KU +5776-81; 3.6 km SW Las Cruces, KU 37740; 17 km S Las Cruces, KU +57793-4; 24 km S Las Cruces, KU 104160 (tadpoles); 11 km S Tapachula, +KU 57782-8, 60000 (young). _Guerrero_: El Limoncito, near La Venta, KU +31392-401; Mexcala, near Balsa River, KU 31391; Organos, S El Trienta, +KU 31390. _Oaxaca_: 26 km N Matías Romero, KU 33878-82; 2.5 km S +Pochutla, KU 59924-7 (skeletons); 5 km S Pochutla, KU 57795-801; 3.2 km +E Tapanatepec, KU 37877-902; 17.6 km WNW Tapanatepec, KU 65033-4; +Temascal, USC 8243 (8); 3.2 km S Tolocita, KU 39657-8; 0.5 km Tuxtepec, +KU 87073-81, 87610 (tadpoles); 17 km S Tuxtepec, KU 65035-7; 1 km W +Zanatepec, KU 104161 (tadpoles). _Quintana Roo_: Isla Cozumel, 3.5 km N +San Miguel, KU 71710-11 (young). _San Luis Potosí_: Valles, KU 31490. +_Tabasco_: Teapa, UMMZ 118887 (3), 119203 (13); 9.6 km N Teapa, UMMZ +119202; 24 km N Teapa, UMMZ 119961 (5); 29 km N Teapa, UMMZ 119960; 3.5 +km S Villahermosa, UMMZ 119201 (2); 17.6 km S Villahermosa, UMMZ 119200 +(8). _Tamaulipas_: 1 km E Chamal, UMMZ 110706; Gómez Farías, UMMZ +110701 (3); 5 km SE Gómez Farías, UMMZ 110705; 8 km NE Gómez Farías, +UMMZ 11282 (2), 11283 (3); Kilometer 615 between Río Limón and Llera, +UMMZ 80455 (2); 5 km W San Geraldo, UMMZ 110702 (4), 110703 (3); 8 km W +San Geraldo, near Río Frío, UMMZ 110704 (5). _Veracruz_: 3 km SW Boca +del Río, KU 10494-8; 5 km SW Boca del Río, KU 23701; 5 km ESE Córdoba, +KU 104162 (tadpoles); Cuautlapán, KU 57098-102, 26787; Hacienda +Tamiahua, Cabo Rojo, KU 62871; 2 km ENE Mata Oscura, KU 105627; 5 km SE +Paso del Toro, KU 40144; Portrero Viejo, KU 23911-2, 26786, 27413, +57094-7. + +Guatemala: _Alta Verapaz_: Chinajá, KU 57769; Finca La Cubilquitz, UMMZ +90871, 90872 (5), 91379 (2). _Baja Verapaz_: 1 km S San Jerónimo, UMMZ +84077 (7), 84078 (14). _Chiquimula_: 1.6 km SE Chiquimula, UMMZ 98114 +(2); Esquipulas, UMMZ 106784 (4), 106785 (14). _El Petén_: No specific +locality, USNM 25143, 24825-6; La Libertad, FMNH 27096-7, KU 57770, +UMMZ 75339 (15), 75340 (15), UMMZ 94341-2. _Esquintla_: 20 km N San +José, AMNH 74369-76. _Guatamala_: 16 km NE Guatamala, KU 43539. Izábal: +Puerto Barrios, TCWC 16671-73, 16646-56; 2.5 km NE Río Blanco, KU +57774-5. _Jalapa_: Jalapa, UMMZ 106788 (44). _Jutiapa_: Finca La +Trinidad, UMMZ 107730 (12), 107731 (16); Jutiapa, UMMZ 106786 (2). +_Zacapa_: 14 km ENE Mayuelas, KU 57773; 7 km ENE Río Hondo, KU 57771-2, +59999 (young). + +British Honduras: BELIZE: Belize, FMNH 4406. _El Cayo_: San Agustín, +UMMZ 80741 (8). _Stann Creek_: 10 km S Stann Creek on Hummingbird +Highway, UMMZ 125720-1. + +El Salvador: _Cuscatlán_: 7 km WNW Cojutepeque, TNHC 32004-10. _La +Libertad_: 16 km NW Santa Tecla, KU 43540-1. _La Union_: 2.5 km Santa +Rosa, TCWC 16669-70. _Morazán_: Dividendero, USNM 73288-92. _San +Salvador_: San Salvador, FMNH 65101-06, KU 61932-44, 61989-92, 62152 +(eggs), USNM 117588, 118391 (3), 118394; 1.6 km NW San Salvador, KU +43162-3. + +Honduras: _Atlantidad_: Ceiba, USNM 117592. _Choluteca_: Choluteca, KU +85361-6; 2 km E Choluteca, UMMZ 118395 (7); 3.2 km NE Choluteca, KU +100500-01; 6.2 km E Choluteca, KU 65046-56; 10 km E Choluteca, KU +65045: 5 km S Choluteca, USC 2700 (4). _Colón_: Isla Guanaja (Islas de +la Bahía), TCWC 21551, TNHC 32011. _Cortés_: Agua Azul, TCWC 19178-9; +East side Lago Yojoa, KU 65038-44. _El Paraiso_: Valle de Jamastrán, +AMNH 54800-04. _Francisco Morazán_: Escuela Agrícola Panamericana, AMNH +54963-73; 14.5 km NW Comayaguela, KU 100499; El Zamorano, KU 103224; 29 +km N Tegucigalpa, TNHC 32003, 32012. + +Nicaragua: _Chinandega_: Finca San Isidro, 10 km S Chinandega, KU +85311-33. _Managua_: 13 km E Managua, KU 85339; 2 km S Tipitapa, KU +85334-8. _Rivas_: 9.5 km SE Rivas, KU 85355-6; 18 km SE Rivas, KU +85354; 7.7 km NE San Juan del Sur, KU 85346-53; 16.5 km NE San Juan del +Sur, KU 85340-5; 5 km SE San Pablo, KU 43151-61. _Zelaya_: Isla Grande +del Maiz, KU 85357-60. + +Costa Rica: _Alajuela_: _Los Chiles_, USC 7215 (2), 7217. _Guanacaste_: +4 km W Bagaces, USC 7019 (5); Finca Taboga, KU 102265-5; 12 km S La +Cruz, USC 8091; Las Cañas, KU 41113 (skeleton); 27 km N Las Cañas, USC +8171 (5); Guardia, Río Tempisque, USC 8214; 10 km N Guardia, KU +102266-7; 1.6 km N Guayabo de Bagaces, USC 7023 (3); Liberia, KU +36510-22; 4 km W Liberia, KU 36449-64, USC 102 (10), 103 (9), 104 (7), +105; 6 km N Liberia. USC 8096; 8 km NNW Liberia, KU 65032; 14.5 km N +Liberia, USC 8079, 8138 (2); 14.5 km S Liberia, USC 8238 (5); 6 km N +Nicoya, USC 8229 (11); 4 km S Nicoya, USC 8230, 8231; Peñas Blancas, KU +102263; 8.6 km ESE Playa del Coco, USC 8137 (14); 21 km E Playa del +Coco, USC 8138 (2); Santa Cruz, USC 8232 (2); 3 km E Santa Rosa, TCWC +16663-68; Tenorio, KU 32159; Tilarán, KU 36509. _Puntarenas_: 10 km WNW +Esparta, KU 65022-9, 68614 (skeleton); 4.5 km WNW Esparta, KU 65030; 12 +km WNW Esparta, KU 65031; 6 km E Esparta, KU 86477; Hotel Maribella, KU +32157-8; 3 km W Puntarenas, TCWC 16657-62. + + +_Hyla staufferi altae_ Dunn, New Combination + + _Hyla altae_ Dunn, Occas. Papers Boston Soc. Nat. Hist., 8:61, June + 7, 1933 [Holotype.--MCZ 17972, Summit, Canal Zone, Panamá; Emmett + R. Dunn collector]. + + _Hyla culex_: Stuart, Misc. Publ. Univ. Michigan Mus. Zool., 29:38, + October 1, 1935. Gaige, Carnegie Inst. Washington Publ., 457:293, + 1936. + + _Hyla staufferi_: Taylor, Univ. Kansas Sci. Bull., 35:862, July 1, + 1952. Duellman, Univ. Kansas Publ., Mus. Nat. Hist., 17:274, June + 17, 1966. + +_Diagnosis._--Small frogs (Male to 26 mm., Female to 27 mm.); +dorsolateral and paravertebral stripes complete; longitudinal dark gray +stripe on shank; thighs unmarked; interorbital bar usually absent; toes +about three fifths webbed; gray to brownish gray above. + +_Variation._--_Hyla staufferi altae_ is less variable in size, +proportions, and color pattern than is _H. s. staufferi_. The size +varies from 21.7 to 26 mm. (23.6) in 72 males. The ratio of tibia to +snout-vent length is 0.42 to 0.50 (0.45), slightly less than in the +northern subspecies. In color pattern 94.5 per cent of the individuals +have complete dorsal stripes, and all have a longitudinal stripe on the +shank (Table 7). + +_Distribution._--This subspecies is restricted to subhumid forests and +savannas on the Pacific lowlands of Panamá. _Hyla s. altae_ is +presently known to occur from Chepo in east-central Panamá through the +Azuero Peninsula to Concepción, Chiriquí, in western Panamá (Fig. 7). + +_Specimens Examined._--Panamá: _Canal Zone_: No specific locality, TNHC +24406; 2.8 km SW Fort Kobbe, KU 101679. _Chiriquí_: 14.4 km E +Concepción, AMNH 69799-801; 6.6 km N David, TNHC 32013-4; 2 km S David, +AMNH 68802. _Coclé_: 1 km NE El Caño, KU 101662-75; El Valle de Antón, +AMNH 59601-5, KU 77333-47; 7 km SSW Penonomé, KU 101654-61. _Los +Santos_: Tonosí, KU 101246 (tadpoles), 101697-701. _Panamá_: 2 km WSW +Chepo, KU 101680-8; 6 km WSW Chepo, KU 77324-27; El Cangrejo (Panamá), +KU 101676-8; Nueva Gorgona, AMNH 69991, 69798; 1.5 km W Pacora, KU +77328-32; 2 km N Tocumen, KU 101689-95; 8 km NE Tocumen, KU 101696. + + + + +EVOLUTIONARY HISTORY + + +My assumptions regarding the evolutionary history of the _Hyla rubra_ +group in Central America were derived partly from interpretations of the +evolutionary history of other animal groups (Simpson, 1943, 1965; Dunn, +1931b; Stuart, 1950; Duellman, 1958, 1960, 1963, 1965; and Duellman and +Trueb, 1966). The origin and early evolution of the group probably +occurred prior to the Mid-Pliocene in the lowlands of South America, +because the greatest diversity of the group is in Brazil. Differentiation +into two or more subgroups took place in South America prior to the late +Pliocene. At the end of the Pliocene, shortly after the closure of the +Colombian Portal, many South American animals migrated into Central +America (Simpson, 1943, Maldonado-Koerdell, 1964, and Savage, 1966). It +is likely that the _Hyla rubra_ group entered Central America at that +time; apparently two stocks (_rubra-elaeochroa-staufferi_ stock and +_boulengeri-foliamorta_ stock) migrated into Central America. + +_Hyla elaeochroa_ is closely related to _rubra_ and probably +differentiated from _rubra_ through spatial isolation. Thus, we have +_elaeochroa_ in Central America and _rubra_ in South America; most +likely only in relatively recent times has _rubra_ migrated into +eastern Panamá from northern South America. The differentiation and +dispersal of _elaeochroa_ and _staufferi_ took place in Central America +after the Pliocene. Probably the events of the Pleistocene resulted in +the isolation of populations. One of these (_Hyla staufferi_ stock) was +restricted in the subhumid Pacific lowlands, whereas the _Hyla +elaeochroa_ stock occupied the tropical wet forests of the Caribbean +lowlands. _Hyla elaeochroa_ apparently more closely resembled the +parental stock by being restricted to the tropical rain forests, +whereas _staufferi_ adapted to subhumid environments and thereby was +able to disperse throughout most of the subhumid regions of Central +America. + +After geographical separation took place the initial genetic divergence +between the two populations was maintained by means of ecological and +ethological isolating mechanisms. Under these circumstances it can be +supposed that the different ecological preferences of _elaeochroa_ and +_staufferi_ depend on the climatic changes that took place during the +Pleistocene. On this basis it may be proposed that when the original +prototype broke up into the two incipient species, the _staufferi_ +stock became physiologically and behaviorally adapted to subhumid +conditions and dispersed into dry areas of the lowlands of Middle +America. The tropical evergreen forests on the Caribbean side of lower +Central America and the uplift of the Talamanca range in the Pliocene +were barriers to the dispersal of _staufferi_. Consequently, this frog +dispersed along the Pacific lowlands. + +At the present time _staufferi_ occupies the length of the Pacific +lowlands in Central America, except in the rainforest of the Golfo Duce +region, which apparently is a relict stand and now separates the ranges +of two subspecies of _Hyla staufferi_. This species crossed the central +Nicaraguan lowlands and reached the Caribbean lowlands of Nicaragua and +nuclear Central America. The species migrated through the subhumid +corridor in northern Honduras and eastern Guatemala (Comayagua Valley +in Honduras and the Motagua Valley of Guatemala) to the Isthmus of +Tehuantepec. Duellman (1960) hypothesized "that during the times of +glacial advances (Pleistocene) the lowlands of the Isthmus probably +were more extensive and had more semiarid tropical environments than at +the present" and that when semiarid environments were continuous from +the Pacific slope across the isthmus to the Gulf lowlands _staufferi_ +and other amphibians migrated northward to southeastern Tamaulipas, +México. + +_Hyla elaeochroa_ dispersed along Caribbean lowland routes. This +species not only occurs in the wet forests of the Golfo Dulce region +but also in Guanacaste. It is possible that _elaeochroa_ entered +Guanacaste and moved to the Golfo Dulce region when the intervening +area was less xeric than now (Duellman, 1966b). _Hyla elaeochroa_ +extended its range to eastern Nicaragua, but even though northeastern +Nicaragua has over 2,000 mm. of precipitation annually (Vivo Escoto, +1964), this species has not spread into Honduras and Guatemala. + +_Hyla boulengeri_ is widespread in Amazonian and northern South +America, whereas _foliamorta_ occurs only in eastern Panamá and in +north-central Colombia. The ancestral _boulengeri-foliamorta_ stock +probably invaded Central America in the late Pliocene and dispersed +through humid forested environments to Nicaragua. Apparently a +peripheral population established itself in the dry Pacific lowlands of +Panamá. This population differentiated from _boulengeri_ of the humid +Caribbean lowlands and evolved into _foliamorta_, which subsequently +expanded its range into Colombia. + + + + +LITERATURE CITED + + +BLAIR, W. F. + +1960. Mating call as evidence of relations in the _Hyla eximia_ group. +Southwestern Nat., 5(3):129-135. November 1. + +BOULENGER, G. A. + +1882. Catalogue of the Batrachia Salientia s. Ecaudata, 2nd. ed., pp. +1-503, pls. 1-30. February. + +BROCCHI, P. + +1881-1883. Etude des batraciens de l'Amerique Centrale. Mission +Scientifique au Mexique et dans l'Amerique Centrale, Liv. 1, pp. 1-122, +pls. 1-21. + +COCHRAN, D. M. + +1955. Frogs of southeastern Brazil. Bull. U.S. Natl. Mus., 206:1-423. + +COPE, E. D. + +1865. Third contribution to the Herpetology of Tropical America. Proc. +Acad. Nat. Sci. Philadelphia, 17:195. + +1876. On the batrachia and reptilia of Costa Rica. Jour. Acad. Nat. +Sci. Philadelphia, new series, 8:93-154, pls. 23-28. 1887. Thirteenth +contribution to the Herpetology of Tropical America. Proc. Amer. +Philos. Society, 23:273. April. + +DAUDIN, F. M. + +1802. Histoire naturelle des rainettes, des grenouilles et des +crapauds. Paris, pp. 1-71, pls. 1-33. + +1803. Histoire naturelle générale et particulière des reptiles. Paris, +8:1-439. + +DUELLMAN, W. E. + +1956. The frogs of the hylid Genus _Phrynohyas Fitzinger_, 1843. Misc. +Publ. Mus. Zool., Univ. Mich., 96:1-47, pls. 1-6. February 21. + +1958. A monographic study of the colubrid snakes Genus _Leptodeira_. +Bull. Amer. Mus. Nat. Hist., 114:1-152, pls. 1-31. February 24. + +1960. A distributional study of the amphibians of the Isthmus of +Tehuantepec, México. Univ. Kansas Publ., Mus. Nat. Hist, 13:19-72, pls. +1-8. August 16. + +1963. Amphibians and reptiles of the rainforests of southern El Petén, +Guatemala. Univ. Kansas Publ., Mus. Nat. Hist., 15:205-249, pls. 7-10. +October 4. + +1965. 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Field Mus. Nat. Hist., Zool. Ser., 24:233-247. + +SEBA, A. + +1734. Locupletissimi rerum naturalium thesauri accurata descriptio, et +iconibus artificissimis expressio, per universam physis historiam. +Amsterdam, I xxxiv + 178 pp., pls. 1-111. + +SIMPSON, G. G. + +1943. Turtles and the origin of the fauna of Latin America. Amer. Jour. +Sci., 24:413-429. + +1965. The geography of evolution. Chilton Books Publishers. +Philadelphia, pp. 1-349. July. + +SMITH, H. M. and E. H. TAYLOR + +1948. An annotated checklist and key to the amphibians of México. Bull. +U.S. Natl. Mus., 194:1-118. + +STARRETT, P. + +1960. Description of tadpoles of Middle American frogs. Misc. Publ. +Mus. Zool., Univ. Michigan, 110:1-37, pl. 1. + +STUART, L. C. + +1935. A contribution to a knowledge of the herpetology of a portion of +the savanna region of Central Petén, Guatemala. Misc. Publ. Mus. Zool., +Univ. Michigan, 29:1-56, pls. 1-4. October 1. + +1948. The amphibians and reptiles of Alta Verapaz, Guatemala. Misc. +Publ. Mus. Zool., Univ. Michigan, 69:1-109. June 12. + +1950. A geographic study of the herpetofauna of Alta Verapaz, +Guatemala. Contr. Lab. Vert. Biol., Univ. Michigan, 45:1-77, pls. 1-9. +May. + +TAYLOR, E. H. + +1938. Frogs of the _Hyla eximia_ group, with description of two new +species. Univ. Kansas Sci. Bull., 25:421-445. June 1. + +1952. The frogs and toads of Costa Rica. Univ. Kansas Sci. Bull., +35:577-942. July 1. + +1958. Additions to the known herpetological fauna of Costa Rica, with +comments on other species. No. III. Univ. Kansas Sci. Bull., 34:3-40. +November 18. + +VIVO ESCOTO, J. A. + +1964. Weather and climate of México and Central America. _In_ R. +Wauchope and R. C. West (Eds.), Handbook of Middle American Indians. +Vol. 1, Univ. Texas Press, Austin, 570 pp. + + +_Transmitted February 7, 1969._ + + + + + + +End of the Project Gutenberg EBook of The Systematics of the Frogs of the +Hyla Rubra Group in Middle America, by Juan R. 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