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diff --git a/.gitattributes b/.gitattributes new file mode 100644 index 0000000..6833f05 --- /dev/null +++ b/.gitattributes @@ -0,0 +1,3 @@ +* text=auto +*.txt text +*.md text diff --git a/34604-8.txt b/34604-8.txt new file mode 100644 index 0000000..b1cf64c --- /dev/null +++ b/34604-8.txt @@ -0,0 +1,2482 @@ +The Project Gutenberg EBook of Middle American Frogs of the Hyla +microcephala Group, by William E. Duellman and M. J. Fouquette + +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: Middle American Frogs of the Hyla microcephala Group + +Author: William E. Duellman + M. J. Fouquette + +Release Date: December 9, 2010 [EBook #34604] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK MIDDLE AMERICAN FROGS *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +https://www.pgdp.net + + + + + + + + + +University of Kansas Publications + +Museum of Natural History + +Volume 17, No. 12, pp. 517-557, pls. 13-16, 9 figs. +March 20, 1968 + +Middle American Frogs +of the Hyla microcephala Group + +BY + +WILLIAM E. DUELLMAN AND M. J. FOUQUETTE, JR. + +University of Kansas +Lawrence +1968 + + + + +University of Kansas Publications, Museum of Natural History + +Editors: E. Raymond Hall, Chairman, Henry S. Fitch, +Frank B. Cross + +Volume 17, No. 12, pp. 517-557, 4 pls. 9 figs. +Published March 20, 1968 + +University of Kansas +Lawrence, Kansas + +PRINTED BY +ROBERT R. (BOB) SANDERS, STATE PRINTER +TOPEKA, KANSAS +1968 + +31-9419 + + + + +Middle American Frogs +of the Hyla microcephala Group + +BY +WILLIAM E. DUELLMAN AND M. J. FOUQUETTE, JR. + + +CONTENTS + + + PAGE + +Introduction 519 + Acknowledgments 520 + Materials and Methods 520 + +Hyla microcephala Group 521 + Key to Species and Subspecies 522 + +Accounts of Species and Subspecies 523 + +Cranial Osteology 540 + +Analysis of Mating Calls 544 + +Life History 550 + +Phylogenetic Relationships 552 + +Literature Cited 556 + + + + + +INTRODUCTION + + +The small yellow tree frogs, _Hyla microcephala_ and its relatives, +are among the most frequently heard and commonly collected frogs in +the lowlands of southern México and Central America. The similarities +in size, proportions, and coloration of the different species have +resulted not so much in a multiplicity of specific names, but in +differences of opinion on the application of existing names to the +various taxa. For example, the populations on the Atlantic lowlands +have been known by three names, two of which have been applied to +other taxa. Much of the confusion has been the result of previous +workers' unfamiliarity with the animals in life and unawareness of the +intraspecific geographic variation in the most widespread species. + +Independently we undertook studies of these frogs in the field. The +second author worked on the interspecific relationships and isolating +mechanisms in Panamá (Fouquette, 1960b) and later studied the species +in southern México. As part of his survey of the hylids of Middle +America, the first author accumulated field and laboratory data on the +frogs throughout their ranges in México and Central America. The +purpose of this report is to present our findings on the four species +of Middle American frogs that we place in the _Hyla microcephala_ +group. In addition to conventional taxonomic characters, we have +utilized the features of the cranial osteology and have relied heavily +on the data obtained from an analysis of the mating calls. +Furthermore, we have included ecological and distributional data in +our synthesis of interspecific relationships. + + +ACKNOWLEDGMENTS + +Examination of specimens was made possible by the provision of working +space at various institutions or through the loan of specimens. For +their generosity in this manner we are grateful to Richard J. Baldauf, +Charles M. Bogert, James E. Böhlke, Doris M. Cochran, Robert F. Inger, +John M. Legler, Alan E. Leviton, Gerald Raun, Jay M. Savage, Hobart M. +Smith, Robert C. Stebbins, Wilmer W. Tanner, Charles F. Walker, Ernest +E. Williams, and Richard G. Zweifel. + +Duellman is especially grateful to Charles W. Myers, Linda Trueb, +Jerome B. Tulecke, and John Wellman for their assistance in the field +and to Linda Trueb for her work on the cranial osteology that is +incorporated in this report. Fouquette is indebted to H. Morgan Smith +and A. C. Collins for assistance in the field, to A. J. Delahoussaye +for assistance in the laboratory, and to W. Frank Blair for use of the +facilities of the sound laboratory at the University of Texas and for +much help in the early stages of this study. + +The research reported herein was accomplished mainly through support +by the National Science Foundation (grants NSF G-9827 and GB-1441 to +Duellman and GB-599 to Fouquette). The latter's field work in México +was assisted in part by NSF Grant G-4956 to W. Frank Blair. Some of +the field studies carried out in Panamá by Duellman were supported by +a grant from the National Institutes of Health (NIH GM-12020). + +We are grateful to many persons, too numerous to mention, who in +various ways aided our field work in Middle America. We are especially +indebted to Dr. Rodolfo Hernandez Corzo and the late Ing. Luis Macías +Arellano of the Dirección General de la Fauna Silvestre of the Mexican +government for providing permits to collect in México. + + +Materials and Methods + +For this report, data has been obtained from 2829 preserved frogs, 42 +skeletal preparations, 8 lots of tadpoles and young, and 4 lots of +eggs. Much of the material was collected in our independent field +work, which has extended over a period of 11 years. + +Measurements were taken in the manner described by Duellman (1956). +Osteological data were obtained from specimens that were cleared in +potassium hydroxide, stained with alizarin red, and stored in +glycerine. Recordings were made by means of Magnemite portable tape +recorders (Amplifier Corp. America). The calls recorded by Fouquette +were analyzed on a Sonagraph (Kay Electric Co.) at the University of +Texas; those recorded by Duellman were analyzed mainly on a Vibralyzer +(Kay Electric Co.) at the University of Kansas and in part on a +Sonagraph at the University of Southwestern Louisiana. Sample calls +were analyzed on all three instruments; the slight differences in +results were found to be less than the error in measurement, so the +data from all sources were combined without correction. The techniques +and terminology of the calls are those defined by Fouquette (1960a, +1960b). + +In the accounts of the species we have attempted to give a complete +synonymy. At the end of each species account the localities from which +specimens were examined are listed alphabetically within each state, +province, or department, which in turn are listed alphabetically +within each country. The countries are arranged from north to south. +Localities preceded by an asterisk (*) are not plotted on the +accompanying maps due to the crowding of symbols that would have +resulted. Abbreviations for museum specimens are listed below: + + AMNH --American Museum of Natural History + ANSP --Academy of Natural Sciences of Philadelphia + BMNH --British Museum (Natural History) + BYU --Brigham Young University + CAS --California Academy of Sciences + FMNH --Field Museum of Natural History + KU --University of Kansas Museum of Natural History + MCZ --Museum of Comparative Zoology + MVZ --Museum of Vertebrate Zoology + SU --Stanford University + UIMNH--University of Illinois Museum of Natural History + UMMZ --University of Michigan Museum of Zoology + USC --University of Southern California + USNM --United States National Museum + UU --University of Utah + TCWC --Texas Cooperative Wildlife Collection + TNHM --Texas Natural History Museum + + + + + +HYLA MICROCEPHALA GROUP + + +_Definition._--Small hylids attaining a maximum snout-vent length of +27 mm. in males and 32 mm. in females; dorsum yellowish tan with brown +markings; thighs uniformly yellow, vocal sac in breeding males yellow; +snout truncate in lateral profile; tympanum distinct, usually slightly +smaller than one-half diameter of eye; vocal sac single, median, +subgular; fingers about one-third webbed; toes webbed nearly to bases +of discs, except only to middle of antepenultimate or base of +penultimate phalanx of fourth toe; tarsal fold weak; inner metatarsal +tubercle low, flat, elliptical; axillary membrane present; pupil +horizontally elliptical; palpebral membrane unmarked; cranial elements +reduced in ossification; sphenethmoid small, short; frontoparietal +fontanelle large; tegmen tympani not extensive; quadratojugal greatly +reduced; anterior arm of squamosal extending only about one-fourth +distance to maxillary; posterior arm of squamosal not having bony +connection with proötic; nasals lacking maxillary processes; medial +ramus of pterygoid not having bony attachment to proötic; maxillary, +premaxilary, and prevomerine teeth present; palatine and parasphenoid +teeth absent; Mentomeckelians ossified; tadpoles having xiphicercal +tails with deep caudal fins and terminal mouth lacking teeth; mating +call consisting of one primary note followed by a series of shorter +secondary notes; haploid number of chromosomes, 15 (known only in _H. +microcephala_ and _H. phlebodes_.) + +_Content._--As recognized here the _Hyla microcephala_ group contains +four species, one having two subspecies. An alphabetical list of the +specific and subspecific names that we consider to be applicable to +the _Hyla microcephala_ group are listed below. + + + Names Proposed Valid Names + +_Hyla cherrei_ Cope, 1894 ? = _H. m. microcephala_ +_Hyla microcephala_ Cope, 1886 = _H. m. microcephala_ +_Hyla microcephala_ Boulenger, + 1898 (_nec_ Cope, 1886) = _H. microcephala underwoodi_ +_Hyla microcephala martini_ Smith, 1951 = _H. microcephala underwoodi_ +_Hyla microcephala sartori_ Smith, 1951 = _H. sartori_ +_Hyla phlebodes_ Stejneger, 1906 = _H. phlebodes_ +_Hyla robertmertensi_ Taylor, 1937 = _H. robertmertensi_ +_Hyla underwoodi_ Boulenger, 1899 = _H. microcephala underwoodi_ + + +_Discussion._--The color pattern is the most useful character in +distinguishing the species of the _Hyla microcephala_ group from one +another. Except in _Hyla microcephala_, little geographic variation in +color pattern is noticeable. The features of color pattern that are +helpful in identifying the species are: 1) presence or absence of +lateral dark brown stripe; 2) longitudinal extent and width of lateral +stripe, if present; 3) presence or absence of a narrow white line just +dorsal to the lateral dark stripe; 4) presence or absence of an +interorbital dark mark; 5) the arrangement of dark markings on the +back, either as longitudinal lines or series of dashes, or in the form +of various kinds of transverse markings; 6) presence of dark flecks, +longitudinal line, or transverse marks on shanks. + +Few consistent differences in measurements and proportions exist among +the species (Table 1). The most obvious morphological difference is +that the head is noticeably narrower in _H. robertmertensi_ than in +the other species. _Hyla phlebodes_ is the smallest species; adult +males attain snout-vent lengths of only 23.6 mm. The body is slender +in _H. microcephala_ and _robertmertensi_, slightly wider in +_phlebodes_, and noticeably broader in _sartori_. + +_Distribution._--The composite range of the Middle American frogs of +the _Hyla microcephala_ group includes the lowlands of southern México +and Central America, in some places to elevations of 1200 meters, +southeastward from southern Jalisco and southern Veracruz, excluding +arid regions (northern Yucatán Peninsula, Balsas-Tepalcatepec Basin, +Plains of Tehuantepec, Grijalva Valley, Salamá Basin, and upper +Motagua Valley) to the Pacific lowlands and the Cauca and Magdalena +valleys in Colombia. + + +Key to Species and Subspecies + + +1. Lateral dark stripe, bordered above by narrow white line, + extending from snout at least to sacral region 2 + + Lateral dark stripe, if present, not extending posteriorly to + sacral region and not bordered above by narrow white line 4 + +2. Lateral dark stripe continuous to groin; dark flecks or + longitudinal line on shanks; interorbital dark bar absent; + dorsal pattern usually consisting of pair of longitudinal dark + lines or series of dashes 3 + + Lateral dark stripe usually extending only to sacral region; + dark transverse bars on shanks; interorbital bar usually + present; dorsal pattern usually consisting of interconnecting + dark lines, sometimes forming transverse marks + _H. microcephala underwoodi_ + +3. Lateral dark stripe narrow, covering only upper edge of + tympanum; dorsal longitudinal stripes continuous, extending to + vent _H. microcephala microcephala_ + + Lateral dark stripe wide, encompassing entire tympanum; dorsal + markings consisting of longitudinal series of flecks or dashes, + or of two lines, usually not extending to vent _H. robertmertensi_ + +4. Lateral dark stripe indistinct, present only above tympanum and + insertion of arm; dorsal markings consisting of narrow lines + and dashes, sometimes interconnected; transverse bars on shanks + narrow relative to interspaces _H. phlebodes_ + + Lateral dark stripe absent; dorsal markings consisting of two broad + chevron-shaped marks; transverse bars on shanks wide relative to + interspaces _H. sartori_ + + + + +ACCOUNTS OF SPECIES AND SUBSPECIES + + +_Hyla microcephala_ Cope + + +_Diagnosis._--Lateral dark stripe narrow, covering only upper edge of +tympanum, bordered above by narrow white stripe; dorsal pattern +consisting of pair of longitudinal brown lines and no interorbital bar +(eastern populations), or of irregular dark markings forming an X- or +)(-shaped mark in scapular region and an interorbital bar (western +populations). + +_Content._--The populations inhabiting the Pacific lowlands of +southeastern Costa Rica eastward to Colombia are recognized herein as +_Hyla microcephala microcephala_ Cope; the populations in western +Costa Rica northward to México are assigned to _Hyla microcephala +underwoodi_ Boulenger. + +_Distribution._--Southern Veracruz and northern Oaxaca southeastward +through the Atlantic lowlands of Central America to north-central +Nicaragua, thence southeastward on the Pacific lowlands to eastern +Panamá, and thence into the Cauca and Magdalena valleys (Caribbean +drainage) of Colombia (Fig. 1). + + + [Illustration: Fig. 1. Map showing locality records for _Hyla + microcephala_.] + + +Table 1.--Variation in Certain Measurements and Properties in the + Hyla microcephala Group. (All Data Based on Adult Males; + Mean and Standard Error of Mean Below Observed Range.) + +======================================================================== + Locality | N | Snout-vent | Tibia length |Foot length| + | | length | ------------ | --------- | + | | (S-V L) | S-V L | S-V L | +------------------------------------------------------------------------ + | _H. m. microcephala_ + | +Panamá: Canal Zone | 25 | 21.5-24.1 | 50.2-56.0 | 40.9-46.6 | + | | 22.8±0.20 | 52.9±0.37 | 43.5±0.28 | + | | | | | +Costa Rica: Golfito | 25 | 18.5-24.5 | 49.1-54.4 | 41.8-48.0 | + | | 22.4±0.27 | 51.6±0.26 | 45.1±0.32 | + | + | _H. m. underwoodi_ + | | +Nicaragua: La Cumplida | 25 | 23.0-25.6 | 51.0-55.7 | 41.3-46.5 | + | | 24.1±0.19 | 52.9±0.25 | 43.7±0.25 | + | | | | | +Guatemala: Finca Chamá | 25 | 21.8-25.0 | 51.0-57.2 | 41.2-47.8 | + | | 23.5±0.16 | 54.3±0.39 | 44.4±0.30 | + | | | | | +Tabasco: Teapa | 25 | 22.7-25.8 | 48.0-54.5 | 40.7-46.8 | + | | 24.3±0.14 | 51.5±0.29 | 43.3±0.25 | + | | | | | +Oaxaca: Donají-Sarabia | 25 | 22.1-25.9 | 49.8-55.6 | 40.5-46.6 | + | | 23.8±0.19 | 52.8±0.33 | 43.4±0.27 | + | | | | | +Veracruz: Alvarado | 25 | 21.9-25.4 | 49.6-54.4 | 40.7-47.5 | + | | 24.1±0.17 | 51.1±0.28 | 42.6±0.34 | + | + | _H. robertmertensi_ + | +Guatemala: La Trinidad | 21 | 21.8-24.6 | 47.1-52.8 | 40.9-51.3 | + | | 23.4±0.15 | 49.9±0.34 | 43.5±0.17 | + | | | | | +Chiapas: Acacoyagua | 25 | 21.4-25.7 | 47.8-52.4 | 41.7-46.3 | + | | 24.1±0.20 | 50.4±0.45 | 43.9±0.23 | + | | | | | +Oaxaca: Tapanatepec | 25 | 22.4-26.4 | 44.1-48.3 | 39.1-44.5 | + | | 24.7±0.18 | 46.4±0.23 | 41.7±0.23 | + | + | _H. phlebodes_ + | +Panamá: Canal Zone | 25 | 19.6-23.2 | 49.1-56.9 | 41.9-47.1 | + | | 22.2±0.16 | 52.8±0.35 | 45.4±0.26 | + | | | | | +Costa Rica: Turrialba | 25 | 19.7-23.6 | 47.4-55.7 | 38.1-46.4 | + | | 22.0±0.18 | 51.1±0.35 | 42.8±0.38 | + | + | _H. sartori_ + | +Guerrero: Tierra Colorada| 25 | 23.7-26.0 | 47.2-51.4 | 42.4-47.8 | + | | 24.8±0.13 | 49.6±0.23 | 45.2±0.27 | +------------------------------------------------------------------------ +Table 1. (continued) +=============================================================== + Locality | Head length | Head width | Tympanum + | ----------- | ---------- | -------- + | S-V L | S-V L | Eye +--------------------------------------------------------------- + | _H. m. microcephala_ + | +Panamá: Canal Zone | 28.5-32.8 | 28.1-30.9 | 44.0-54.1 + | 31.0±0.22 | 29.4±0.11 | 49.0±0.55 + | +Costa Rica: Golfito | 30.2-35.5 | 29.0-32.7 | 40.0-57.8 + | 33.1±0.25 | 30.8±0.16 | 48.4±1.10 + | + | _H. m. underwoodi_ + | +Nicaragua: La Cumplida | 29.7-33.5 | 28.9-31.8 | 42.3-60.0 + | 31.6±0.19 | 30.4±0.17 | 49.3±0.97 + | +Guatemala: Finca Chamá | 30.8-35.3 | 29.6-33.6 | 37.5-56.4 + | 33.0±0.16 | 31.3±0.36 | 45.2±0.89 + | +Tabasco: Teapa | 29.5-33.0 | 28.7-31.8 | 40.7-53.8 + | 31.7±0.17 | 30.3±0.16 | 45.5±0.38 + | +Oaxaca: Donají-Sarabia | 30.4-34.8 | 28.9-32.6 | 37.0-54.1 + | 32.8±0.19 | 30.8±0.17 | 45.1±0.76 + | +Veracruz: Alvarado | 29.9-33.8 | 29.1-32.9 | 40.7-53.8 + | 31.4±0.18 | 30.5±0.17 | 46.6±0.65 + | + | _H. robertmertensi_ + | +Guatemala: La Trinidad | 30.0-33.3 | 27.3-29.8 | 44.4-50.0 + | 31.3±0.20 | 28.5±0.23 | 47.4±0.46 + | | | +Chiapas: Acacoyagua | 29.1-32.7 | 26.0-30.3 | 42.8-53.8 + | 31.2±0.29 | 28.1±0.20 | 46.5±0.50 + | | | +Oaxaca: Tapanatepec | 26.1-30.4 | 25.4-28.1 | 45.8-58.3 + | 28.4±0.16 | 26.8±0.14 | 52.9±0.77 + | + | _H. phlebodes_ + | +Panamá: Canal Zone | 33.6-37.4 | 32.3-36.0 | 37.9-46.4 + | 34.8±0.18 | 33.8±0.18 | 41.6±0.49 + | | | +Costa Rica: Turrialba | 32.6-35.9 | 30.5-35.0 | 35.7-48.2 + | 34.1±0.16 | 32.9±0.17 | 40.1±0.53 + | + | _H. sartori_ + | +Guerrero: Tierra Colorada| 29.4-31.8 | 28.9-31.0 | 42.3-52.0 + | 30.6±0.13 | 30.0±0.12 | 47.4±0.59 +--------------------------------------------------------------- + + + + +_Hyla microcephala microcephala_ Cope + + + _Hyla microcephala_ Cope, Proc. Amer. Philos. Soc., 23:281, February + 11, 1886 [Syntypes.--USNM 13473 (2 specimens, now lost) from + Chiriquí, Panamá; Mr. MacNeil collector]; Bull. U.S. Natl. Mus., + 32:14, 1887. Günther, Biologia-Centrali Americana, Reptilia and + Batrachia, p. 265, June, 1901. Dunn, Occas. Papers Boston Soc. + Nat. Hist., 5:413, October 10, 1931; Occas. Papers Boston Soc. + Nat. Hist., 8:72, June 7, 1933. Stebbins and Hendrickson, Univ. + California Publ. Zool., 56:524, February 17, 1959. Fouquette, + Evolution, 14:484, December 16, 1960. Busack, Copeia, 2:371, + June 21, 1966. + + ? _Hyla cherrei_ Cope, Proc. Acad. Nat. Sci. Philadelphia, 1894, + p. 195, 1894 [Holotype.--location unknown, apparently lost; + type-locality: "Alajuela, Costa Rica;" R. Alfaro collector]. + Günther, Biologia Centrali-Americana: Reptilia and Batrachia, + p. 264, June, 1901. Taylor, Univ. Kansas Sci. Bull., 35:846, + July 1, 1952. + + _Hyla underwoodi_, Ruthven, Misc. Publ. Mus. Zool., Univ. Michigan, + 8:55, September 15, 1922. Barbour, Proc. New England Zool. Club, + 10:31, March 2, 1928. + + _Hyla microcephala microcephala_, Smith, Herpetologica, 7:185, + December 31, 1951. Taylor, Univ. Kansas Sci. Bull., 39:23, + November 18, 1958. + + +_Diagnosis._--Brown lateral stripe narrow, extending from nostril +along canthus, along upper edge of tympanum to groin, bordered above +by narrow white line; pair of dark brown longitudinal lines on dorsum +extending to vent; shanks having dark longitudinal line or flecks, no +transverse bars; interorbital dark mark lacking. + +_Description and Variation._--The color pattern is nearly constant. Of +103 males from the Canal Zone, all lack an interorbital dark bar, and +all have a dark longitudinal line on the dorsal surface of the shank +and a narrow lateral dark stripe, bordered above by a narrow white +line, extending to the groin. The longitudinal dark lines on the +dorsum are continuous to the groin in 95 specimens and fragmented in +two specimens. In two others the lines converge and fuse in the +scapular region, and in four specimens auxiliary, fragmented lines are +present dorsolaterally. + +In all specimens from southeastern Costa Rica (Golfito, Palmar Sur, +and Villa Neilly) the pattern is constant, except that in about 10 per +cent of the specimens the longitudinal line on the dorsal surface of +the shank is replaced by a row of brown flecks. + +Of the limited number of Colombian specimens examined, all are +patterned normally, except three from Sautata, Chocó, three from +Curumani, and three from Arcataca, Magdalena, which have flecks on the +dorsal surfaces of the shanks, and one from Espinal, Tolima, which has +no markings on the shanks. + +When active at night most individuals are pale yellowish tan dorsally; +the white dorsolateral line is noticeable, but the brown lateral +stripe, dorsal brown lines, and lines on shanks are so pale that often +they are barely discernible. By day the dorsum changes to tan or pale +reddish brown; the stripes are dark brown, and the dorsolateral stripe +that is white at night becomes creamy yellow (Pl. 13). Small brown +flecks are present on the dorsum of most individuals. The venter +always is white, and the iris is pale bronze with a brown tint +immediately anterior and posterior to the pupil. In breeding males the +vocal sac is pale yellow. + +_Tadpoles._--Tadpoles of this species have been found in weed-choked +ponds in eastern Panamá Province. The following description is based +on KU 104097, a specimen in developmental stage 34 (Gosner, 1960). + +Total length, 20.5 mm.; body length, 8.2 mm.; body slightly wider +than deep; snout pointed; nostrils large, situated dorsally, much +closer to snout than eyes, directed anteriorly; eyes moderately +small, situated dorsolaterally and directed laterally; spiracle +sinistral, located just posteroventral to eye; anal tube dextral. +Tail xiphicercal; caudal musculature moderately deep, becoming +slender posteriorly, extending beyond caudal fin; fins deepest at +about one-third distance from body to tip of tail; dorsal fin +extending onto body, deeper than deepest part of caudal +musculature; ventral fin slightly shallower than musculature. +Mouth small, terminal, lacking teeth and fringing papillae, but +having finely serrate beaks. In preservative, top of head pale +brown; dark stripe from tip of snout through eye to posterior edge +of body, narrowing to thin line on proximal one-fourth of tail; +venter white; tail creamy tan with fine black flecks most numerous +posteriorly; posterior two-thirds of fins edged with black. In +life, top of head yellowish tan; lateral stripe brown; belly +white; anterior half of tail lacking pigment; posterior half deep +orange; iris pale bronze (Pl. 15). + +_Remarks._--Evidence for intergradation of _Hyla microcephala_ with +_H. underwoodi_ is provided by four specimens [USC 818 (2), 6081-2] +from 6.1 kilometers northeast of the mouth of the Río Tarcoles, and +nine specimens [USC 8254 (2), 8255, 8256 (4), 8258 (2)] from Parrita, +both in Puntarenas Province, Costa Rica. These localities lie about +two-thirds the distance from the northwesternmost locality for _H. +m. microcephala_ (Palmar Sur) to the southeasternmost locality for +_H. m. underwoodi_ (Barranca). Although in most aspects of coloration +the frogs are more nearly like _H. m. underwoodi_ than _H. m. +microcephala_, some specimens have longitudinal lines on their shanks, +such as are characteristic of _H. m. microcephala_. The dorsal pattern +varies from nearly complete longitudinal lines to broken lines, fused +into an X-shaped scapular mark or not. + +As noted by Rivero (1961:135), _Hyla microcephala_ seems to be closely +related to _Hyla misera_ Werner, a species having a wide distribution +east of the Andes in South America. Despite the similarity in color +pattern, size, and structure, we are reluctant to place the two taxa +in the same species until data on coloration in life, mating calls, +and life history are available for _Hyla misera_ and compared with +those of _Hyla microcephala_. + +The status of Cope's _Hyla cherrei_ is questionable. Since the type, +the only specimen ever referred to the species, apparently is lost, +the only extant information regarding the taxon is contained in the +original description (Cope, 1894). There the species was characterized +as having a narrow dorsolateral white stripe and lacking pigment on +the upper arms and thighs. These characteristics of the color pattern +combined with the statements "vomerine teeth few, opposite the middle +of the very large choanae" and "tympanic drum distinct, one half the +area of eye" serve to distinguish _H. cherrei_ from all other Costa +Rican hylids, except _H. m. microcephala_ and _H. m. underwoodi_. No +statements in the type description will definitely associate _cherrei_ +with one or the other of these subspecies. Since it seems obvious that +_H. cherrei_ can be associated with _H. microcephala_, we prefer to +place the name in the synonymy of the nominate subspecies, thereby +preserving the commonly used name _H. underwoodi_ (Boulenger, 1899) as +a subspecies of _H. microcephala_. + +_Distribution._--_Hyla microcephala microcephala_ inhabits coastal +lowlands from the area of Golfo Dulce (apparently absent from the +Osa Peninsula) in southeastern Costa Rica eastward in Panamá, +including the Azuero Peninsula to northern Colombia and thence +southward in the valleys of the Río Cauca and Río Magdalena in +Colombia (Fig. 1). Except for the central area of the Canal Zone +the subspecies is unknown from the Caribbean drainage in Central +America, but in Colombia the subspecies occurs only in the +Caribbean drainage. In Central America this frog occurs mostly on +the coastal lowlands; the highest recorded elevation is 560 meters +at El Valle, Coclé, Panamá. Throughout most of its range _Hyla +microcephala microcephala_ occurs in disturbed habitats--cut-over +forests, secondary growth, and pastureland. It does not seem to be +an inhabitant of either primary forest or of _Curatella_-savanna. + +_Specimens examined._--522, as follows: +Costa Rica+: Puntarenas: +Golfito, KU 32172-207; 3 km. E Golfito, KU 86399, USC 2757-8; +Palmar Sur, KU 64591-608, USC 2650 (14), UU 3907-32; *1.5-2.5 km. +ESE Palmar Sur, KU 68293-7 (skeletons), 93957-62; Parrita, USC +8254 (2), 8255, 8256 (4), 8258 (2) [intergrades with _H. m. +underwoodi_]; 3 km. NW Piedras Blancas, KU 103689; 6.1 km. NE +mouth of Río Tárcoles, USC 818 (2), 6081-2 [intergrades with _H. +m. underwoodi_]; Villa Neilly, USC 2651; *1-5 km. WNW Villa +Neilly, USC 6182-4, 8003 (4), 8031 (3), 8032; *10.5 km. WNW Villa +Neilly, KU 64609-27, 68398 (eggs). + ++Panama+: Canal Zone: Albrook Air Base, TNHC 23389, 23497; Balboa, +ANSP 19555-6; *Fort Clayton, UIMNH 42008-12; *2.8 km. SW Fort +Kobbe, KU 96015-25; *Frijoles, MCZ 19208; *Bamboa, MCZ 21507; *8.3 +km. N Gatún Locks, TNHC 23441; *Juan Diaz, MCZ 13747; *Juan Mina, +AMNH 55436-7, ANSP 21811-2, UMMZ 126734, 126735 (6), UU 3900-6; +*8-14 km. N Miraflores Locks, TNHC 23374-88, 23390-409, 23411-38, +23440, 23442-60, 23462-76; 23478-83, 23492, 23555-60, 23562-76; +*Río Chagres, AMNH 55430, 55439; *Río Cocolí, 3.5 km. N Miraflores +Locks, TNHC 23410; *Summit, ANSP 23365-71, FMNH 22966-9, KU +97783-87. Chiriqui: 5.5 km. E Concepción, AMNH 69772; *14.4 km. E +Concepción, AMNH 69773-8; 2 km. S David, AMNH 69779; *Progreso, +UMMZ 58252, 58253 (2), 58254, 58436; Río Gariché, 8.3 km. ESE Paso +Canoas, KU 103065-8. Coclé: 1 km. SE El Caño, KU 103042-51; El +Valle de Antón, AMNH 59614-18 (10), 69785, ANSP 23502-5, KU +77201-14, MVZ 66578-83, UIMNH 46532. Colón: Cement Plant, +Transisthmian Highway, FMNH 60394-5. Darién: El Real, KU 80454-5, +103052-64, UMMZ 125036 (10), USNM 140567-8; Río Canclon at Río +Chucunaque, UMMZ 125035; *Río Chucunaque, near Yavisa, AMNH 59523. +Los Santos: Tonosí, KU 101606-9. Panamá: 5 km. S Bejuco, AMNH +69782; 3 km. W Chepo, KU 77172-4, 104097-8 (tadpoles); *6 km. WSW +Chepo, KU 77175; *Chico, Río La Jagua, USNM 129070; *La Joya, +Cacora, ANSP 25129-33; Madden Dam, FMNH 67819; Nueva Gorgona, AMNH +69780-1; *1.6 km. W Nueva Gorgona, AMNH 69783-4; 1.5 km. W Pacora, +77176-200; *Río La Laja, near Chamé, ANSP 21845; *Río Tapia, MCZ +10048; *Tapia, AMNH 18930, 18950, 18952-3; *18 km. E Tocumen, MVZ +78662. + ++Colombia+: Chocó: Sautatá, Atrato, FMNH 74918 (2), 74919. +Magdalena: Aracataca, ANSP 19755-7; Curumani, MCZ 21465-74, UIMNH +28855; UMMZ 90168, USNM 118247; El Banco, Río Magdalena, ANSP +25061; Fundación, UMMZ 48281-2. Tolima: Espinal, MCZ 15068; +Mariquita, FMNH 81822-3. Valle: Sevilla, MCZ 13751-3. + + + + +_Hyla microcephala underwoodi_ Boulenger + + + _Hyla microcephala_ Boulenger, Proc. Zool. Soc. London, p. 481, + October 1, 1898 [Syntypes.--BMNH 94. 11. 1532-33 from Bebedero, + Guanacaste Province, Costa Rica; C. F. Underwood collector] (not + _Hyla microcephala_ Cope, Proc. Amer. Philos. Soc., 23:281, + February 11, 1886, from Chiriquí, Panamá). + + _Hyla underwoodi_ Boulenger, Ann. Mag. Nat. Hist., ser. 7, 3:277, + April, 1899 (substitute name for _Hyla microcephala_ Boulenger, + preoccupied). Günther, Biologia-Centrali Americana, Reptilia and + Batrachia, p. 278, September, 1901. Dunn and Emlen, Proc. Acad. + Nat. Sci. Philadelphia, 84:25, March 22, 1932. Stuart, Misc. Publ. + Mus. Zool., Univ. Michigan, 29:39, October 1, 1935. Taylor, Proc. + Biol. Soc. Washington, 50:44, April 21, 1937. Stuart, Occas. + Papers Mus. Zool., Univ. Michigan, 471:15, May 17, 1943. Taylor + and Smith, Proc. U. S. Natl. Mus., 95:586, June 30, 1945. Stuart, + Misc. Publ. Mus. Zool., Univ. Michigan, 69:35, June 12, 1948. + Smith and Taylor, Bull. U. S. Natl. Mus., 194:85, June 17, 1948; + Univ. Kansas Sci. Bull., 33:316, March 20, 1950. Stuart, Contr. + Lab. Vert. Biol., Univ. Michigan, 45:48, May, 1950. Taylor, Univ. + Kansas Sci. Bull., 35:891, July 1, 1952; Univ. Kansas Sci. Bull., + 39:25, November 18, 1958. + + _Hyla phlebodes_, Cole and Barbour, Bull. Mus. Comp. Zool., 50:154, + November, 1906. Kellogg, Bull. U. S. Natl. Mus., 160:172, + March 31, 1932. + + _Hyla microcephala martini_ Smith, Herpetologica, 7:187, December + 31, 1951 [Holotype.--UIMNH 20965 from Encarnacion, Campeche, + México; H. M. Smith collector]. Stuart, Contr. Lab. Vert. Biol., + Univ. Michigan, 68:46, November, 1954. Fugler and Webb, + Herpetologica, 13:105, July 10, 1957. Stuart, Contr. Lab. Vert. + Biol., Univ. Michigan, 75:17, June, 1958. Neill and Allen, Publ. + Research Div., Ross Allen's Reptile Inst., 2:26, November 10, + 1959. Duellman, Univ. Kansas Publ., Mus. Nat. Hist., 13:62, + August 16, 1960. Stuart, Herpetologica, 17:74, July 11, 1961. + Hensley and Smith, Herpetologica, 18:70, April 9, 1962. Stuart, + Misc. Publ. Mus. Zool., Univ. Michigan, 122:36, April 2, 1963. + Holman and Birkenholz, Herpetologica, 19:144, July 3, 1963. + Duellman, Univ. Kansas Publ., Mus. Nat. Hist., 15:225, October 4, + 1963; Univ. Kansas Publ., Mus. Nat. Hist., 15:588, June 22, 1965. + + _Hyla microcephala underwoodi_, Smith, Herpetologica, 7:188, + December 31, 1951. + + +_Diagnosis._--Brown lateral stripe narrow, extending to groin or +only to sacral region, bordered above by narrow white line; dorsal +pattern bold, consisting of X- or )(-shaped mark in scapular +region or pair of interconnected dark lines on back; interorbital +dark mark usually present; shanks usually having dark transverse +bars. + +_Description and Variation._--The dorsal color pattern is highly +variable. The various permutations of the X-shaped scapular mark +and dark sacral marks differ proportionately in different samples. +The variation in color pattern in 12 samples is summarized in +Table 2. In samples from the southern part of the range (southern +Nicaragua and Guanacaste Province, Costa Rica) more (40-93%) +individuals have the lateral stripes extending to the groin than +in northern samples (0-42%) from southern México and Guatemala. +Likewise, the percentage of specimens lacking bars on the shanks and +a dark interorbital bar is higher in the Costa Rican samples than +elsewhere in the range. The X- or )(-shaped scapular markings and +/\- or / \-shaped sacral markings are most prevalent in northern +samples, whereas to the south the dorsal markings are more commonly +arranged in a pattern of paired lines, which usually are discontinuous +and usually extend posteriorly only to the sacral region. Thus, the +color pattern in _H. m. underwoodi_ in the southern part of its range +shows trends towards the pattern characteristic of _H. m. +microcephala_. Intergrades between these two subspecies have +been discussed in the account of the nominate subspecies. + + + + + Table 2.--Variation in Color Pattern in Hyla microcephala underwoodi +========================================================================== + Population | N | Shanks || Interorbital || Dorsolateral | + | | || bar || stripe | + | |-------------||----------------||--------------| + | | Bars |Flecks|| Present| Absent|| Groin| Sacrum| +-------------------------------------------------------------------------- +Oaxaca: | 27 | 22 | 5 || 27 | 0 || 0 | 27 | + Donají-Sarabia | | | || | || | | + | | | || | || | | +Tabasco: | 55 | 46 | 9 || 55 | 0 || 0 | 55 | + Teapa-Villahermosa| | | || | || | | + | | | || | || | | +Guatemala: | 51 | 51 | 0 || 51 | 0 || 17 | 34 | + La Libertad | | | || | || | | + | | | || | || | | +Guatemala: | 32 | 32 | 0 || 32 | 0 || 0 | 32 | + Finca Chamá | | | || | || | | + | | | || | || | | +Guatemala: | 31 | 31 | 0 || 31 | 0 || 14 | 17 | + Puerto Barrios | | | || | || | | + | | | || | || | | +Honduras: | 13 | 13 | 0 || 13 | 0 || 9 | 4 | + Lago Yojoa | | | || | || | | + | | | || | || | | +Nicaragua: | 56 | 44 | 12 || 54 | 2 || 13 | 43 | + La Cumplida | | | || | || | | + | | | || | || | | +Nicaragua: | 10 | 10 | 0 || 10 | 0 || 8 | 2 | + Tipitapa | | | || | || | | + | | | || | || | | +Nicaragua: | 10 | 10 | 0 || 10 | 0 || 8 | 2 | + Santo Thomás | | | || | || | | + | | | || | || | | +Costa Rica: | 12 | 0 | 12 || 6 | 6 || 7 | 5 | + Tenorio-Tilarán | | | || | || | | + | | | || | || | | +Costa Rica: | 38 | 21[A]| 15 || 34 | 4 || 25 | 13 | + Las Cañas-Liberia | | | || | || | | + | | | || | || | | +Costa Rica: | 32 | 26 | 6 || 29 | 3 || 30 | 2 | + Esparta | | | || | || | | +-------------------------------------------------------------------------- + +========================================================================== + Population | Scapular markings || Sacral | + | || markings | + |----------------------------||----------------------| + | X | )( | ][ | Other || /\ | / \ | Other | +-------------------------------------------------------------------------- +Oaxaca: | 23 | 4 | 0 | 0 || 7 | 6 | 14 | + Donají-Sarabia | | | | || | | | + | | | | || | | | +Tabasco: | 53 | 2 | 0 | 0 || 19 | 11 | 23 | + Teapa-Villahermosa| | | | || | | | + | | | | || | | | +Guatemala: | 45 | 6 | 0 | 0 || 16 | 14 | 21 | + La Libertad | | | | || | | | + | | | | || | | | +Guatemala: | 32 | 0 | 0 | 0 || 26 | 2 | 4 | + Finca Chamá | | | | || | | | + | | | | || | | | +Guatemala: | 23 | 0 | 4 | 4 || 6 | 4 | 21 | + Puerto Barrios | | | | || | | | + | | | | || | | | +Honduras: | 3 | 2 | 3 | 5 || 2 | 1 | 10 | + Lago Yojoa | | | | || | | | + | | | | || | | | +Nicaragua: | 11 | 35 | 8 | 2 || 0 | 19 | 37 | + La Cumplida | | | | || | | | + | | | | || | | | +Nicaragua: | 0 | 5 | 3 | 2 || 0 | 3 | 7 | + Tipitapa | | | | || | | | + | | | | || | | | +Nicaragua: | 3 | 0 | 7 | 0 || 0 | 5 | 5 | + Santo Thomás | | | | || | | | + | | | | || | | | +Costa Rica: | 0 | 0 | 12 | 0 || 0 | 0 | 12 | + Tenorio-Tilarán | | | | || | | | + | | | | || | | | +Costa Rica: | 0 | 11 | 19 | 8 || 0 | 0 | 38 | + Las Cañas-Liberia | | | | || | | | + | | | | || | | | +Costa Rica: | 0 | 0 | 14 | 18 || 0 | 0 | 32 | + Esparta | | | | || | | | +-------------------------------------------------------------------------- + +[Footnote A: Longitudinal stripes present in two specimens.] + + + +When this frog is active at night its dorsum is pale yellow; faint +flecks are present in some individuals. The white dorsolateral line +usually is evident in the tympanic region, but in many individuals a +dorsal pattern of lines and other marks is not evident. By day the +dorsum changes to yellowish tan or pale brown with dark brown or +reddish brown markings (Pl. 13). The venter is white, and the vocal +sac in breeding males is yellow. The iris is pale bronze with a brown +tint anterior and posterior to the pupil. + +_Remarks._--_Hyla microcephala underwoodi_ has had a confused +nomenclatural history. The taxon was first named _Hyla microcephala_ +by Boulenger (1898); this name was preoccupied by _Hyla microcephala_ +Cope (1886). Cole and Barbour (1906) and Kellogg (1932) used the name +_Hyla phlebodes_ Stejneger (1906) for specimens of this frog from +México. Dunn (1931, 1933, 1934) applied the name _Hyla underwoodi_ to +Panamanian specimens that we identify as _Hyla phlebodes_. Smith +(1951) named _Hyla microcephala martini_ from southern México and +Guatemala and considered the northern populations to represent a +subspecies distinct from the Costa Rican _Hyla microcephala +underwoodi_, despite the fact the Stuart (1935:39) stated that +comparisons of specimens from El Petén, Guatemala, with the holotype +of _Hyla underwoodi_ showed only trivial differences. + +Much of the confusion regarding the name _Hyla underwoodi_ stems from +the illustration given by Boulenger (1898:pl. 39, fig. 3) and +reproduced by Taylor (1952:892), which shows a frog having a unicolor +dorsum, dorsolateral white lines, and dark flanks. This pattern is in +marked contrast to the pattern seen in most preserved specimens, which +have the dorsum variously marked by dark brown lines or irregular +marks. Smith (1951:185), in his description of _Hyla microcephala +martini_ from southern México, considered _H. underwoodi_ to be a +subspecies of _H. microcephala_ that lacked dorsal dark markings. + +Data accumulated in 1961 through field studies by the senior author at +the type locality, Bebedero, and other localities in Guanacaste and +Puntarenas provinces in Costa Rica provide a reasonable explanation of +the differences in color pattern. As noted in the preceding description +of this subspecies, at night the dorsal markings are not evident in +many living individuals, whereas by day the dorsal markings are +prominent. Most collectors prepare their specimens by day; consequently +the majority of specimens have a pronounced dorsal pattern. Of the +frogs collected in Costa Rica in 1961, some specimens were preserved +at night; others from the same series were preserved by day. The +differences are striking. In those preserved at night, dorsal markings +are faint, if present at all. Some specimens closely match the figure +given by Boulenger (1898). + +It is extremely doubtful if the frog described and illustrated by +Boulenger could be associated with either _Hyla phlebodes_ or _H. +microcephala microcephala_. Individuals of the former species lack +a dorsolateral white line and always have some dorsal markings +evident at night; furthermore, _H. phlebodes_ is not known to occur +on the Pacific lowlands. _Hyla microcephala microcephala_ occurs +farther southeast. Since there is no reason to doubt the type locality +of _H. underwoodi_, since specimens from the area around the type +locality that have been preserved at night are like the holotype in +pattern, and since the characteristics of the populations of the frogs +in Guanacaste are the same as, or gradually blend into those of, +populations in northern Central America and southern México, the +frogs from throughout the entire range can be referred to one taxon, +the earliest name for which is _Hyla underwoodi_ Boulenger, which +herein is considered to be a subspecies of _H. microcephala_ Cope. + +_Distribution._--_Hyla microcephala underwoodi_ inhabits the +Atlantic slopes and lowlands from southern Veracruz and extreme +northern Oaxaca eastward across the base of the Yucatan Peninsula +(possibly the species is extant in the northern part of the +peninsula) to British Honduras and thence southeastward through +the Caribbean lowlands and interior valleys in Honduras to central +Nicaragua, where it apparently avoids the forested Caribbean +lowlands and the dry Pacific lowlands of northwestern Nicaragua, +but in the vicinity of Managua invades the Pacific lowlands and +continues southward into northwestern Costa Rica as far as the +Puntarenas Peninsula (Fig. 1). In México and Guatemala the species +has not been taken at elevations of more than 350 meters, whereas +farther south it occurs at higher elevations--780 meters at +Silencio, Costa Rica, 830 meters on Montaña de Guaimaca, Honduras, +960 meters at Finca Tepeyac, Nicaragua, and 1200 meters at Finca +Venecia, Nicaragua. + +_Specimens examined._--1270, as follows: +Mexico+: Campeche: Balchacaj, +FMNH 100406, UIMNH 20944-6; Encarnación, FMNH 27069-70, 75784, +MCZ 28360, 29637, UIMNH 20948-58, 20965, USNM 134264-5; Escárcega, +UMMZ 122999; *7.5 km. W Escárcega, KU 71229-43; Laguna Alvarado, 65 km. +S Xpujil, KU 75084-9; Pacaitún, Río Candelaria, FMNH 83118-20; +*Tres Brazos, FMNH 113101-22, UIMNH 20947; 10 km. W Xpujil, KU 75082-3. +Chiapas: Palenque, UIMNH 47984, 49139-50, USNM 114973-8. Oaxaca: *5 km. +N Chiltepec, KU 87015-23; 3 km. N Donají UMMZ 115249 (9); *3.7 km. +N Donají, UMMZ 115250 (5); *43 km. N Matías Romero, UIMNH 42550-68; +*3.5 km. N Palomares, TNHC 25185, 25321-31, 25341-68; 4.6 km. N +Sarabia, UMMZ 115247 (2); *6.1 km. N Sarabia, UMMZ 115248 (11), *3 km. +N Tolocita, KU 39655; Tuxtepec, KU 87024-40. Tabasco: 24 km. N +Frontera, MCZ 35665-70; 0.8 km. E Río Tonolá, TNHC 25189; Teapa, UMMZ +119218 (4); *2.7 km. N Teapa, UMMZ 119216 (4); *10 km. N Teapa, UMMZ +119217 (6); *11.5 km. N Teapa, UMMZ 119219; *15.2 km. N Teapa, UMMZ +119220 (4); *17.6 km. N Teapa, UMMZ 119221 (12), 3.3 km. S Villahermosa, +UMMZ 119215 (12), *17.6 km. S Villahermosa, UMMZ 119214 (12). +Veracruz: 2.1 km. N Acayucan, UIMNH 42547-9; *6.4 km. NW Acayucan, +UMMZ 115254 (14); 1.6 km. ESE Alvarado, UMMZ 115258 (39); *2.4 km. ESE +Alvarado, UMMZ 115251 (2); *4.5 km. S Aquilera, UMMZ 115252 (21); +*8 km. SW Coatzacoalcos, UMMZ 119213 (10); 2.2 km. E Cosoleacaque, +UMMZ 119222 (26); 10 km. SE Hueyapan, UMMZ 115255; 0.8 km. S Lerdo de +Tejada, UMMZ 122778; *3.6 km. NE Minatítlán, TNHC 25150-2; 1.9 km. S +Naranja, UMMZ 115253 (3); 4.5 km. NE Novillero, UMMZ 115256; San Andrés +Tuxtla, FMNH 113124-8, UIMNH 20942-3. Yucatán: Chichén-Itzá, FMNH 36570, +MCZ 2463 (2). + ++British Honduras+: Cayo: 6.2 km. S El Cayo, MCZ 37885-92. Stann Creek: +Stann Creek, FMNH 49068. + ++Guatemala+: Alta Verapaz: 28.3 km. N Campur, KU 64578-90; Chinajá, +KU 57425; Cubilquitz, UMMZ 90887, 90888 (4); Finca Chamá, UMMZ 90879 +(13), 90880 (4), 90881, 90882 (28), 90883 (12), 90884 (46), 90885 (39), +90886 (20); *Finca Tinaja, BYU 16032; Panzós, UMMZ 90889 (2). +Chiquimula: Chiquimula, UMMZ 98113; 2 km. N Esquipulas, UMMZ 106844. +El petén: La Libertad, KU 57447-97, 59907-11 (skeletons), MCZ 21461, +UMMZ 75332 (13), 75333 (11), 75334 (14), 75335 (10); Piedras Negras, +FMNH 113123, UIMNH 20966; *5 km. S Piedras Negras, USNM 114951-72; +Tikal, UMMZ 117981 (2); Toocog, 15 km. SE La Libertad, KU 57426-46. El +Quiché: Finca Tesoro, UMMZ 89165 (5). Huehuetenango: Finca San Rafael, +16 km. SE Barillas, FMNH 40917-9. Izabal: Puerto Barrios, FMNH +20004-7; 8 km. S Puerto Barrios, KU 57507-37, 59991 (eggs), 59992 +(tadpoles); Quirigua, CAS 69657-701; 2.5 km. NE Río Blanco, KU 57539; +San Felípe, FMNH 35065. Zacapa: 14 km. ENE Mayuelas, KU 57502-6; 8 km. +ENE Río Hondo, KU 57498-501. + ++Honduras+: Atlantidad: La Ceiba, UMMZ 91948 (2), USNM 117593-600; +Lancetilla, MCZ 17981. Cortes: Lago Yojoa, AMNH 54917-9, 54957, 55134, +KU 64563-77. El Paraiso: Valle de Jamastran, AMNH 54807-12. +Francisco-Moranza: El Zamorano, AMNH 54873-81, KU 103223, UMMZ 123101; +Montaña de Guaimaca, AMNH 54900-4 (8); Ranch San Diego, 19 km. SW +Guaimaca, AMNH 53939. Itibucá: Vieja Itibucá, AMNH 54912-3. + ++Nicaragua+: Chontales: 3 km. SW Santo Tomás, KU 64770-9, 68308 +(skeleton). Esteli: Finca Venecia, 7 km. N, 16 km. E Condega, KU +85296; 2.4 km. N Estelí, MCZ 28933-7. MANAGUA: 12-13 km. E Managua, +KU 85297-301; *10 km. SW Tipitapa, UMMZ 119977 (10). Matagalpa: *Finca +Tepeyac, 10.5 km. N, 9 km. E Matagalpa, KU 85302-3; Hacienda La +Cumplida, KU 64780-96, 68309-11 (skeletons), UMMZ 116482 (8), 116483 +(23), 116484 (3), 116485 (5), 119984 (3). Rivas: *Finca Amayo, 13 km. +S, 14 km. E Rivas, KU 85304-7; 16 km. S Rivas, MCZ 29011-7; *20.5 km. +SE Rivas, KU 85308-10; 5 km. SE San Pablo, KU 43111-4. + ++Costa Rica+: Guanacaste: Arenal, USC 6254 (2); *3 km. W Bagaces, USC +7019 (10); *3 km. NE Boca del Barranca, USC 8017 (21), *Finca San +Bosco, USC 6272 (6), 6276 (3); *Guayabo de Bagaces, USC 7022 (4), 7023 +(3), 7025; 12 km. S La Cruz, USC 8091 (2); *Laguna Arenal, USC 6262; +*27 km. N Las Cañas, USC 8171 (6); *16 km. E Las Cañas, KU 102252-8; +16 km. SSE Las Cañas, KU 65090-5; *20 km. SE Las Cañas, KU 102251; +Liberia, KU 30827-39; *7.3 km. N Liberia, USC 8096 (4); *10 km. N +Liberia, USC 8085 (9); *7.5 km. SE Liberia, KU 65102-8, 68621-2 +(skeletons); *14.7 km. S Liberia, USC 8238 (3); *4 km. W Liberia, KU +36847-57; 2 km. S Nicoya, USC 8230; *3-10 km. ESE Playa del Coco, USC +8012 (16), 8137 (14); *21.6 km. ESE Playa del Coco, USC 8138 (13); +*Peñas Blancas, KU 102247-50; *Río Bebedero, 5 km. S Bebedero, KU +65089; *Río Higuerón, USC 7168 (2); Santa Cruz, USC 8232 (2); +*Silencio, USC 6248; *Tenorio, KU 32313; Tilarán, KU 36858-60; *2 km. +E Tilarán, KU 86403, *5 km. NE Tilarán, KU 36840-6 USC 6269. +Puntarenas: Barranca, KU 32305-12, *5 km. WNW Barranca, UMMZ 119976 +(2); *10 km. E Esparta, KU 86400-2; 1 km. WNW Esparta, KU 65101; *4 km. +WNW Esparta, KU 65088; *10 km. WNW Esparta, KU 65063-87, 68616-20 +(skeletons); *12 km. WNW Esparta, KU 65096-100, USC 8251; 21.8 km. +W San Ramón, USC 8242 (15). + + + + ++Hyla robertmertensi+ Taylor + + + _Hyla robertmertensi_ Taylor, Proc. Biol. Soc. Washington, 50:43, + April 21, 1937 [Holotype.--CNHM 100096 (formerly EHT-HMS 2270) + from Tapachula, Chiapas, México; H. M. Smith and E. H. Taylor + collectors]. Smith and Taylor, Bull. U. S. Natl. Mus., 194:84, + June 17, 1948; Univ. Kansas Sci. Bull., 33:326, March 20, 1950. + Mertens. Senckenbergiana, 33:170, June 15, 1952; + Senckenbergischen Naturf. Gesell., 487:30, December 1, 1952. + Stuart, Contr. Lab. Vert. Biol., Univ. Michigan, 68:47, + November, 1954. Duellman, Univ. Kansas Publ., Mus. Nat. Hist., + 13:63, August 16, 1960. Duellman and Hoyt, Copeia, 1961 (2): 417, + December 19, 1961. Porter, Herpetologica, 18:168, October 17, + 1962. Stuart, Misc. Publ. Mus. Zool., Univ. Michigan, 122:36, + April 2, 1963. Duellman and Trueb, Univ. Kansas Publ., Mus. Nat. + Hist., 17:348, July 14, 1966. + + +_Diagnosis._--Brown lateral stripe wide, including loreal region and +entire tympanum, extending to groin, bordered above by narrow white +line; dorsum unicolor or with pair of dark lines (or rows of dashes) +usually extending only to the sacral region; shanks having dark flecks, +no transverse bars; interorbital bar lacking. + +_Description and Variation._--Males attain a maximum snout-vent length +of 26.4 mm. in Oaxaca, whereas in a sample from Acacoyagua, Chiapas, +the largest male has a snout-vent length of 25.7 mm., and from La +Trinidad, Guatemala, 24-6 mm. Specimens from the western part of the +range (eastern Oaxaca) have slightly smaller heads and proportionately +larger tympani than the more eastern populations (Table 1). + +The color pattern shows little variation, except in the nature of the +dorsal markings. In a few specimens from throughout the range, but +especially in the eastern part of the range, the dorsum lacks markings +between the dorsolateral white lines. In most specimens the dorsal +pattern consists of flecks or dashes arranged in two parallel +longitudinal rows, and in some specimens the marks are fused into +parallel lines. Small brown flecks are present on the dorsal surfaces +of the shanks; in some specimens these flecks tend to form a +longitudinal stripe on the shank. An interorbital dark mark is +invariably absent. + +When active at night _Hyla robertmertensi_ is pale yellow above with a +white dorsolateral line and pale brown lateral stripe; the dorsal +markings are faint. By day the dorsum is yellowish tan with brown +markings. The dorsolateral stripe is creamy white, and the lateral +stripe is dark brown (Pl. 14). The venter is white, and the iris is +dull bronze. In breeding males the vocal sac is yellow. + +_Remarks._--Although this species superficially resembles _Hyla +microcephala microcephala_, the latter is easily distinguished by the +narrow brown lateral stripe, as compared with the much wider stripe +in _H. robertmertensi_. No other hylids in northern Central America +and southern México can be confused with this species. + +_Distribution._--_Hyla robertmertensi_ inhabits the Pacific slopes (to +elevations of 700 meters) and lowlands from eastern Oaxaca (east of +the Plains of Tehuantepec) southeastward to central El Salvador. The +species also occurs in the Cintalapa Valley (Atlantic drainage) in +southwestern Chiapas (Fig. 2.) The distribution seems to be limited on +the northwest and southeast by arid environments. The region in which +_Hyla robertmertensi_ lives is characterized by higher rainfall and +more luxuriant vegetation than occur on the Plains of Tehuantepec or +on the Pacific lowlands of eastern El Salvador and southern Honduras. +In addition to the localities listed below, Mertens (1952:30) +recorded the species from Hacienda Cuyan-Cuya, Depto. Sonsonate, El +Salvador. + + + [Illustration: Fig. 2. Map showing locality records for + _Hyla robertmertensi_.] + + +_Specimens examined._--490, as follows: +Mexico+: Chiapas: Acacoyagua, +USNM 114754-61; *2 km. W Acacoyagua, UMMZ 87843 (28), 87844 (50), +87845 (50), 87846 (45), 87847 (27), 87848 (3); 32 km. N Arriaga, KU +57619-24, 59917-8 (skeletons); Asunción, FMNH 100413, 100501-4, UIMNH +26989-90, USNM 134267; *La Esperanza, USNM 114737-48, 114750-3, 17 km. +S Las Cruces, KU 57625-49, 59997 (eggs); 8.5 km. N Puerto Madero, UMMZ +119981 (2); *11.7 km. N Puerto Madero, UMMZ 119982; Tapachula, FMNH +100096, UIMNH 26987; *11 km. S Tapachula, KU 57605-18, 59916 (skeleton); +Tonolá, FMNH 27073, 100505-10, UIMNH 26988. Oaxaca: Tapanatepec, +UMMZ 115245 (2), *1.6 km. E Tapanatepec, UMMZ 115244 (14); *4.3 km. +E Tapanatepec, UIMNH 38368-9; *7.5 km. W Tapanatepec, UMMZ 115246 +(39); 12.8 km. W Tapanatepec, KU 65007-14; 7.2 km. WNW Zanatepec, +UMMZ 115243 (77); *13.6 km. WNW Zanatepec, TNHC 25213-22; 22.7 km. +WNW Zanatepec, TNHC 25203-9. + ++Guatemala+: Jutiapa: Jutiapa, UMMZ 106848; La Trinidad, UMMZ +107733 (23). Retalhueleu: Casa Blanca, UMMZ 107732. + ++El Salvador+: La Libertad: 16 km. NW Santa Tecla, KU 44112. San +Salvador: 21.9 km. N San Salvador, UMMZ 119983 (6). + + + + ++Hyla phlebodes+ Stejneger + + + _Hyla phlebodes_ Stejneger, Proc. U. S. Natl. Mus., 30:817, June 4, + 1906 [Holotype.--USNM 2997 from "San Carlos," Costa Rica; + Burgdorf and Schild collectors]. Taylor, Proc. Biol. Soc. + Washington, 50:44, April 21, 1937; Univ. Kansas Sci. Bull., + 35:888, July 1, 1952; Univ. Kansas Sci. Bull., 39:25, November + 18, 1958. Fouquette, Evolution, 14:484, December 16, 1960. + Duellman and Trueb, Univ. Kansas Publ., Mus. Nat. Hist., 17:348, + July 14, 1966. + + _Hyla underwoodi_, Dunn, Occas. Papers Boston Soc. Nat. Hist., 5:413, + October 10, 1931; Occas. Papers Boston Soc. Nat. Hist. 8:72, + June 7, 1933; Amer. Mus. Novitiates, 747.2, September 17, 1934, + Gaige, Hartweg, and Stuart, Occas. Papers Mus. Zool., Univ. + Michigan, 357:5, October 26, 1937. Breder, 1946, Bull. Amer. Mus. + Nat. Hist., 86:416, August 22, 1946. + + +_Diagnosis._--Dark brown lateral stripe, if present, usually extending +only to insertion of forearm, never posteriorly to sacral region; +white line above brown stripe absent or faint; dorsal pattern weak, +usually consisting of irregular dashes or interconnected lines; +interorbital dark mark present; shanks having weakly defined +transverse bars. + +_Description and variation._--In the majority of specimens (70%) the +lateral dark stripe extends from the nostril to the eye and thence +above the tympanum to a point above the insertion of the arm; in 17 +per cent the stripe extends to the mid-flank, whereas in 13 per cent +the stripe is absent. A narrow and faint white line is present on the +canthus in some specimens, but no distinct white stripe is present +above the lateral dark line posterior to the eye. An interorbital bar +and transverse marks on the shanks are invariably present. The dorsal +markings are variable, but in most specimens (92%) consist of either +an X- or )(-shaped mark in the scapular region; in the other specimens +the markings are irregular short lines or absent. Approximately equal +numbers of specimens have a transverse bar, chevron, or broken lines +in the sacral region, whereas about eight per cent of the specimens +lack markings in the sacral region. + +When active at night, individuals are pale yellowish tan with faint +brown dorsal markings. By day they are tan with more distinct brown +markings (Pl. 14). The thighs are pale yellow; the belly is white. The +iris is pale creamy tan with brown flecks. In breeding males the vocal +sac is yellow. + +_Tadpoles._--Tadpoles of this species have been found in an extensive +grassy pond at Puerto Viejo, Costa Rica. The following description is +based on KU 104099, a specimen in development stage 36 (Gosner, 1960). + +Total length, 21.0 mm.; body length, 6.7 mm.; body slightly wider than +deep, snout pointed; nostrils large, directed anteriorly, situated +near end of snout; eyes small, situated dorsolaterally, directed +laterally; spiracle sinistral, located just posteroventral to eye; +anal tube dextral. Tail xiphicercal; caudal musculature moderately +deep, extending far beyond posterior edge of fins; fins deepest at +about midlength; dorsal fin extending onto body, slightly deeper than +caudal musculature; ventral fin slightly shallower than musculature. +Mouth small, terminal, lacking teeth and fringing papillae, but having +finely serrate beaks. In preservative top of head olive-tan with brown +flecks; dark stripe from snout through eye to posterior edge of body; +belly white, flecked with brown anteriorly; tail creamy tan with +grayish brown blotches. In life, dorsum of body reddish tan mottled +with darker brown; lateral stripe dark brown; belly white, mottled +with brown and black; caudal musculature heavily pigmented with +grayish tan; posterior tip of tail marked with dark gray; caudal fins +heavily blotched with grayish tan; iris orange-tan peripherally, red +centrally (Pl. 15). + +_Remarks._--This species has been confused with _Hyla microcephala +underwoodi_ by many workers. Dunn (1931, 1933, 1934) and Breder (1946) +referred Panamanian specimens of _H. phlebodes_ to _H. underwoodi_; +likewise, Gaige, Hartweg, and Stuart (1937) made the same error. Cole +and Barbour (1906) and Kellog (1932) used the name _H. phlebodes_ for +Mexican specimens of _H. microcephala underwoodi_. The similarity in +color pattern of _H. microcephala underwoodi_ and _H. phlebodes_ +easily accounts for the misapplication of names. Although both species +have nearly identical dorsal color patterns, that of _H. microcephala +underwoodi_ usually is bolder. Furthermore, in that species a narrow +white line usually is present above the well-defined lateral dark +stripe, whereas the lateral dark stripe is short and posterior to the +eye is not bordered above by a white line in _H. phlebodes_. + +The type locality "San Carlos, Costa Rica" given by Stejneger +(1906:817) apparently refers to a region, the Llanuras de San Carlos, +in the northern part of Alajuela Province, Costa Rica. + + + [Illustration: Fig. 3. Map showing locality records for + _Hyla phlebodes_.] + + +_Distribution._--_Hyla phlebodes_ inhabits humid tropical forests from +southeastern Nicaragua southeastward on the Caribbean slopes and +lowlands to the Canal Zone in Panamá, thence eastward in the Chucunaque +Basin of eastern Panamá and onto the Pacific lowlands of Colombia +(Fig. 3). The species also reaches the Pacific slopes in the Arenal +Depression in northwestern Costa Rica and in the Panamanian isthmus, +where it occurs in humid forests on the Pacific slope of El Valle and +Cerro La Campana. Mostly the species is found at low elevations, but +it occurs at 600 meters at Turrialba and at 700 meters at Finca San +Bosco in Costa Rica. + +_Specimens examined._--410, as follows: +Nicaragua+: Zelaya: Isla +Grande del Maíz, MCZ 14848; Río Mico, El Recrero, UMMZ 79720 (6). + ++Costa Rica+: Alajuela: 12.4 km. N Florencia, MVZ 76108-10, USC 2628; +*Las Playuelas, 11 km. S Los Chiles, USC 7216; Los Chiles, USC 7217, +7219; 3 km. NE Muelle de Arenal, USC 2644 (2); *"San Carlos," USNM +29970. Cartago: Chitaría, KU 103690; *1.6 km. E Río Reventazón Bridge, +east of Turrialba, UMMZ 119978 (2); *Tunnel Camp, near Peralta, +KU 32456, 32458-69, 41098 (skeleton); Turrialba, FMNH 101794, 103188-9, +KU 25725-9, 32439-48, 41095-7 (skeletons), 64797-827, 68300-2 +(skeletons), 68403 (eggs), 68404 (tadpoles), MCZ 29224-5, 29310-2, +UMMZ 119979 (6), USC 31, 256 (2), 458 (2), 580, 594, 599 (7), 7074 (2), +USNM 29933. Guanacaste: Arenal, USC 6254; *Finca San Bosco, USC 62724, +6276 (3), Guayabo de Bagaces, USC 7022 (3), 7023; *Laguna Arenal, +USC 6262 (4); 3 km. NE Tilarán, USC 524; *5 km. NE Tilarán, USC 6269; +*6 km. NE Tilarán, UMMZ 122653 (6), S-2680 (skeleton), USC 523 (8). +Heredia: Puerto Viejo, KU 64828-63, 68303-7 (skeletons), 68405-6 +(tadpoles), 104099-100 (tadpoles); *1.5 km. N Puerto Viejo, KU 64871; +*1 km. S Puerto Viejo, KU 86432-40; *4.2 km. W Puerto Viejo, KU +64864-5; *5.9 km. W Puerto Viejo, KU 64866-70; *7.5 km. W Puerto Viejo, +KU 86431. Limón: Batán, UMMZ 119980 (2); La Castilla, ANSP 23707; +Puerto +Limón+, KU 32449-55. + ++Panama+: Bocas del Toro: 3.2 km. NW Almirante, KU 96026; Cayo de +Agua, KU 96027-31; Fish Creek, KU 96032-4. Canal Zone: Barro Colorado +Island, AMNH 69790, ANSP 23244-50; FMNH 13380, 22972-4; Juan Mina, +AMNH 55429, UU 3899; *8.6-13.8 km. N Miraflores Locks, TNHC 23439, +23477, 23484-8, 23491, 23494-9, 23501-2, 23504-8, 23510-17, 23519-30, +23532-8, 23541-54, 23561. *Rio Chagres, AMNH 55431-4; Río Cocolí, +3.5 km. N Miraflores Locks, TNHC 23461, 23489-90, 23493, 23500, 23503, +23509, 23518, 23531, 23539-40; *Summit, ANSP 23361, KU 97788; *Three +Rivers Plantation, SU 2130. Coclé: El Valle de Antón, AMNH 55435, +69786-9, ANSP 23506-9. Colón: Achiote, KU 77215-78; Ciricito, CAS +71499-500, 71505-6. Darién: Río Canclon at Río Chucunaque, UMMZ 126733; +Río Chucunaque, near Yavisa, AMNH 51783. Panamá: Cero La Campana, FMNH +67847-50. + ++Colombia+: Chocó: Andagoya, FMNH 81856; Boca de Raspadura, AMNH +13570-8. + + + + ++Hyla sartori+ Smith + + + _Hyla underwoodi_ (in part), Smith and Taylor, Bull. U. S. Natl. + Mus., 194:85, June 17, 1948. + + _Hyla microcephala sartori_ Smith, Herpetologica, 7:186, + December 31, 1951 [Holotype.--UIMNH 20934 from 1 mile north of + Organos, south of El Treinte, Guerrero, México; H. M. Smith and + E. H. Taylor collectors]. Duellman, Univ. Kansas Publ., Mus. Nat. + Hist., 15:124, December 20, 1961. Porter, Herpetologica, 18:168, + October 17, 1962. Davis and Dixon, Herpetologica, 20:230, + January 25, 1965. Duellman, Univ. Kansas Publ. Mus. Nat. Hist., + 15:652, December 30, 1965. + +_Diagnosis._--Dorsum tan with broad dark brown chevrons or transverse +bars; shanks marked with two or three broad transverse bars; +dorsolateral stripes absent. + +_Description and variation._--No noticeable geographic variation is +apparent in either structural features or coloration in this species. +All specimens lack a dorsolateral dark stripe and white line, although +a dark line is present on the canthus and dissipates in the loreal +region. A broad interorbital brown bar is present in all specimens. +The color pattern on the dorsum invariably consists of a broad, dark, +chevron-shaped mark in the scapular region and a broad dark chevron or +transverse bar in the sacral region. The shanks invariably have two or +three dark brown transverse bars. + +When active at night individuals are yellowish tan above with chocolate +brown markings (Pl. 14). The belly is white, and the thighs are pale +yellowish tan. The iris is dark bronze-color. In breeding males the +vocal sac is yellow. By day some individuals were observed to change +to creamy gray with distinct darker markings. + +_Remarks._--Although tadpoles of this species have not been found, +observations on the breeding sites indicate that the tadpoles probably +develop in ponds. Except for calling males observed around a pool in a +stream-bed 11.8 kilometers west-northwest of Tierra Colorada, Guerrero, +all breeding congregations have been found at temporary ponds. + +Smith (1951:186) named _Hyla sartori_ as a subspecies of _Hyla +microcephala_. This subspecific relationship seemed reasonable until +analysis of the mating calls showed that the call of _H. sartori_ is +more nearly like that of _H. phlebodes_ than that of _H. microcephala_. +The broad hiatus separating the ranges of _H. microcephala_ and _H. +sartori_ is additional evidence for considering _H. sartori_ as a +distinct species. + + + [Illustration: Fig. 4. Map showing locality records for + _Hyla sartori_.] + + +_Distribution._--_Hyla sartori_ occurs in mesophytic forests to +elevations of about 300 meters on the Pacific slopes of southern México +from southwestern Jalisco to south-central Oaxaca (Fig. 4). The lack of +specimens from Colima and Michoacán probably reflects inadequate +collecting instead of the absence of the species there. On the basis of +available habitat the species would be expected to occur in Nayarit, +but extensive collecting there has failed to reveal its presence. The +semi-arid Plains of Tehuantepec apparently limit the distribution to +the east. + +_Specimens examined._--190, as follows: +México+: Guerrero: 5 km. E +Acapulco, AMNH 54611-2; 23.2 km. N Acapulco, UIMNH 26404-7; Colonia +Buenas Aires, 23 km. E Tecpán de Galeana, UMMZ 119223 (7); *El +Limoncito, FMNH 75785, 100390-402, 104631, 104633, UMMZ 117250, USNM +134266; El Treinte, FMNH 100403, UIMNH 20935-7; Laguna Coyuca, AMNH +59686; La Venta, MCZ 29635; *Morjonares, UIMNH 26392-402; 1.6 km. +N Organos, FMNH 100404-5, UIMNH 20933-4; 19.2 km. S Petaquillas, +UIMNH 26408; 6.1 km. E. Tecpán de Galeana, TNHC 23396-408; *11.2 km. +N Tierra Colorada, UIMNH 26403; 11.8 km. WNW Tierra Colorada, UMMZ +119225 (51), S-2677-9 (skeletons); Zacualpán, UMMZ 119224 (6). Jalisco: +6.4 km. NE La Resolana, KU 67853-69; 24 km NE La Resolana, KU 67870-3. +Oaxaca: 3 km. N Pochutla, KU 57539; 13.4 km. N Pochutla, UMMZ +123495 (40). + + + + +CRANIAL OSTEOLOGY + + +The frogs of the _Hyla microcephala_ group have a minimal amount +of cranial ossification as compared to more generalized hylid skulls, +such as _Smilisca_ (Duellman and Trueb, 1966). In the _Hyla +microcephala_ group the sphenethmoid is small and short, and a large +frontoparietal fontanelle is present. The quadratojugal exists only +as a small spur and is not in contact with the maxillary. The proötics +are poorly developed. The anterior and posterior arms of the squamosal +are short; the anterior arm extends no more than one-fourth of the +distance to the maxillary, and the posterior arm does not have a bony +connection with the proötic. The nasal lacks a maxillary process, and +the medial ramus of the pterygoid lacks a bony connection to the +proötic. + +Teeth are absent on the parasphenoid and palatines, but present on the +maxillaries, premaxillaries, and prevomers. The teeth are simple, +pointed, and slightly curved. Although the number of teeth varies +(Table 3), no consistent differences between the species are apparent. + + +Table 3.--Variation in the Number of Teeth in the Species of the Hyla + Microcephala Group. (N=Number of Jaws, or Twice the Number of + Individuals; Means are Given in Parentheses After the Observed + Ranges). + +========================+====+=============+==============+========== + Species | N | Maxillary | Premaxillary | Prevomer +------------------------+----+-------------+--------------+---------- +_H. microcephala_ | 32 | 31-47(37.8) | 4-13(8.9) | 2-4(3.2) + | | | | +_H. phlebodes_ | 10 | 38-45(40.1) | 8-13(10.3) | 2-5(3.9) + | | | | +_H. robertmertensi_ | 6 | 23-43(32.8) | 7-12(10.5) | 2-3(2.7) + | | | | +_H. sartori_ | 6 | 27-43(38.2) | 9-10(9.3) | 3-4(3.7) +------------------------+----+-------------+--------------+---------- + + + [Illustration: PLATE 13 + Upper figure, _Hyla microcephala microcephala_ (KU 64593); + middle figure, _H. microcephala underwoodi_ (KU 64565); + lower figure, _H. microcephala underwoodi_ (UMMZ 115247). + All approximately ×3.] + + + [Illustration: PLATE 14 + Upper figure, _Hyla robertmertensi_ (UMMZ 115243); + middle figure, _H. phlebodes_ (KU 64798); lower figure, + _H. sartori_ (UMMZ 119225). All approximately ×3.] + + + [Illustration: PLATE 15 + Tadpoles of _Hyla microcephala_ group: upper figure, _H. m. + microcephala_ (KU 104097); lower figure, _H. phlebodes_ + (KU 104099). Both ×4.] + + + [Illustration: PLATE 16 + Audiospectrograms and sections of mating calls of _Hyla + microcephala_ group: + (a) _H. m. microcephala_ (KU Tape No. 19); + (b) _H. robertmertensi_ (KU Tape No. 41); + (c) _H. phlebodes_ (KU Tape No. 6); + (d) _H. sartori_ (KU Tape No. 190).] + + +Table 4.--Comparative Cranial Osteology of Hyla microcephala Group + +===============+=======================+========================+ + Character | _H. microcephala_ | _H. robertmertensi_ | +---------------+-----------------------+------------------------+ +Frontoparietal | Minimally ossified | Ossification extensive | + | with large fontanelle | anteriorly with narrow | + | extending from | medial separation; | + | sphenethmoid to | fontanelle largest in | + | occipital ridge. | parietal region. | + | | | + | | | +Nasals | Moderately long and | Moderate in size; | + | slender; arcuate in | slightly wider | + | dorsal view. | anteriorly than | + | | posteriorly in dorsal | + | | view. | + | | | +Sphenethmoid | Extremely short in | Moderately short in | + | dorsal view. | dorsal view. | + | | | + | | | + | | | +Columella | Distal and greatly | Distal and slightly | + | expanded. | expanded or not. | +---------------+-----------------------+------------------------+ +Table 4. (Continued) +===============+========================+======================== + Character | _H. phlebodes_ | _H. sartori_ +---------------+------------------------+------------------------ +Frontoparietal | Ossification extensive | Ossification moderately + | anteriorly with narrow | extensive anteriorly; + | medial separation; | medial separation of + | fontanelle largest in | about uniform width + | parietal region. | throughout length of + | | fontanelle. + | | +Nasals | Moderate in size; | Long and broad; + | slightly wider | arcuate in dorsal + | anteriorly than | view. + | posteriorly in dorsal | + | view. | + | | +Sphenethmoid | Moderately short in | Moderately short in + | dorsal view. | dorsal view; ossified + | | anteriorly between + | | nasals. + | | +Columella | Distal and not | Distal and not + | expanded. | expanded. +---------------+------------------------+------------------------ + + + [Illustration: Fig. 5. Dorsal views of the skulls of (a) _Hyla m. + microcephala_ (KU 68293) and (b) _H. sartori_ (UMMZ S-2677). + Both × 12.] + + + [Illustration: Fig. 6. Dorsal views of skulls of (a) _Hyla phlebodes_ + (KU 68303) and (b) _H. robertmertensi_ (KU 59917). Both × 12.] + + +Despite the great reduction in the ossification of the cranial +elements, certain apparently consistent differences exist between the +species seem to be consistent. The most notable differences are: +1) amount of ossification of the frontoparietals and consequent shape +and size of the frontoparietal fontanelle, 2) shape of the nasals, +3) shape and extent of the sphenethmoid, and 4) shape of the columella +(Table 4, Figs. 5-6). On the basis of these characters, _Hyla +microcephala_ can be set apart from the other species and characterized +as having a poorly ossified frontoparietal and correspondingly large +frontoparietal fontanelle; long, slender, arcuate nasals; extremely +short sphenethmoid; and expanded distal end of the columella. The other +species in the group (_phlebodes_, _robertmertensi_, and _sartori_) +have more ossification of the frontoparietals, broader nasals, only a +moderately short sphenethmoid, and an unexpanded distal end of the +columella. Among these three species, the skulls of _phlebodes_ and +_robertmertensi_ are most nearly alike, whereas the skull of _sartori_ +differs by having a differently shaped frontoparietal fontanelle, +broader nasals, and an ossified anterior extension of the sphenethmoid +between the nasals (compare Fig. 5b with Fig. 6 a-b). + +Although all skulls examined belong to breeding adults, the extent of +the ossification of the frontoparietals and the resulting shape of the +frontoparietal fontanelle might be correlated with the age of the frog. +Nevertheless, in the 24 skulls of _Hyla microcephala_ examined, the +frontoparietals are less extensively ossified than in the skulls of the +other species. The trivial differences among the other three species +certainly are suggestive of close relationship, but on the basis of +present knowledge of the evolutionary trends in hylid cranial +osteology, the differences offer little evidence for determining +phylogenetic lineage. + + + + +ANALYSIS OF MATING CALLS + + +Calls of all five taxa were compared in several characteristics, of +which three are deemed most significant systematically. These are +1) the pattern and duration of the notes of a call-group, 2) the +fundamental frequency, and 3) the dominant frequency. Air temperatures +were noted at the time the calls were recorded, but no valid +correlation could be determined between this factor and any of the +parameters of the calls; consequently recordings made at all +temperatures (21-29° C.) were grouped together. + +_Pattern and duration of notes._--In all five taxa the basic pattern +consists of a call-group made up of one primary note followed by a +series of shorter secondary notes. In some species the secondary notes +differ from the primary in other characteristics. Both subspecies of +_Hyla microcephala_ have a long, unpaired primary note followed by 0 +to 18 (usually about 4) somewhat shorter paired secondary notes. In +calls of _Hyla m. microcephala_ the mean duration of the primary is +0.131 (0.10-0.16) second and that of the secondaries is 0.101 +(0.05-0.14) second, whereas in _H. m. underwoodi_ the mean duration of +the primary is 0.018 (0.05-0.15) second and that of the secondaries is +0.086 (0.06-0.11) second. + +_Hyla robertmertensi_ has a reverse of this pattern in that the primary +note is paired and the secondaries are unpaired. In the sample studied +a call-group contains 0-28 secondary notes (generally about 3). The +mean duration of the primary is 0.091 (0.07-0.11) second and that of +the secondaries is 0.040 (0.025-0.06) second. + +_Hyla phlebodes_ and _sartori_ have call-groups composed of a rather +short, unpaired primary and several short, unpaired secondaries +(0-28 in _phlebodes_, 0-23 in _sartori_). The mean duration of the +primary of _phlebodes_ is 0.105 (0.07-0.16) second and that of the +secondaries is 0.067 (0.035-0.12) second. The mean duration of the +primary of _sartori_ is 0.080 (0.07-0.09) second and that of the +secondaries is 0.053 (0.035-0.07) second. + +The two subspecies of _H. microcephala_ are identical in call pattern +and agree closely in duration of notes, although those of the nominate +subspecies tend to be slightly longer. _Hyla robertmertensi_ is +distinctive in call pattern in that it is the only species having a +paired primary; the duration of the primary is completely overlapped by +that in the other species, but the secondaries tend to be the shortest +in the group. The call patterns of _H. phlebodes_ and _H. sartori_ are +identical and the range of duration of notes of _phlebodes_ completely +overlaps that of _sartori_, although both the primary and secondary +notes of the latter tend to be somewhat shorter (Table 5, Pl. 16). + +_Fundamental frequency._--This parameter was analyzed for the primary +notes. It was measured for the secondaries as well and was found to +differ in magnitude in the same way as the primary note. In a few +examples of both subspecies of _H. microcephala_ a high primary note, +in which the fundamental frequency is exceptionally high, is sometimes +emitted (Fouquette, 1960b). None of these notes was used in this +analysis; only the fundamental frequencies of normal primary notes are +compared (Table 5, Fig. 7). + + +Table 5.--Comparison of Normal Mating Calls in the Hyla microcephala + Group. (Observed Range Given in Parentheses Below Mean; + Unless Otherwise Noted Data Are for Primary Notes.). + +----------------+--+---------+---------+-------------------+-------------- + | |Dominant | Funda- |Duration of notes | Repetition + | | | mental| (seconds) | rate of + Species |N |frequency|frequency+---------+---------+ secondaries + | | (cps) | (cps) | Primary |Secondary|(notes/minute) +----------------+--+---------+---------+---------+---------+-------------- +_H. m. |44| 5637 | 205 | 0.13 | 0.10 | 268 + microcephala_ | |(5150 |(184-244)|(0.11 |(0.05 | (192-353) + | | -5962)| | -0.16)| -0.14)| + | | | | | | +_H. m. |47| 5772 | 220 | 0.11 | 0.09 | 283 + underwoodi_ | |(5177 |(192-275)|(0.05 |(0.06 | (197-384) + | | -6200)| | -0.15)| -0.11)| + | | | | | | +_H. |25| 5388 | 162 | 0.09 | 0.04 | 418 + robertmertensi_| |(5150 |(140-178)|(0.07 |(0.03 | (368-570) + | | -5785)| | -0.11)| -0.06)| + | | | | | | +_H. phlebodes_ |34| 3578 | 148 | 0.11 | 0.07 | 284 + | |(3220 |(125-158)|(0.07 |(0.04 | (210-350) + | | -4067)| | -0.16)| -0.12)| + | | | | | | +_H. sartori_ |10| 3217 | 126 | 0.08 | 0.05 | 434 + | |(2950 |(116-135)|(0.07 |(0.04 | (396-477) + | | -3600)| | -0.09)| -0.07)| +----------------+--+---------+---------+---------+---------+-------------- + + +The two subspecies of _H. microcephala_ agree closely in fundamental +frequency. There is considerable overlap, but the difference +between the means is significant at the 0.001 level of probability +(t = 4.2406). The call of _H. robertmertensi_ does not overlap that +of _H. sartori_ or either subspecies of _H. microcephala_ in this +parameter; but it does overlap that of _H. phlebodes_, although again +the difference between the means is significant at the 0.001 level +(t = 9.360). _Hyla phlebodes_ and _sartori_ have the lowest fundamental +frequencies, and there is some overlap, but here too the difference +between the means is significant at the 0.001 level (t = 4.923). + +_Dominant frequency._--A dominant band of frequencies cuts across +the harmonics of the fundamental, obscuring the harmonic pattern and +generally shifting upward in frequency. The midpoint of this band is +measured at the terminal border as the dominant frequency. As with the +fundamental frequency, only the normal primary notes were utilized in +the comparisons (Table 5, Fig 8). + + + [Illustration: Fig. 7. Variation in the fundamental frequency of the + normal primary notes in the _Hyla microcephala_ group. The + horizontal lines = range of variation, vertical lines = mean, + solid bars = twice the standard error of the mean, and open + bars = one standard deviation. The number of specimens in each + sample is indicated in parentheses after the name of the taxon.] + + +The two subspecies of _H. microcephala_ agree more closely in this +parameter than in fundamental frequency. The overlap is great, but the +difference between the means is significant at the 0.001 level +(t = 3.658). The calls of both subspecies completely overlap that of +_robertmertensi_ in this parameter, but the difference between the +means is significant at the 0.001 level. The calls of _H. phlebodes_ +and _H. sartori_ overlap considerably in this characteristic, although +the difference between the means is significant at the 0.001 level +(t = 7.504) (Fig. 9). The call of neither species overlaps those of +_H. microcephala_ and _robertmertensi_. + + + [Illustration: Fig. 8. Variation in the mid-point of the dominant + frequency band of the normal primary notes in the _Hyla + microcephala_ group. The horizontal lines = range of variation, + vertical lines = mean, solid bars = twice the standard error of + the mean, and open bars = one standard deviation. The number of + specimens in each sample is indicated in parentheses after the + name of the taxon.] + + + [Illustration: Fig. 9. Scatter diagram relating the dominant and + fundamental frequencies of the normal primary notes in the + _Hyla microcephala_ group. Each symbol represents a different + individual.] + + +_Repetition rate._--The repetition rate of the secondary notes, in +calls consisting of more than one secondary, was measured for each +form. A considerable amount of variation in this parameter was found +in all of the taxa (Table 5). This variation probably is due in part +to the effect of temperature differences. Repetition rate is the only +parameter analyzed for which there is a correlation with the +air-temperature, but even here the correlation is weak, probably due +to the microenvironmental effects of humidity, air-movement, and other +factors in addition to the ambient air temperature that influences the +body temperature of the frogs. These rates are nearly alike in both +subspecies of _H. microcephala_ and in _phlebodes_. The repetition +rates in _H. robertmertensi_ and _H. sartori_ are considerably faster +than in the other three taxa. _Hyla sartori_ has the fastest +repetition rate of the group. + +In all characteristics of the mating calls the two subspecies of +_H. microcephala_ agree closely, as might be expected, although the +differences are statistically significant. _Hyla robertmertensi_ is +distinctive in call pattern and seems to be closer to _microcephala_ +in dominant frequency but closer to _H. phlebodes_ in fundamental +frequency. Thus, it is somewhat intermediate between _microcephala_ +and _phlebodes_. The identical pattern and similarity in fundamental +and dominant frequencies of the calls of _H. phlebodes_ and _H. sartori_ +possibly indicate close relationship. + +_Geographic variation in call._--_Hyla m. microcephala_ has higher +fundamental and dominant frequencies in Costa Rica than in Panamá. In +Costa Rican _H. m. underwoodi_ the fundamental and dominant frequencies +are lower than in other parts of the range. Frogs of this subspecies +recorded in Nicaragua and Honduras have slightly lower dominant +frequencies and higher fundamental frequencies than those recorded in +Guatemala or Oaxaca. The duration of both primary and secondary notes +decreases to the south; samples from Nicaragua and Costa Rica have the +shortest notes. Comparison of duration of notes in the two subspecies +shows that the Panamanian _H. m. microcephala_ have slightly longer +notes than do any _H. m. underwoodi_; the more northern populations of +_H. m. underwoodi_ from México most closely approach _H. m. +microcephala_ in this characteristic. + +The calls of _H. robertmertensi_ in Oaxaca have higher dominant and +fundamental frequencies and longer secondary notes than do those in +Chiapas. + +The calls of _H. phlebodes_ recorded at Puerto Viejo, Costa Rica, +have slightly lower dominant frequencies than do those recorded at +Turrialba, Costa Rica, and in Panamá, whereas those recorded at +Turrialba have lower fundamental frequencies than in other samples. +The duration of notes is slightly shorter in both Costa Rican samples +than in those recorded in Panamá. + + + + +LIFE HISTORY + + +The frogs of the _Hyla microcephala_ group breed in shallow grassy +ponds. In some places they breed in permanent ponds, but usually +congregate around temporary pools, such as depressions in forests, +flooded fields, and roadside ditches. At the height of their breeding +season, usually in the early part of the rainy season, the +congregations are made up of large numbers of individuals. In April, +1961, and in June, 1966, the senior author noted nearly continuous +choruses of _H. m. microcephala_ in roadside ditches along the 75 +kilometers of road between Villa Neily and Palmar Sur, Puntarenas +Province, Cost Rica; on June 20, 1966, at Puerto Viejo, Heredia +Province, Costa Rica, he estimated approximately 900 _Hyla phlebodes_ +in one pond, and two nights later noticed that the number of +individuals had increased substantially. Other observations by the +first author on size of breeding congregations include nearly +continuous choruses of _H. m. underwoodi_ between Villahermosa and +Teapa, Tabasco, in July of 1958, an estimated 400 _Hyla robertmertensi_ +in a road side ditch 7.2 kilometers west-northwest of Zanatepec, +Oaxaca, on July 13, 1956, and approximately 150 _Hyla sartori_ around +a rocky pool in a riverbed, 11.8 kilometers west-northwest of Tierra +Colorada, Guerrero, on June 28, 1958. + +The length of the breeding season seemingly is more dependent on +climatic conditions in various parts of Middle America than on +behavioral differences in the various species. Thus, Fouquette (1960b) +found in the Canal Zone that _H. m. microcephala_ formed breeding +choruses from May through January, the entire rainy season in that +area. In the wetter coastal region of Puntarenas Province, Costa Rica, +the species breeds as early as mid-March, whereas in the drier region +encompassing Guanacaste Province, Costa Rica, and southwestern +Nicaragua breeding activity is initiated by the first heavy rains of +the season, usually in June. + +_Hyla phlebodes_ inhabits regions having rainfall throughout the year. +Although large breeding congregations are most common in the early +parts of the rainy season, males probably call throughout the year. At +Puerto Viejo in Costa Rica the senior author has heard _Hyla phlebodes_ +in February, April, June, July, and August. Charles W. Myers noted +calling males of this species in the area around Almirante, Bocas del +Toro Province, Panamá, in September, October, and February. An +exception to the correlation between rainfall and breeding activity +was noted by the junior author in _Hyla phlebodes_ in the Canal Zone, +where he noticed a decrease in activity of that species in October and +November, when the rains are heaviest and most frequent. Furthermore, +independent observations made by both of us indicate that _H. +phlebodes_ does not reach peaks of activity during or immediately +after heavy rains, but instead builds up to peaks of activity two or +three days after a heavy rain. This is in contrast to the other +species, all of which characteristically inhabit drier environments +than does _H. phlebodes_. Peaks of breeding activity in the other +species occur immediately after, or even during, heavy rains. + +The calling location of the males generally is on vegetation above, +or at the edge of, the water. _Hyla microcephala_ and _H. phlebodes_ +call almost exclusively from grasses and sedges; _phlebodes_ usually +calls from taller and more dense grasses than does _microcephala_. +Except for some minor differences in calling location observed by +the junior author (Fouquette, 1960b) in the Canal Zone, the differences +in density and height of grasses utilized for calling-locations +probably is dependent primarily on the nature of the available +vegetation. Although bushes and broad-leafed herbs are usually present +at the breeding sites, males of these species seldom utilize them for +calling locations. Both _H. robertmertensi_ and _H. sartori_ have been +observed calling from grasses, herbs, bushes, and low trees. Calling +males of _robertmertensi_ have been found two meters above the ground +in small trees. + +Daytime retreats in the breeding season sometimes are no more than +shaded clumps of vegetation adjacent to a pond or in clumps of grass +in a pond. Individuals of _H. m. underwoodi_ were found by day under +the outer sheaths of banana plants next to a water-filled ditch. Dry +season refuges are unknown. + +Amplexus is axillary in all four species. Egg deposition has been +observed in _H. m. microcephala_, _m. underwoodi_, and _phlebodes_. +In all three the eggs are deposited in small masses that float near +the surface of the water and usually are at least partly attached to +emergent vegetation. Each clutch does not represent the entire egg +complement of the female. + +Tadpoles are definitely known of only _H. m. microcephala_ and +_phlebodes_; these have been described in the preceding accounts of +the species. The tadpoles of these two species can be distinguished +readily (Pl. 15). The tadpole of _H. microcephala_ has a uniformly +white venter and nearly transparent tail, whereas in _H. phlebodes_ +the venter is flecked anteriorly and the tail is mottled. In life, _H. +microcephala_ is easily recognized by the orange posterior half of +the tail, whereas the tail in _H. phlebodes_ is mottled tan and grayish +brown. + + + + +PHYLOGENETIC RELATIONSHIPS + + +The evidence already presented on osteology, external structure, +coloration, mating call, and life history emphatically show that the +four species under consideration are a closely related assemblage. +Now the question arises: To what other groups in the genus is the +_Hyla microcephala_ group related? Furthermore, it is pertinent to +this discussion to attempt a reconstruction of the phylogeny of the +group as a whole and of the individual species in the _Hyla +microcephala_ group. With regard to the relationships of the group we +must take into account certain species in South America. Our endeavors +there are hampered by the absence of data on the mating calls and life +histories of most of the relevant species. + +As mentioned in the account of _Hyla m. microcephala_, the species +_microcephala_ possibly is subspecifically related to _Hyla misera_, a +frog widespread in the Amazon Basin. _Hyla misera_ resembles +_microcephala_ in coloration, external structure, and cranial +characters. The frontoparietals are equally poorly ossified, and the +frontoparietal fontanelle is extensive. Our principal reason for not +considering the two taxa conspecific at this time is our lack of +knowledge concerning the color of living _H. misera_, the structure of +the tadpoles, and the characteristics of the mating call. Even with the +absence of such data that we think essential to establish the +nomenclature status of the taxa, we are confident that the two are +sufficiently closely related that any discussion of the phylogenetic +relationships of one species certainly must involve consideration of +the other. + +_Hyla misera_ possibly is allied to other small yellowish tan South +American _Hyla_ that lack dark pigmentation on the thighs. Probable +relatives are _Hyla elongata_, _minuta_ (with _goughi_, _pallens_, +_suturata_, _velata_, and possibly others as synonyms), _nana_, and +_werneri_. The consideration of the interspecific relationships of +these taxa is beyond the scope of this paper, but we can say that each +of these species has a pale yellowish tan dorsum, relatively broad +dorsolateral brown stripe, and narrow longitudinal brown lines or +irregular marks on the dorsum. Furthermore, examination of the skulls +of _elongata_, _nana_, and _werneri_ reveals that they are like +_misera_ and _microcephala_ in the nature of the frontoparietal +fontanelle and in having a greatly reduced quadratojugal. Thus, on the +basis of cranial and external characters the _Hyla microcephala_ group +can be associated with _Hyla misera_ and its apparent allies in South +America. This association can be only tentative until the mating calls, +tadpoles, and chromosome numbers of the South American species are +known. + +Among the Middle American hylids, only the _Hyla microcephala_ group +and _H. ebraccata_ have a haploid number of 15 chromosomes (Duellman +and Cole, 1965). All other New World _Hyla_, for which the number is +known, have a haploid number of 12; the only other _Hyla_ having 15 +is a Papuan _Hyla angiana_ (Duellman, 1967). + +_Hyla ebraccata_ occurs in the humid tropical lowlands of Middle +America and the Pacific lowlands of northwestern South America. It is +the northernmost, and only Central American, representative of the +_Hyla leucophyllata_ group, which is diverse (about 10 species +currently recognized) and widespread in tropical South America east of +the Andes. This group is characterized by having broad, flat skulls +with larger nasals and more ossification of the frontoparietals than +in the _Hyla microcephala_ group. The quadratojugal is present as a +small anteriorly projecting spur that does not connect with the +maxillary. Externally, the _Hyla leucophyllata_ group is characterized +by having a well-developed axillary membrane, uniformly yellow thighs, +and a dorsal color pattern in many species consisting of a dark lateral +band, a pale dorsolateral band or dorsal ground color, and a large +middorsal dark mark. In some species, the dorsal pattern consists of +small dark markings or is nearly uniformly pale. At least in the +Central American _Hyla ebraccata_, the mating call consists of a +single primary note followed by a series of shorter secondary notes, +the tadpoles have xiphicercal tails and lack teeth, and the haploid +number of chromosomes is 15. On the strength of these observations it +seems imperative to consider the _Hyla leucophyllata_ group as a close +ally to the _Hyla microcephala_ group. Successful artificial +hybridization supports the close relationship of _H. m. microcephala_ +and _phlebodes_; partial success of artificial hybridization of these +two with _ebraccata_ (Fouquette, 1960b) provides further evidence for +close relationship between the _Hyla leucophyllata_ and _Hyla +microcephala_ groups. + +In México and northern Central America two small species, _Hyla picta_ +and _Hyla smithi_, comprise the _Hyla picta_ group. These frogs +resemble members of the _Hyla microcephala_ group by having a +yellowish tan dorsum with a dorsolateral white stripe and uniformly +yellow thighs. Furthermore the mating call is not unlike those of +the species in the _Hyla microcephala_ group. Despite these +similarities, the _Hyla picta_ group differs from the _Hyla +microcephala_ group by having a well-developed quadratojugal that +connects to the maxillary, tadpoles with teeth present and caudal fins +completely enclosing the caudal musculature, and a haploid number of +12 chromosomes. In all of these characteristics the frogs of the + +_Hyla picta_ group more closely resemble other Middle American _Hyla_ +than they do the _Hyla microcephala_ group. Therefore, it can best be +presumed that the superficial resemblances of coloration and the +mating call are the result of convergence. + +Since the _Hyla microcephala_ and _leucophyllata_ groups apparently +are related and since the greatest diversity of these frogs is in +South America (if _Hyla misera_ and its relatives are placed with the +_Hyla microcephala_ group), it seems appropriate to place the centers +of origins of these groups in South America. Therefore, the _Hyla +microcephala_ group and _Hyla ebraccata_ of the _Hyla leucophyllata_ +group either have immigrated into Central America, or they are +representatives of those groups that were isolated in Central America +during most of the Cenozoic when South America was separated from +Central America. + +The interspecific relationships of the species in the _Hyla +microcephala_ group are not clear. On the basis of coloration, _H. m. +microcephala_ and _H. robertmertensi_ are close, and _H. m. underwoodi_ +and _H. phlebodes_ are nearly identical. The mating calls of _H. +phlebodes_ and _sartori_ closely resemble one another, whereas the call +of _robertmertensi_ is intermediate between these and _microcephala_. + +In most respects _Hyla microcephala_ is distinct from the other +species, and with the exception of the amount of ossification of the +frontoparietals, the other species can be easily derived from a +_microcephala_-like ancestor. Possibly the slightly increased +ossification of the frontoparietals in _robertmertensi_, _phlebodes_, +and _sartori_ is secondary, or possibly after differentiation of the +species the amount of ossification was further reduced in +_microcephala_. If so, the species fall into a reasonable phylogenetic +scheme that has _microcephala_ as the extant species most like the +ancestral stock. + +We visualize the evolutionary history of the group to have followed +a course that began with the invasion of Central America by a +_microcephala_ ancestral stock that differentiated into two populations +in lower Central America--a _microcephala_-like frog on the Pacific +lowlands and a _phlebodes_-like frog on the Caribbean lowlands. +Differentiation could have been brought about by isolation by montaine +or marine barriers. The population on the Pacific lowlands either was +preadapted for subhumid conditions or became so adapted and dispersed +northward onto the Pacific lowlands of northern Central America. +Simultaneously the frogs on the Caribbean lowlands, which were adapted +to humid environments, dispersed northward in the humid forested +regions to southern México and crossed the Isthmus of Tehuantepec onto +the Pacific slopes of Oaxaca and Guerrero northward to Jalisco. +Subsequent development of arid conditions, possibly in the Pliocene, +Pleistocene, or even as late as the Thermal Maximum in post-Wisconsin +time, resulted in a restriction of the ranges in northern Central +America, thereby isolating part of the _phlebodes_-stock on the +Pacific slopes of México, where it adapted to drier conditions and +evolved into _sartori_. The rest of the _phlebodes_-stock was +restricted to the humid forests on the Caribbean lowlands of lower +Central America. The increased aridity on the Pacific lowlands +eliminated the _microcephala_-stock from southern Honduras and +northwestern Nicaragua and in so doing left an isolated population on +the lowlands of Chiapasand Guatemala, which differentiated into +_robertmertensi_. The original stock on the Pacific lowlands of Panamá +and southeastern Costa Rica became _microcephala_. + +If the _microcephala_-stock was, as we believe, better adapted for +existence under subhumid conditions than was the _phlebodes_-stock, +the development of subhumid conditions in much of the lowland region +of northern Central America and southern México would have permitted +the expansion of the range of _microcephala_ into the area now +inhabited by _H. m. underwoodi_, while _phlebodes_ was being eliminated +from this area by climatic conditions that were unsuited to its +survival there. Perhaps the similarity in coloration of _H. m. +underwoodi_ and _phlebodes_ is the result of convergence or possibly +hybridization occurred at the time the former was expanding its range +and the latter's range was being restricted. If hybridization did +occur, the differences in mating call subsequently were enhanced, +thereby providing a valid isolating mechanism in sympatric populations. + +_Hyla microcephala_ and _phlebodes_ range into northern South America. +Probably both species entered South America in relatively recent times +after they had differentiated from one another in Central America. + + + + +LITERATURE CITED + + +Boulenger, G. A. + 1898. Fourth report on additions to the batrachian collection in the + Natural-History Museum. Proc. Zool. Soc. London, 1898, + pp. 373-482, pls. 38-39. October 1. + 1899. Descriptions of new batrachians in the collection of the + British Museum (Natural History). Ann. Mag. Nat. Hist, ser. + 7, 3:273-277, pls. 11-12. + +Breder, C. M. Jr. + 1946. Amphibians and reptiles of the Rio Chucunaque Drainage, Darien, + Panama, with notes on their life histories and habits. Bull. + Amer. Mus. Nat. Hist, 86:375-436, pls. 42-60, August 26. + +Cole, L. J. and Barbour, T. + 1906. Vertebrata from Yucatan: Reptilia; Amphibia; Pisces. Bull. Mus. + Comp. Zool., 50:146-159. November. + +Cope, E. D. + 1886. Thirteenth contribution to the herpetology of tropical America. + Proc. Amer. Philos. Soc, 23:271-287. February 11. + 1894. Third addition to a knowledge of the Batrachia and Reptilia of + Costa Rica. Proc. Acad. Nat. Sci. Philadelphia, 1894, pp. + 194-206. + +Duellman, W. E. + 1956. The frogs of the hylid genus _Phrynohyas_ Fitzinger, 1843. + Misc. Publ. Mus. Zool., Univ. Michigan, 96:1-47, pls. 1-6. + February 21. + 1967. Additional studies of chromosomes of anuran amphibians. Syst. + Zool., 16:38-43, March 17. + +Duellman, W. E. and Cole, C. J. + 1965. Studies of chromosomes of some anuran amphibians (Hylidae and + Centrolenidae). Syst. Zool., 14:139-143. July 9. + +Duellman, W. E. and Trueb, L. + 1966. Neotropical hylid frogs, genus Smilisca. Univ. Kansas Publ., + Mus. Nat. Hist., 17:281-375, pls. 1-12. July 14. + +Dunn, E. R. + 1931. The amphibians of Barro Colorado Island. Occas. Papers Boston + Soc. Nat. Hist., 5:403-421. October 10. + 1933. Amphibians and reptiles from El Valle de Anton, Panamá. + _Ibid._, 8:65-79. June 7. + 1934. Two new frogs from Darien. Amer. Mus. Novit., 747:1-2. + September 17. + +Fouquette, M. J. Jr. + 1960a. Call structure in frogs of the family Leptodactylidae. Texas + Jour. Sci., 12:201-215. October. + 1960b. Isolating mechanisms in three sympatric tree frogs in the Canal + Zone. Evolution, 14:484-497. December 16. + +Gaige, H. T., Hartweg, N. and Stuart, L. C. + 1937. Notes on a collection of amphibians and reptiles from + eastern Nicaragua. Occas. Papers Mus. Zool., Univ. Michigan, + 357:1-18. October 26. + +Gosner, K. L. + 1960. A simplified table for staging anuran embryos and larvae with + notes on identification. Herpetologica, 16:183-190. + September 23. + +Kellogg, R. + 1932. Mexican tailless amphibians in the United States National + Museum. Bull. U.S. Natl. Mus., 160:1-224. March 31. + +Rivero, J. A. + 1961. Salientia of Venezuela. Bull. Mus. Comp. Zool., 126:1-207. + November. + +Smith, H. M. + 1951. The identity of _Hyla underwoodi_ Auctorum of Mexico. + Herpetologica, 7:184-190. December 31. + +Stejneger, L. + 1906. A new tree toad from Costa Rica. Proc. U. S. Natl. Mus., + 30:817-818. June 4. + +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 4. + +Taylor, E. H. + 1952. The frogs and toads of Costa Rica. Univ. Kansas Sci. Bull., + 35-577-942. July 1. + + +_Transmitted July 11, 1967._ + + + +Transcriber's Notes + +This file was derived from scanned images. With the exception of the +list of typographical errors that were corrected below, the original +text is presented. + +In the copy of the original, the Plate text contains the notation +"X 2" after the caption to let the reader know that the image was +enlarged by a factor of two. + + +Emphasis Notation + + _Text_ = Italic + + +Text+ = Bold + + +Typographical Errors Corrected: + +Several minor typographical corrections were made (missing periods, +commas, incomplete italicization, etc.); but are not indicated here. +More substantial changes are listed below: + +Page 533 - UMZ => UMMZ +Page 534 - Diganosis => Diagnosis +Page 544 - fontanells => fontanelle +Page 545 - prrimary => primary +Page 547 - band of of frequencies => band of frequencies +Page 550 - ad => had +Page 551 - clumbs => clumps +Page 552 - acount => account +Page 557 - Minchigan => Michigan + + + + + + + +End of the Project Gutenberg EBook of Middle American Frogs of the Hyla +microcephala Group, by William E. 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Fouquette, Jr. +</title> +<style type="text/css"> + +body {margin-left: 10%; margin-right: 10%;} + + h1,h2,h3,h4,h5,h6 {text-align: center; clear: both;} + +p { text-align: justify; text-indent: 2em;} + + hr {color: #000;} + .hr75 {width:75%;} + + table {margin-left: auto; margin-right: auto; vertical-align:text-top;} + .data {text-align:center; white-space: nowrap; font-size:.8em; border-collapse: collapse;} + +.pagenum { /* uncomment the next line for invisible page numbers */ + /* visibility: hidden; */ + position: absolute; + left: 92%; + font-size: 0.75em; + text-align: right; + color: #a0a0a0; +} /* page numbers */ + + .remarks {font-size:1.25em;} + + .bb {border-bottom: solid #000 1px;} + .bl {border-left: solid #000 1px;} + .bt {border-top: solid #000 1px;} + .br {border-right: solid #000 1px;} + .bbox {border: solid #000 1px;} + + .toc td {background: url('images/dot.png') repeat-x 0;} + .toc {text-align:left; width:100%; border:1px; background:#fff; margin-bottom:6px;} + .toc .blank {background:#ffffff;} + .vtop {vertical-align: top;} + .descrp2 {text-align: right; float:right;} + .species p {margin-left: 4em; text-indent: -2em;} + .references p {padding-left: 5em; text-indent:-3em; margin-right: 10%;} + .center {text-align: center;} + .text_lf {text-align: left;} + .text_rt {text-align: right;} + .smcap {font-variant: small-caps;} + .caption1 {font-weight: bold; font-size:2.00em; text-align: center;} + .caption2 {font-weight: bold; font-size:1.50em; text-align: center;} + .caption3 {font-weight: bold; font-size:1.15em; text-align: center;} + .caption3nb {font-size:1.15em; text-align: center;} + .caption4 {font-weight: bold; font-size:0.75em; text-align: center;} + .trans_notes {background:#d0d0d0; padding: 7px; border:solid black 1px;} + +/* Footnotes */ +.footnote {margin-left: 10%; margin-right: 10%; font-size: 0.75em;} + +.footnote .label {position: absolute; right: 84%; text-align: right;} + +.fnanchor { + vertical-align: super; + font-size: .8em; + text-decoration: + none; +} + +</style> + </head> +<body> + + +<pre> + +The Project Gutenberg EBook of Middle American Frogs of the Hyla +microcephala Group, by William E. Duellman and M. J. Fouquette + +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: Middle American Frogs of the Hyla microcephala Group + +Author: William E. Duellman + M. J. Fouquette + +Release Date: December 9, 2010 [EBook #34604] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK MIDDLE AMERICAN FROGS *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +https://www.pgdp.net + + + + + + +</pre> + + +<div class="trans_notes"> +<div class="caption2">Transcriber's Notes</div> + + +<p>Typographical Corrections</p> + +<a name="typos"></a> +<table summary="Typos"> +<tr><td>Page 533 -</td><td>UMZ</td><td>=></td><td><a href="#UMMZ">UMMZ</a></td></tr> +<tr><td>Page 534 -</td><td>Diganosis</td><td>=></td><td><a href="#Diagnosis">Diagnosis</a></td></tr> +<tr><td>Page 544 -</td><td>fontanells</td><td>=></td><td><a href="#fontanelle">fontanelle</a></td></tr> +<tr><td>Page 545 -</td><td>prrimary</td><td>=></td><td><a href="#primary">primary</a></td></tr> +<tr><td>Page 547 -</td><td>band of of frequencies</td><td>=></td><td><a href="#of_of">band of frequencies</a></td></tr> +<tr><td>Page 550 -</td><td>ad</td><td>=></td><td><a href="#had">had</a></td></tr> +<tr><td>Page 551 -</td><td>clumbs</td><td>=></td><td><a href="#clumps">clumps</a></td></tr> +<tr><td>Page 552 -</td><td>acount</td><td>=></td><td><a href="#account">account</a></td></tr> +<tr><td>Page 557 -</td><td>Minchigan</td><td>=></td><td><a href="#Michigan">Michigan</a></td></tr> +</table> +<p> </p> +</div> +<p> </p> +<p> </p> + + + +<p><span class="pagenum"><a name="Page_517" id="Page_517">[Pg 517]</a></span> + +<p> </p> +<img src="images/bar_double.png" width="100%" height="15" border="0" alt="double bar"> +<div class="caption2"><div class="smcap">University of Kansas Publications<br> +Museum of Natural History</div></div> +<hr class="hr75"><br> +<div class="caption2">Volume 17, No. 12, pp. 517-557, pls. 13-16, 9 figs.</div><br> +<div class="center"><img src="images/bar_single.png" width="28%" height="15" title="bar" alt="bar"> <span class="caption2">March 20, 1968</span> <img src="images/bar_single.png" width="28%" height="15" title="bar" alt="bar"></div> +<p> </p> +<p> </p> +<p> </p> +<div class="caption1"> +Middle American Frogs +of the Hyla microcephala Group<br> +</div> +<p> </p> +<p> </p> + +<div class="caption3"> +BY<br> +<p> </p> +WILLIAM E. DUELLMAN AND M. J. FOUQUETTE, JR.<br> +</div> +<p> </p> +<p> </p> +<p> </p> +<p> </p> +<p> </p> +<p> </p> + +<div class="caption2"> +<span class="smcap">University of Kansas</span><br> +<span class="smcap">Lawrence</span><br> +1968 +</div> +<p> </p> +<p> </p> + +<span class="pagenum"><a name="Page_518" id="Page_518">[Pg 518]</a></span> + +<div class="center"> +<div class="caption3"> +<span class="smcap">University of Kansas Publications, Museum of Natural History</span><br> +<br> +Editors: E. Raymond Hall, Chairman, Henry S. Fitch,<br> +Frank B. Cross<br> +</div> +<p> </p> +<p> </p> + +Volume 17, No. 12, pp. 517-557, 4 pls. 9 figs.<br> +<br> +Published March 20, 1968<br> +<p> </p> +<p> </p> + +<span class="smcap">University of Kansas</span><br> +Lawrence, Kansas<br> +<p> </p> +<p> </p> +<div class="caption4"> +PRINTED BY<br> +ROBERT R. (BOB) SANDERS, STATE PRINTER<br> +TOPEKA, KANSAS<br> +1968<br> +<img src="images/union_label.png" width="71" height="26" border="0" alt="Look for the Union Label" title="Look for the Union Label"><br> +31-9419<br> +</div> +</div> +<p> </p> +<p> </p> + +<span class="pagenum"><a name="Page_519" id="Page_519">[Pg 519]</a></span> +<p> </p> +<p> </p> + +<div class="caption2">Middle American Frogs<br> +of the Hyla microcephala Group</div> +<p> </p> + +<div class="caption3">BY<br> +WILLIAM E. DUELLMAN AND M. J. FOUQUETTE, JR.</div> +<p> </p> +<p> </p> + +<a name="TOC"></a><a name="CONTENTS" id="CONTENTS"></a> +<div class="caption2">CONTENTS</div> +<br> +<table class="toc" summary="Table of Contents"> +<tr><td class="blank"> </td><td class="blank text_rt"> PAGE</td></tr> +<tr><td><span class="blank smcap"><a href="#INTRODUCTION">Introduction</a> </span></td><td class="blank text_rt">519</td></tr> +<tr><td><span class="blank"> <a href="#ACKNOWLEDGMENTS">Acknowledgments</a> </span></td><td class="blank text_rt">520</td></tr> +<tr><td><span class="blank"> <a href="#MATERIALS_AND_METHODS">Materials and Methods</a> </span></td><td class="blank text_rt">520</td></tr> +<tr><td><span class="blank smcap"><a href="#HYLA_MICROCEPHALA_GROUP">Hyla microcephala Group</a> </span></td><td class="blank text_rt">521</td></tr> +<tr><td><span class="blank"> <a href="#KEY_TO_SPECIES">Key to Species and Subspecies</a> </span></td><td class="blank text_rt">522</td></tr> +<tr><td><span class="blank smcap"><a href="#ACCOUNTS">Accounts of Species and Subspecies</a> </span></td><td class="blank text_rt">523</td></tr> +<tr><td><span class="blank smcap"><a href="#CRANIAL_OSTEOLOGY">Cranial Osteology</a> </span></td><td class="blank text_rt">540</td></tr> +<tr><td><span class="blank smcap"><a href="#ANALYSIS_OF_MATING_CALLS">Analysis of Mating Calls</a> </span></td><td class="blank text_rt">544</td></tr> +<tr><td><span class="blank smcap"><a href="#LIFE_HISTORY">Life History</a> </span></td><td class="blank text_rt">550</td></tr> +<tr><td><span class="blank smcap"><a href="#PHYLOGENETIC_RELATIONSHIPS">Phylogenetic Relationships</a> </span></td><td class="blank text_rt">552</td></tr> +<tr><td><span class="blank smcap"><a href="#LITERATURE_CITED">Literature Cited</a> </span></td><td class="blank text_rt">556</td></tr> +</table> +<p> </p> +<p> </p> + +<a name="INTRODUCTION" id="INTRODUCTION"></a> +<div class="caption2">INTRODUCTION</div> + +<p>The small yellow tree frogs, <i>Hyla microcephala</i> and its relatives, +are among the most frequently heard and commonly collected frogs in +the lowlands of southern México and Central America. The similarities +in size, proportions, and coloration of the different species have +resulted not so much in a multiplicity of specific names, but in +differences of opinion on the application of existing names to the +various taxa. For example, the populations on the Atlantic lowlands +have been known by three names, two of which have been applied to +other taxa. Much of the confusion has been the result of previous +workers' unfamiliarity with the animals in life and unawareness of the +intraspecific geographic variation in the most widespread species.</p> + +<p>Independently we undertook studies of these frogs in the field. The +second author worked on the interspecific relationships and isolating +mechanisms in Panamá (Fouquette, 1960b) and later studied the species +in southern México. As part of his survey of the hylids of Middle +America, the first author accumulated field and laboratory data on the +frogs throughout their ranges in México and Central America. The +purpose of this report is to present our findings on the four species +of Middle American frogs that we place in the <i>Hyla microcephala</i> +group. In addition to conventional taxonomic characters, we have +utilized the features of the cranial osteology and have relied heavily +on the data obtained from an analysis of the mating calls. +Furthermore, we have included ecological and distributional data in +our synthesis of interspecific relationships. +<p> </p> +<p> </p> + +<span class="pagenum"><a name="Page_520" id="Page_520">[Pg 520]</a></span> +<a name="ACKNOWLEDGMENTS" id="ACKNOWLEDGMENTS"></a> +<div class="caption2">ACKNOWLEDGMENTS</div> + +<p>Examination of specimens was made possible by the provision of working +space at various institutions or through the loan of specimens. For +their generosity in this manner we are grateful to Richard J. Baldauf, +Charles M. Bogert, James E. Böhlke, Doris M. Cochran, Robert F. Inger, +John M. Legler, Alan E. Leviton, Gerald Raun, Jay M. Savage, Hobart M. +Smith, Robert C. Stebbins, Wilmer W. Tanner, Charles F. Walker, Ernest +E. Williams, and Richard G. Zweifel.</p> + +<p>Duellman is especially grateful to Charles W. Myers, Linda Trueb, +Jerome B. Tulecke, and John Wellman for their assistance in the field +and to Linda Trueb for her work on the cranial osteology that is +incorporated in this report. Fouquette is indebted to H. Morgan Smith +and A. C. Collins for assistance in the field, to A. J. Delahoussaye +for assistance in the laboratory, and to W. Frank Blair for use of the +facilities of the sound laboratory at the University of Texas and for +much help in the early stages of this study.</p> + +<p>The research reported herein was accomplished mainly through support +by the National Science Foundation (grants NSF G-9827 and GB-1441 to +Duellman and GB-599 to Fouquette). The latter's field work in México +was assisted in part by NSF Grant G-4956 to W. Frank Blair. Some of +the field studies carried out in Panamá by Duellman were supported by +a grant from the National Institutes of Health (NIH GM-12020).</p> + +<p>We are grateful to many persons, too numerous to mention, who in +various ways aided our field work in Middle America. We are especially +indebted to Dr. Rodolfo Hernandez Corzo and the late Ing. Luis Macías +Arellano of the Dirección General de la Fauna Silvestre of the Mexican +government for providing permits to collect in México.</p> +<p> </p> + +<a name="MATERIALS_AND_METHODS" id="MATERIALS_AND_METHODS"></a> +<div class="caption3">Materials and Methods</div> + +<p>For this report, data has been obtained from 2829 preserved frogs, 42 +skeletal preparations, 8 lots of tadpoles and young, and 4 lots of +eggs. Much of the material was collected in our independent field +work, which has extended over a period of 11 years.</p> + +<p>Measurements were taken in the manner described by Duellman (1956). +Osteological data were obtained from specimens that were cleared in +potassium hydroxide, stained with alizarin red, and stored in +glycerine. Recordings were made by means of Magnemite portable tape +recorders (Amplifier Corp. America). The calls recorded by Fouquette +were analyzed on a Sonagraph (Kay Electric Co.) at the University of +Texas; those recorded by Duellman were analyzed mainly on a Vibralyzer +(Kay Electric Co.) at the University of Kansas and in part on a +Sonagraph at the University of Southwestern Louisiana. Sample calls +were analyzed on all three instruments; the slight differences in +results were found to be less than the error in measurement, so the +data from all sources were combined without correction. The techniques +and terminology of the calls are those defined by Fouquette (1960a, +1960b).</p> + +<p>In the accounts of the species we have attempted to give a complete +synonymy. At the end of each species account the localities from which +specimens were examined are listed alphabetically within each state, +province, or department, which in turn are listed alphabetically +within each country. The countries are arranged from north to south. +Localities preceded by an +<span class="pagenum"><a name="Page_521" id="Page_521">[Pg 521]</a></span> +asterisk (*) are not plotted on the accompanying maps due to the +crowding of symbols that would have resulted. Abbreviations for museum +specimens are listed below:</p> + +<table summary="Specimen Sources"> +<tr><td>AMNH</td><td>—American Museum of Natural History</td></tr> +<tr><td>ANSP</td><td>—Academy of Natural Sciences of Philadelphia</td></tr> +<tr><td>BMNH</td><td>—British Museum (Natural History)</td></tr> +<tr><td>BYU</td><td>—Brigham Young University</td></tr> +<tr><td>CAS</td><td>—California Academy of Sciences</td></tr> +<tr><td>FMNH</td><td>—Field Museum of Natural History</td></tr> +<tr><td>KU</td><td>—University of Kansas Museum of Natural History</td></tr> +<tr><td>MCZ</td><td>—Museum of Comparative Zoology</td></tr> +<tr><td>MVZ</td><td>—Museum of Vertebrate Zoology</td></tr> +<tr><td>SU</td><td>—Stanford University</td></tr> +<tr><td>UIMNH</td><td>—University of Illinois Museum of Natural History</td></tr> +<tr><td>UMMZ</td><td>—University of Michigan Museum of Zoology</td></tr> +<tr><td>USC</td><td>—University of Southern California</td></tr> +<tr><td>USNM</td><td>—United States National Museum</td></tr> +<tr><td>UU</td><td>—University of Utah</td></tr> +<tr><td>TCWC</td><td>—Texas Cooperative Wildlife Collection</td></tr> +<tr><td>TNHM</td><td>—Texas Natural History Museum</td></tr> +</table> +<p> </p> +<p> </p> + +<a name="HYLA_MICROCEPHALA_GROUP" id="HYLA_MICROCEPHALA_GROUP"></a> +<div class="caption2">HYLA MICROCEPHALA GROUP</div> + +<div class="blockquot"><p><i>Definition.</i>—Small hylids attaining a maximum snout-vent length of +27 mm. in males and 32 mm. in females; dorsum yellowish tan with brown +markings; thighs uniformly yellow, vocal sac in breeding males yellow; +snout truncate in lateral profile; tympanum distinct, usually slightly +smaller than one-half diameter of eye; vocal sac single, median, +subgular; fingers about one-third webbed; toes webbed nearly to bases +of discs, except only to middle of antepenultimate or base of +penultimate phalanx of fourth toe; tarsal fold weak; inner metatarsal +tubercle low, flat, elliptical; axillary membrane present; pupil +horizontally elliptical; palpebral membrane unmarked; cranial elements +reduced in ossification; sphenethmoid small, short; frontoparietal +fontanelle large; tegmen tympani not extensive; quadratojugal greatly +reduced; anterior arm of squamosal extending only about one-fourth +distance to maxillary; posterior arm of squamosal not having bony +connection with proötic; nasals lacking maxillary processes; medial +ramus of pterygoid not having bony attachment to proötic; maxillary, +premaxilary, and prevomerine teeth present; palatine and parasphenoid +teeth absent; Mentomeckelians ossified; tadpoles having xiphicercal +tails with deep caudal fins and terminal mouth lacking teeth; mating +call consisting of one primary note followed by a series of shorter +secondary notes; haploid number of chromosomes, 15 (known only in <i>H. +microcephala</i> and <i>H. phlebodes</i>.)</p> + +<p><i>Content.</i>—As recognized here the <i>Hyla microcephala</i> group contains +four species, one having two subspecies. An alphabetical list of the +specific and subspecific names that we consider to be applicable to +the <i>Hyla microcephala</i> group are listed below. +</p></div> +<p> </p> + +<div class="center"> +<table summary="Proposed Names"> +<tr><td><span class="smcap">Names Proposed</span></td><td><span class="smcap">Valid Names</span></td></tr> +<tr><td><i>Hyla cherrei</i> Cope, 1894</td><td>? = <i>H. m. microcephala</i></td></tr> +<tr><td><i>Hyla microcephala</i> Cope, 1886</td><td> = <i>H. m. microcephala</i></td></tr> +<tr><td><i>Hyla microcephala</i> Boulenger,<br> +<span style="margin-left: 1em;">1898 (<i>nec</i> Cope, 1886)</span></td><td> = <i>H. microcephala underwoodi</i></td></tr> +<tr><td><i>Hyla microcephala martini</i> Smith, 1951</td><td> = <i>H. microcephala underwoodi</i></td></tr> +<tr><td><i>Hyla microcephala sartori</i> Smith, 1951</td><td> = <i>H. sartori</i></td></tr> +<tr><td><i>Hyla phlebodes</i> Stejneger, 1906</td><td> = <i>H. phlebodes</i></td></tr> +<tr><td><i>Hyla robertmertensi</i> Taylor, 1937</td><td> = <i>H. robertmertensi</i></td></tr> +<tr><td><i>Hyla underwoodi</i> Boulenger, 1899</td><td> = <i>H. microcephala underwoodi</i></td></tr> +</table> +</div> +<p> </p> + +<p><span class="pagenum"><a name="Page_522" id="Page_522">[Pg 522]</a></span> +<i>Discussion.</i>—The color pattern is the most useful character in +distinguishing the species of the <i>Hyla microcephala</i> group from one +another. Except in <i>Hyla microcephala</i>, little geographic variation in +color pattern is noticeable. The features of color pattern that are +helpful in identifying the species are: 1) presence or absence of +lateral dark brown stripe; 2) longitudinal extent and width of lateral +stripe, if present; 3) presence or absence of a narrow white line just +dorsal to the lateral dark stripe; 4) presence or absence of an +interorbital dark mark; 5) the arrangement of dark markings on the +back, either as longitudinal lines or series of dashes, or in the form +of various kinds of transverse markings; 6) presence of dark flecks, +longitudinal line, or transverse marks on shanks.</p> + +<p>Few consistent differences in measurements and proportions exist among +the species (<a href="#Table_1">Table 1</a>). The most obvious morphological difference is +that the head is noticeably narrower in <i>H. robertmertensi</i> than in +the other species. <i>Hyla phlebodes</i> is the smallest species; adult +males attain snout-vent lengths of only 23.6 mm. The body is slender +in <i>H. microcephala</i> and <i>robertmertensi</i>, slightly wider in +<i>phlebodes</i>, and noticeably broader in <i>sartori</i>.</p> + +<p><i>Distribution.</i>—The composite range of the Middle American frogs of +the <i>Hyla microcephala</i> group includes the lowlands of southern México +and Central America, in some places to elevations of 1200 meters, +southeastward from southern Jalisco and southern Veracruz, excluding +arid regions (northern Yucatán Peninsula, Balsas-Tepalcatepec Basin, +Plains of Tehuantepec, Grijalva Valley, Salamá Basin, and upper +Motagua Valley) to the Pacific lowlands and the Cauca and Magdalena +valleys in Colombia.</p> +<p> </p> + +<a name="KEY_TO_SPECIES" id="KEY_TO_SPECIES"></a> +<div class="caption3">Key to Species and Subspecies</div> + +<table width="100%" summary="TTable Key"> +<tr><td class="vtop">1. </td><td class="txt-lf rt2em">Lateral dark stripe, bordered above by narrow white line, extending from snout at least to sacral region<span class="descrp2">2</span></td></tr> +<tr><td> </td><td class="txt-lf rt2em">Lateral dark stripe, if present, not extending posteriorly to sacral region and not bordered above by narrow white line<span class="descrp2">4</span></td></tr> + +<tr><td class="vtop">2.</td><td class="txt-lf rt2em">Lateral dark stripe continuous to groin; dark flecks or longitudinal line on shanks; interorbital dark bar absent; dorsal pattern usually consisting of pair of longitudinal dark lines or series of dashes<span class="descrp2">3</span></td></tr> + +<tr><td> </td><td class="txt-lf rt2em">Lateral dark stripe usually extending only to sacral region; dark transverse bars on shanks; interorbital bar usually present; dorsal pattern usually consisting of interconnecting dark lines, sometimes forming transverse marks<span class="descrp2"><i>H. microcephala underwoodi</i></span></td></tr> + +<tr><td class="vtop">3.</td><td class="txt-lf rt2em">Lateral dark stripe narrow, covering only upper edge of tympanum; dorsal longitudinal stripes continuous, extending to vent<span class="descrp2"><i>H. microcephala microcephala</i></span></td></tr> + +<tr><td> </td><td class="txt-lf rt2em">Lateral dark stripe wide, encompassing entire tympanum; dorsal markings consisting of longitudinal series of flecks or dashes, or of two lines, usually not extending to vent <span class="descrp2"><i>H. robertmertensi</i></span></td></tr> + +<tr><td colspan=2><p><span class="pagenum"><a name="Page_523" id="Page_523">[Pg 523]</a></span></p></td></tr> +<tr><td class="vtop">4.</td><td class="txt-lf rt2em">Lateral dark stripe indistinct, present only above tympanum and insertion of arm; dorsal markings consisting of narrow lines and dashes, sometimes interconnected; transverse bars on shanks narrow relative to interspaces<span class="descrp2"><i>H. phlebodes</i></span></td></tr> + +<tr><td> </td><td class="txt-lf rt2em">Lateral dark stripe absent; dorsal markings consisting of two broad chevron-shaped marks; transverse bars on shanks wide relative to interspaces<span class="descrp2"><i>H. sartori</i></span></td></tr> +</table> +<p> </p> +<p> </p> + +<a name="ACCOUNTS" id="ACCOUNTS"></a> +<div class="caption2">ACCOUNTS OF SPECIES AND SUBSPECIES</div> +<p> </p> + +<div class="caption2"><i>Hyla microcephala</i> Cope</div> + +<div class="blockquot"> +<p><i>Diagnosis.</i>—Lateral dark stripe narrow, covering only upper edge of +tympanum, bordered above by narrow white stripe; dorsal pattern +consisting of pair of longitudinal brown lines and no interorbital bar +(eastern populations), or of irregular dark markings forming an X- or +)(-shaped mark in scapular region and an interorbital bar (western +populations).</p> + +<p><i>Content.</i>—The populations inhabiting the Pacific lowlands of +southeastern Costa Rica eastward to Colombia are recognized herein as +<i>Hyla microcephala microcephala</i> Cope; the populations in western +Costa Rica northward to México are assigned to <i>Hyla microcephala +underwoodi</i> Boulenger.</p> + +<p><i>Distribution.</i>—Southern Veracruz and northern Oaxaca southeastward +through the Atlantic lowlands of Central America to north-central +Nicaragua, thence southeastward on the Pacific lowlands to eastern +Panamá, and thence into the Cauca and Magdalena valleys (Caribbean +drainage) of Colombia (<a href="#Fig_1">Fig. 1</a>).</p> +</div> +<p> </p> +<p> </p> + +<a name="Fig_1"></a> +<div class="center"> +<img src="images/fig_1.png" width="600" height="469" title="Locality Map - Hyla microcephala" alt="Locality Map - Hyla microcephala"><br><br> +<span class="smcap">Fig. 1.</span> Map showing locality records for <i>Hyla microcephala</i>.<br> +</div> +<p> </p> +<p> </p> + +<a name="Table_1"></a> +<span class="pagenum"><a name="Page_524" id="Page_524">[Pg 524]</a></span> +<span class="smcap">Table 1.</span>—Variation in Certain Measurements and Properties in the Hyla microcephala Group. (All Data Based on Adult Males;<br> +Mean and Standard Error of Mean Below Observed Range.)<br> +<br> + +<table class="data" width="100%" summary="Size Variation in Hyla microcephala Group"> +<tr><td colspan=8 class="bb"> </td></tr> +<tr><td class="bt bb">Locality</td><td class="bt bl bb">N</td><td class="bt bl bb">Snout-vent<br>length (S-V L)</td><td class="bt bl bb">Tibia length<br><img src="images/bar_single.png" width="55" height="15" title="bar" alt="bar"><br>S-V L</td><td class="bt bl bb">Foot length<br><img src="images/bar_single.png" width="55" height="15" title="bar" alt="bar"><br>S-V L</td><td class="bt bl bb">Head length<br><img src="images/bar_single.png" width="55" height="15" title="bar" alt="bar"><br>S-V L</td><td class="bt bl bb">Head width<br><img src="images/bar_single.png" width="55" height="15" title="bar" alt="bar"><br>S-V L</td><td class="bt bl bb">Tympanum<br><img src="images/bar_single.png" width="55" height="15" title="bar" alt="bar"><br>Eye</td></tr> +<tr><td> </td><td colspan=7 class="center bl"><br><i>H. m. microcephala</i></td></tr> +<tr><td class="text_lf">Panamá: Canal Zone</td><td class="bl">25</td><td class="bl">21.5-24.1</td><td class="bl">50.2-56.0</td><td class="bl">40.9-46.6</td><td class="bl">28.5-32.8</td><td class="bl">28.1-30.9</td><td class="bl">44.0-54.1</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">22.8±0.20</td><td class="bl">52.9±0.37</td><td class="bl">43.5±0.28 </td><td class="bl">31.0±0.22 </td><td class="bl">29.4±0.11</td><td class="bl">49.0±0.55</td></tr> +<tr><td> </td><td colspan=7 class="bl"> </td></tr> +<tr><td class="text_lf">Costa Rica: Golfito</td><td class="bl">25</td><td class="bl">18.5-24.5</td><td class="bl">49.1-54.4</td><td class="bl">41.8-48.0</td><td class="bl">30.2-35.5</td><td class="bl">29.0-32.7</td><td class="bl">40.0-57.8</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">22.4±0.27</td><td class="bl">51.6±0.26</td><td class="bl">45.1±0.32 </td><td class="bl">33.1±0.25 </td><td class="bl">30.8±0.16</td><td class="bl">48.4±1.10</td></tr> +<tr><td> </td><td colspan=7 class="center bl"><br><i>H. m. underwoodi</i><br></td></tr> +<tr><td class="text_lf">Nicaragua: La Cumplida </td><td class="bl">25 </td><td class="bl">23.0-25.6 </td><td class="bl">51.0-55.7</td><td class="bl">41.3-46.5</td><td class="bl">29.7-33.5</td><td class="bl">28.9-31.8 </td><td class="bl">42.3-60.0</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">24.1±0.19</td><td class="bl">52.9±0.25</td><td class="bl">43.7±0.25 </td><td class="bl">31.6±0.19 </td><td class="bl">30.4±0.17</td><td class="bl">49.3±0.97</td></tr> +<tr><td> </td><td colspan=7 class="bl"> </td></tr> +<tr><td class="text_lf">Guatemala: Finca Chamá </td><td class="bl">25 </td><td class="bl">21.8-25.0 </td><td class="bl">51.0-57.2</td><td class="bl">41.2-47.8</td><td class="bl">30.8-35.3</td><td class="bl">29.6-33.6 </td><td class="bl">37.5-56.4</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">23.5±0.16</td><td class="bl">54.3±0.39</td><td class="bl">44.4±0.30 </td><td class="bl">33.0±0.16 </td><td class="bl">31.3±0.36</td><td class="bl">45.2±0.89</td></tr> +<tr><td> </td><td colspan=7 class="bl"> </td></tr> +<tr><td class="text_lf">Tabasco: Teapa</td><td class="bl">25 </td><td class="bl">22.7-25.8 </td><td class="bl">48.0-54.5</td><td class="bl">40.7-46.8</td><td class="bl">29.5-33.0</td><td class="bl">28.7-31.8 </td><td class="bl">40.7-53.8</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">24.3±0.14</td><td class="bl">51.5±0.29</td><td class="bl">43.3±0.25 </td><td class="bl">31.7±0.17 </td><td class="bl">30.3±0.16</td><td class="bl">45.5±0.38</td></tr> +<tr><td> </td><td colspan=7 class="bl"> </td></tr> +<tr><td class="text_lf">Oaxaca: Donají-Sarabia </td><td class="bl">25 </td><td class="bl">22.1-25.9 </td><td class="bl">49.8-55.6</td><td class="bl">40.5-46.6</td><td class="bl">30.4-34.8</td><td class="bl">28.9-32.6 </td><td class="bl">37.0-54.1</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">23.8±0.19</td><td class="bl">52.8±0.33</td><td class="bl">43.4±0.27 </td><td class="bl">32.8±0.19 </td><td class="bl">30.8±0.17</td><td class="bl">45.1±0.76</td></tr> +<tr><td> </td><td colspan=7 class="bl"> </td></tr> +<tr><td class="text_lf">Veracruz: Alvarado</td><td class="bl">25 </td><td class="bl">21.9-25.4 </td><td class="bl">49.6-54.4</td><td class="bl">40.7-47.5</td><td class="bl">29.9-33.8</td><td class="bl">29.1-32.9 </td><td class="bl">40.7-53.8</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">24.1±0.17</td><td class="bl">51.1±0.28</td><td class="bl">42.6±0.34 </td><td class="bl">31.4±0.18 </td><td class="bl">30.5±0.17</td><td class="bl">46.6±0.65</td></tr> +<tr><td> </td><td colspan=7 class="center bl"><p><span class="pagenum"><a name="Page_525" id="Page_525">[Pg 525]</a></span></p><i>H. robertmertensi</i><br></td></tr> +<tr><td class="text_lf">Guatemala: La Trinidad</td><td class="bl">21</td><td class="bl">21.8-24.6</td><td class="bl">47.1-52.8</td><td class="bl">40.9-51.3</td><td class="bl">30.0-33.3</td><td class="bl">27.3-29.8</td><td class="bl">44.4-50.0</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">23.4±0.15</td><td class="bl">49.9±0.34</td><td class="bl">43.5±0.17</td><td class="bl"> 31.3±0.20</td><td class="bl"> 28.5±0.23</td><td class="bl"> 47.4±0.46</td></tr> +<tr><td> </td><td colspan=7 class="bl"> </td></tr> +<tr><td class="text_lf">Chiapas: Acacoyagua</td><td class="bl">25</td><td class="bl"> 21.4-25.7</td><td class="bl"> 47.8-52.4</td><td class="bl">41.7-46.3</td><td class="bl">29.1-32.7</td><td class="bl">26.0-30.3</td><td class="bl">42.8-53.8</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">24.1±0.20</td><td class="bl">50.4±0.45</td><td class="bl">43.9±0.23</td><td class="bl">31.2±0.29</td><td class="bl">28.1±0.20</td><td class="bl"> 46.5±0.50</td></tr> +<tr><td> </td><td colspan=7 class="bl"> </td></tr> +<tr><td class="text_lf">Oaxaca: Tapanatepec</td><td class="bl">25</td><td class="bl">22.4-26.4</td><td class="bl">44.1-48.3</td><td class="bl">39.1-44.5</td><td class="bl">26.1-30.4</td><td class="bl"> 25.4-28.1</td><td class="bl"> 45.8-58.3</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">24.7±0.18</td><td class="bl">46.4±0.23</td><td class="bl">41.7±0.23</td><td class="bl">28.4±0.16</td><td class="bl">26.8±0.14</td><td class="bl"> 52.9±0.77</td></tr> +<tr><td> </td><td colspan=7 class="center bl"><br><i>H. phlebodes</i><br><br> +<tr><td class="text_lf">Panamá: Canal Zone</td><td class="bl">25</td><td class="bl">19.6-23.2</td><td class="bl"> 49.1-56.9</td><td class="bl">41.9-47.1</td><td class="bl">33.6-37.4</td><td class="bl">32.3-36.0</td><td class="bl">37.9-46.4</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">22.2±0.16</td><td class="bl">52.8±0.35</td><td class="bl"> 45.4±0.26 </td><td class="bl">34.8±0.18</td><td class="bl">33.8±0.18</td><td class="bl"> 41.6±0.49</td></tr> +<tr><td> </td><td colspan=7 class="bl"> </td></tr> +<tr><td class="text_lf">Costa Rica: Turrialba</td><td class="bl">25</td><td class="bl">19.7-23.6</td><td class="bl">47.4-55.7</td><td class="bl">38.1-46.4</td><td class="bl">32.6-35.9</td><td class="bl">30.5-35.0</td><td class="bl">35.7-48.2</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl"> 22.0±0.18</td><td class="bl">51.1±0.35</td><td class="bl">42.8±0.38</td><td class="bl">34.1±0.16</td><td class="bl">32.9±0.17</td><td class="bl"> 40.1±0.53</td></tr> +<tr><td> </td><td colspan=7 class="center bl"><br><i>H. sartori</i></td></tr> +<tr><td class="text_lf">Guerrero: Tierra Colorada </td><td class="bl"> 25 </td><td class="bl">23.7-26.0</td><td class="bl">47.2-51.4</td><td class="bl"> 42.4-47.8</td><td class="bl">29.4-31.8</td><td class="bl"> 28.9-31.0</td><td class="bl">42.3-52.0</td></tr> +<tr><td class="bb"> </td><td class="bl bb"> </td><td class="bl bb">24.8±0.13</td><td class="bl bb">49.6±0.23</td><td class="bl bb">45.2±0.27</td><td class="bl bb">30.6±0.13</td><td class="bl bb"> 30.0±0.12</td><td class="bl bb"> 47.4±0.59</td></tr> +</table> +<p> </p> +<p> </p> + +<p><span class="pagenum"><a name="Page_526" id="Page_526">[Pg 526]</a></span></p> + +<div class="caption3nb"><span class="bold"><i>Hyla microcephala microcephala</i></span> Cope</div> + +<div class="species"> +<p><i>Hyla microcephala</i> Cope, Proc. Amer. Philos. Soc., 23:281, February +11, 1886 [Syntypes.—USNM 13473 (2 specimens, now lost) from Chiriquí, +Panamá; Mr. MacNeil collector]; Bull. U.S. Natl. Mus., 32:14, 1887. +Günther, Biologia-Centrali Americana, Reptilia and Batrachia, p. 265, +June, 1901. Dunn, Occas. Papers Boston Soc. Nat. Hist., 5:413, October +10, 1931; Occas. Papers Boston Soc. Nat. Hist., 8:72, June 7, 1933. +Stebbins and Hendrickson, Univ. California Publ. Zool., 56:524, +February 17, 1959. Fouquette, Evolution, 14:484, December 16, 1960. +Busack, Copeia, 2:371, June 21, 1966.</p> + +<p>? <i>Hyla cherrei</i> Cope, Proc. Acad. Nat. Sci. Philadelphia, 1894, p. +195, 1894 [Holotype.—location unknown, apparently lost; +type-locality: "Alajuela, Costa Rica;" R. Alfaro collector]. Günther, +Biologia Centrali-Americana: Reptilia and Batrachia, p. 264, June, +1901. Taylor, Univ. Kansas Sci. Bull., 35:846, July 1, 1952.</p> + +<p><i>Hyla underwoodi</i>, Ruthven, Misc. Publ. Mus. Zool., Univ. Michigan, +8:55, September 15, 1922. Barbour, Proc. New England Zool. Club, +10:31, March 2, 1928.</p> + +<p><i>Hyla microcephala microcephala</i>, Smith, Herpetologica, 7:185, +December 31, 1951. Taylor, Univ. Kansas Sci. Bull., 39:23, November +18, 1958.</p> +</div> + +<p><i>Diagnosis.</i>—Brown lateral stripe narrow, extending from nostril +along canthus, along upper edge of tympanum to groin, bordered above +by narrow white line; pair of dark brown longitudinal lines on dorsum +extending to vent; shanks having dark longitudinal line or flecks, no +transverse bars; interorbital dark mark lacking.</p> + +<p><i>Description and Variation.</i>—The color pattern is nearly constant. Of +103 males from the Canal Zone, all lack an interorbital dark bar, and +all have a dark longitudinal line on the dorsal surface of the shank +and a narrow lateral dark stripe, bordered above by a narrow white +line, extending to the groin. The longitudinal dark lines on the +dorsum are continuous to the groin in 95 specimens and fragmented in +two specimens. In two others the lines converge and fuse in the +scapular region, and in four specimens auxiliary, fragmented lines are +present dorsolaterally.</p> + +<p>In all specimens from southeastern Costa Rica (Golfito, Palmar Sur, +and Villa Neilly) the pattern is constant, except that in about 10 per +cent of the specimens the longitudinal line on the dorsal surface of +the shank is replaced by a row of brown flecks.</p> + +<p>Of the limited number of Colombian specimens examined, all are +patterned normally, except three from Sautata, Chocó, three from +Curumani, and three from Arcataca, Magdalena, which have flecks on the +dorsal surfaces of the shanks, and one from Espinal, Tolima, which has +no markings on the shanks.</p> + +<p>When active at night most individuals are pale yellowish tan dorsally; +the white dorsolateral line is noticeable, but the brown lateral +stripe, dorsal brown lines, and lines on shanks are so pale that often +they are barely discernible. By day the dorsum changes to tan or pale +reddish brown; the stripes are dark brown, and the dorsolateral stripe +that is white at night becomes creamy yellow (<a href="#Pl_13">Pl. 13</a>). Small brown +flecks are present on the dorsum of most individuals. The venter +always is white, and the iris is pale bronze with a brown tint +immediately anterior and posterior to the pupil. In breeding males the +vocal sac is pale yellow.</p> + +<p><i>Tadpoles.</i>—Tadpoles of this species have been found in weed-choked +ponds in eastern Panamá Province. The following description is based +on KU 104097, a specimen in developmental stage 34 (Gosner, 1960).</p> + +<p><span class="pagenum"><a name="Page_527" id="Page_527">[Pg 527]</a></span> +Total length, 20.5 mm.; body length, 8.2 mm.; body slightly wider +than deep; snout pointed; nostrils large, situated dorsally, much +closer to snout than eyes, directed anteriorly; eyes moderately +small, situated dorsolaterally and directed laterally; spiracle +sinistral, located just posteroventral to eye; anal tube dextral. +Tail xiphicercal; caudal musculature moderately deep, becoming +slender posteriorly, extending beyond caudal fin; fins deepest at +about one-third distance from body to tip of tail; dorsal fin +extending onto body, deeper than deepest part of caudal +musculature; ventral fin slightly shallower than musculature. +Mouth small, terminal, lacking teeth and fringing papillae, but +having finely serrate beaks. In preservative, top of head pale +brown; dark stripe from tip of snout through eye to posterior edge +of body, narrowing to thin line on proximal one-fourth of tail; +venter white; tail creamy tan with fine black flecks most numerous +posteriorly; posterior two-thirds of fins edged with black. In +life, top of head yellowish tan; lateral stripe brown; belly +white; anterior half of tail lacking pigment; posterior half deep +orange; iris pale bronze (<a href="#Pl_15">Pl. 15</a>).</p> + +<div class="remarks"> +<p><i>Remarks.</i>—Evidence for intergradation of <i>Hyla microcephala</i> with +<i>H. underwoodi</i> is provided by four specimens [USC 818 (2), 6081-2] +from 6.1 kilometers northeast of the mouth of the Río Tarcoles, and +nine specimens [USC 8254 (2), 8255, 8256 (4), 8258 (2)] from Parrita, +both in Puntarenas Province, Costa Rica. These localities lie about +two-thirds the distance from the northwesternmost locality for <i>H. +m. microcephala</i> (Palmar Sur) to the southeasternmost locality for +<i>H. m. underwoodi</i> (Barranca). Although in most aspects of coloration +the frogs are more nearly like <i>H. m. underwoodi</i> than <i>H. m. +microcephala</i>, some specimens have longitudinal lines on their shanks, +such as are characteristic of <i>H. m. microcephala</i>. The dorsal pattern +varies from nearly complete longitudinal lines to broken lines, fused +into an X-shaped scapular mark or not.</p> + +<p>As noted by Rivero (1961:135), <i>Hyla microcephala</i> seems to be closely +related to <i>Hyla misera</i> Werner, a species having a wide distribution +east of the Andes in South America. Despite the similarity in color +pattern, size, and structure, we are reluctant to place the two taxa +in the same species until data on coloration in life, mating calls, +and life history are available for <i>Hyla misera</i> and compared with +those of <i>Hyla microcephala</i>.</p> + +<p>The status of Cope's <i>Hyla cherrei</i> is questionable. Since the type, +the only specimen ever referred to the species, apparently is lost, +the only extant information regarding the taxon is contained in the +original description (Cope, 1894). There the species was characterized +as having a narrow dorsolateral white stripe and lacking pigment on +the upper arms and thighs. These characteristics of the color pattern +combined with the statements "vomerine teeth few, opposite the middle +of the very large choanae" and +<span class="pagenum"><a name="Page_528" id="Page_528">[Pg 528]</a></span> +"tympanic drum distinct, one half the area of eye" serve to +distinguish <i>H. cherrei</i> from all other Costa Rican hylids, except <i>H. +m. microcephala</i> and <i>H. m. underwoodi</i>. No statements in the type +description will definitely associate <i>cherrei</i> with one or the other +of these subspecies. Since it seems obvious that <i>H. cherrei</i> can be +associated with <i>H. microcephala</i>, we prefer to place the name in the +synonymy of the nominate subspecies, thereby preserving the commonly +used name <i>H. underwoodi</i> (Boulenger, 1899) as a subspecies of <i>H. +microcephala</i>.</p> +</div> + +<p><i>Distribution.</i>—<i>Hyla microcephala microcephala</i> inhabits coastal +lowlands from the area of Golfo Dulce (apparently absent from the +Osa Peninsula) in southeastern Costa Rica eastward in Panamá, +including the Azuero Peninsula to northern Colombia and thence +southward in the valleys of the Río Cauca and Río Magdalena in +Colombia (<a href="#Fig_1">Fig. 1</a>). Except for the central area of the Canal Zone +the subspecies is unknown from the Caribbean drainage in Central +America, but in Colombia the subspecies occurs only in the +Caribbean drainage. In Central America this frog occurs mostly on +the coastal lowlands; the highest recorded elevation is 560 meters +at El Valle, Coclé, Panamá. Throughout most of its range <i>Hyla +microcephala microcephala</i> occurs in disturbed habitats—cut-over +forests, secondary growth, and pastureland. It does not seem to be +an inhabitant of either primary forest or of <i>Curatella</i>-savanna.</p> + +<p><i>Specimens examined.</i>—522, as follows: <b>Costa Rica</b>: <span class="smcap">Puntarenas</span>: +Golfito, KU 32172-207; 3 km. E Golfito, KU 86399, USC 2757-8; +Palmar Sur, KU 64591-608, USC 2650 (14), UU 3907-32; *1.5-2.5 km. +ESE Palmar Sur, KU 68293-7 (skeletons), 93957-62; Parrita, USC +8254 (2), 8255, 8256 (4), 8258 (2) [intergrades with <i>H. m. +underwoodi</i>]; 3 km. NW Piedras Blancas, KU 103689; 6.1 km. NE +mouth of Río Tárcoles, USC 818 (2), 6081-2 [intergrades with <i>H. +m. underwoodi</i>]; Villa Neilly, USC 2651; *1-5 km. WNW Villa +Neilly, USC 6182-4, 8003 (4), 8031 (3), 8032; *10.5 km. WNW Villa +Neilly, KU 64609-27, 68398 (eggs).</p> + +<p><b>Panama</b>: <span class="smcap">Canal Zone</span>: Albrook Air Base, TNHC 23389, 23497; Balboa, +ANSP 19555-6; *Fort Clayton, UIMNH 42008-12; *2.8 km. SW Fort +Kobbe, KU 96015-25; *Frijoles, MCZ 19208; *Bamboa, MCZ 21507; *8.3 +km. N Gatún Locks, TNHC 23441; *Juan Diaz, MCZ 13747; *Juan Mina, +AMNH 55436-7, ANSP 21811-2, UMMZ 126734, 126735 (6), UU 3900-6; +*8-14 km. N Miraflores Locks, TNHC 23374-88, 23390-409, 23411-38, +23440, 23442-60, 23462-76; 23478-83, 23492, 23555-60, 23562-76; +*Río Chagres, AMNH 55430, 55439; *Río Cocolí, 3.5 km. N Miraflores +Locks, TNHC 23410; *Summit, ANSP 23365-71, FMNH 22966-9, KU +97783-87. <span class="smcap">Chiriqui</span>: 5.5 km. E Concepción, AMNH 69772; *14.4 km. E +Concepción, AMNH 69773-8; 2 km. S David, AMNH 69779; *Progreso, +UMMZ 58252, 58253 (2), 58254, 58436; Río Gariché, 8.3 km. ESE Paso +Canoas, KU 103065-8. <span class="smcap">Coclé</span>: 1 km. SE El Caño, KU 103042-51; El +Valle de Antón, AMNH 59614-18 (10), 69785, ANSP 23502-5, KU +77201-14, MVZ 66578-83, UIMNH 46532. <span class="smcap">Colón</span>: Cement Plant, +Transisthmian Highway, FMNH 60394-5. <span class="smcap">Darién</span>: El Real, KU 80454-5, +103052-64, UMMZ 125036 (10), USNM 140567-8; Río Canclon at Río +Chucunaque, UMMZ 125035; *Río Chucunaque, near Yavisa, AMNH 59523. +<span class="smcap">Los Santos</span>: Tonosí, KU 101606-9. <span class="smcap">Panamá</span>: 5 km. S Bejuco, AMNH +69782; 3 km. W Chepo, KU 77172-4, 104097-8 (tadpoles); *6 km. WSW +Chepo, KU 77175; *Chico, Río La Jagua, USNM 129070; *La Joya, +Cacora, ANSP 25129-33; Madden Dam, FMNH 67819; Nueva Gorgona, AMNH +69780-1; *1.6 km. W Nueva Gorgona, AMNH 69783-4; 1.5 km. W Pacora, +77176-200; *Río La Laja, near Chamé, ANSP 21845; *Río Tapia, MCZ +10048; *Tapia, AMNH 18930, 18950, 18952-3; *18 km. E Tocumen, MVZ +78662.</p> + +<p><span class="pagenum"><a name="Page_529" id="Page_529">[Pg 529]</a></span> +<b>Colombia</b>: <span class="smcap">Chocó</span>: Sautatá, Atrato, FMNH 74918 (2), 74919. +<span class="smcap">Magdalena</span>: Aracataca, ANSP 19755-7; Curumani, MCZ 21465-74, UIMNH +28855; UMMZ 90168, USNM 118247; El Banco, Río Magdalena, ANSP +25061; Fundación, UMMZ 48281-2. <span class="smcap">Tolima</span>: Espinal, MCZ 15068; +Mariquita, FMNH 81822-3. <span class="smcap">Valle</span>: Sevilla, MCZ 13751-3.</p> +<p> </p> +<p> </p> + +<div class="caption3nb"><i>Hyla microcephala underwoodi</i> Boulenger</div> + +<div class="species"> +<p><i>Hyla microcephala</i> Boulenger, Proc. Zool. Soc. London, p. 481, +October 1, 1898 [Syntypes.—BMNH 94. 11. 1532-33 from Bebedero, +Guanacaste Province, Costa Rica; C. F. Underwood collector] (not +<i>Hyla microcephala</i> Cope, Proc. Amer. Philos. Soc., 23:281, +February 11, 1886, from Chiriquí, Panamá).</p> + +<p><i>Hyla underwoodi</i> Boulenger, Ann. Mag. Nat. Hist., ser. 7, 3:277, +April, 1899 (substitute name for <i>Hyla microcephala</i> Boulenger, +preoccupied). Günther, Biologia-Centrali Americana, Reptilia and +Batrachia, p. 278, September, 1901. Dunn and Emlen, Proc. Acad. +Nat. Sci. Philadelphia, 84:25, March 22, 1932. Stuart, Misc. Publ. +Mus. Zool., Univ. Michigan, 29:39, October 1, 1935. Taylor, Proc. +Biol. Soc. Washington, 50:44, April 21, 1937. Stuart, Occas. +Papers Mus. Zool., Univ. Michigan, 471:15, May 17, 1943. Taylor +and Smith, Proc. U. S. Natl. Mus., 95:586, June 30, 1945. Stuart, +Misc. Publ. Mus. Zool., Univ. Michigan, 69:35, June 12, 1948. +Smith and Taylor, Bull. U. S. Natl. Mus., 194:85, June 17, 1948; +Univ. Kansas Sci. Bull., 33:316, March 20, 1950. Stuart, Contr. +Lab. Vert. Biol., Univ. Michigan, 45:48, May, 1950. Taylor, Univ. +Kansas Sci. Bull., 35:891, July 1, 1952; Univ. Kansas Sci. Bull., +39:25, November 18, 1958.</p> + +<p><i>Hyla phlebodes</i>, Cole and Barbour, Bull. Mus. Comp. Zool., +50:154, November, 1906. Kellogg, Bull. U. S. Natl. Mus., 160:172, +March 31, 1932.</p> + +<p><i>Hyla microcephala martini</i> Smith, Herpetologica, 7:187, December +31, 1951 [Holotype.—UIMNH 20965 from Encarnacion, Campeche, +México; H. M. Smith collector]. Stuart, Contr. Lab. Vert. Biol., +Univ. Michigan, 68:46, November, 1954. Fugler and Webb, +Herpetologica, 13:105, July 10, 1957. Stuart, Contr. Lab. Vert. +Biol., Univ. Michigan, 75:17, June, 1958. Neill and Allen, Publ. +Research Div., Ross Allen's Reptile Inst., 2:26, November 10, 1959. +Duellman, Univ. Kansas Publ., Mus. Nat. Hist., 13:62, August 16, +1960. Stuart, Herpetologica, 17:74, July 11, 1961. Hensley and +Smith, Herpetologica, 18:70, April 9, 1962. Stuart, Misc. Publ. +Mus. Zool., Univ. Michigan, 122:36, April 2, 1963. Holman and +Birkenholz, Herpetologica, 19:144, July 3, 1963. Duellman, Univ. +Kansas Publ., Mus. Nat. Hist., 15:225, October 4, 1963; Univ. +Kansas Publ., Mus. Nat. Hist., 15:588, June 22, 1965.</p> + +<p><i>Hyla microcephala underwoodi</i>, Smith, Herpetologica, 7:188, +December 31, 1951.</p> +</div> + +<p><i>Diagnosis.</i>—Brown lateral stripe narrow, extending to groin or +only to sacral region, bordered above by narrow white line; dorsal +pattern bold, consisting of X- or )(-shaped mark in scapular +region or pair of interconnected dark lines on back; interorbital +dark mark usually present; shanks usually having dark transverse +bars.</p> + +<p><i>Description and Variation.</i>—The dorsal color pattern is highly +variable. The various permutations of the X-shaped scapular mark +and dark sacral marks differ proportionately in different samples. +The variation in color pattern in 12 samples is summarized in +<a href="#Table_2">Table 2</a>. In samples from the southern part of the range (southern +Nicaragua and Guanacaste Province, Costa Rica) more (40-93%) +individuals have the lateral stripes extending to the groin than +in northern samples (0-42%) from southern México and Guatemala. +Likewise, the percentage of specimens lacking bars on the shanks +and a dark interorbital bar is higher in the Costa Rican samples +than elsewhere in the range. The X- or )(-shaped scapular markings +and /\- or / \-shaped sacral markings are most prevalent in northern samples, +whereas to the south the dorsal markings are more commonly arranged in a +pattern of paired lines, which usually are discontinuous and usually extend +posteriorly only to the sacral region. Thus, the color pattern in <i>H. m. +underwoodi</i> in the southern part of its range shows trends towards the +pattern characteristic of <i>H. m. microcephala</i>. Intergrades between +these two subspecies have been discussed in the account of the +nominate subspecies.</p> +<p> </p> +<p> </p> + +<a name="Table_2"></a> +<p><span class="pagenum"><a name="Page_530" id="Page_530">[Pg 530]</a></span></p> +<div class="center"><span class="smcap">Table 2.</span>--Variation in Color Pattern in Hyla microcephala underwoodi</div> +<div> +<table width="100%" class="data" cellpadding=4 summary="Variation in Color Pattern of Hyla microcephala"> +<tr><td class="bb" colspan=19><hr style="height:2px; color:#000;"></td></tr> +<tr><td class="bb" rowspan=2><span class="smcap">Population</span></td><td rowspan=2 class="bl bb">N</td><td colspan=2 class="bb bl">Shanks</td><td class="bl"> </td><td colspan=2 class="bb bl">Interorbital bar</td><td class="bl"> </td><td colspan=2 class="bb bl">Dorsolateral stripe</td><td class="bl"> </td><td colspan=4 class="bb bl">Scapular markings</td><td class="bl"> </td><td colspan=3 class="bb bl">Sacral markings</td></tr> +<tr><td class="bb bl"> Bars </td><td class="bb bl">Flecks</td><td class="bl"> </td><td class="bb bl">Present</td><td class="bb bl">Absent</td><td class="bl"> </td><td class="bb bl">Groin</td><td class="bb bl">Sacrum</td><td class="bl"> </td><td class="bb bl">X</td><td class="bb bl">)(</td><td class="bb bl">][</td><td class="bb bl">Other</td><td class="bl"> </td><td class="bb bl">/\</td><td class="bb bl">/ \</td><td class="bb bl">Other</td></tr> +<tr><td class="text_lf">Oaxaca: Donají-Sarabia</td><td class="bl">27</td><td class="bl">22</td><td class="bl">5</td><td class="bl"> </td><td class="bl">27</td><td class="bl">0</td><td class="bl"> </td><td class="bl">0</td><td class="bl">27</td><td class="bl"> </td><td class="bl">23</td><td class="bl">4</td><td class="bl">0</td><td class="bl">0</td><td class="bl"> </td><td class="bl">7</td><td class="bl">6</td><td class="bl">14</td></tr> +<tr><td class="text_lf">Tabasco: Teapa-Villahermosa</td><td class="bl">55 </td><td class="bl">46</td><td class="bl">9</td><td class="bl"> </td><td class="bl">55</td><td class="bl">0</td><td class="bl"> </td><td class="bl">0</td><td class="bl">55</td><td class="bl"> </td><td class="bl">53 </td><td class="bl">2</td><td class="bl">0</td><td class="bl">0</td><td class="bl"> </td><td class="bl">19</td><td class="bl">11</td><td class="bl">23</td></tr> +<tr><td class="text_lf">Guatemala: La Libertad</td><td class="bl">51 </td><td class="bl">51</td><td class="bl">0</td><td class="bl"> </td><td class="bl">51</td><td class="bl">0</td><td class="bl"> </td><td class="bl">17</td><td class="bl">34</td><td class="bl"> </td><td class="bl">45</td><td class="bl">6</td><td class="bl">0</td><td class="bl">0</td><td class="bl"> </td><td class="bl">16 </td><td class="bl">14</td><td class="bl">21</td></tr> +<tr><td class="text_lf">Guatemala: Finca Chamá</td><td class="bl">32 </td><td class="bl">32</td><td class="bl">0</td><td class="bl"> </td><td class="bl">32</td><td class="bl">0</td><td class="bl"> </td><td class="bl">0</td><td class="bl">32</td><td class="bl"> </td><td class="bl">32 </td><td class="bl">0</td><td class="bl">0</td><td class="bl">0</td><td class="bl"> </td><td class="bl">26 </td><td class="bl">2</td><td class="bl">4</td></tr> +<tr><td class="text_lf">Guatemala: Puerto Barrios</td><td class="bl">31</td><td class="bl">31</td><td class="bl">0</td><td class="bl"> </td><td class="bl">31</td><td class="bl">0</td><td class="bl"> </td><td class="bl">14</td><td class="bl">17</td><td class="bl"> </td><td class="bl">23 </td><td class="bl">0</td><td class="bl">4</td><td class="bl">4</td><td class="bl"> </td><td class="bl">6</td><td class="bl">4</td><td class="bl">21</td></tr> +<tr><td class="text_lf">Honduras: Lago Yojoa</td><td class="bl">13</td><td class="bl">13 </td><td class="bl">0</td><td class="bl"> </td><td class="bl">13</td><td class="bl">0</td><td class="bl"> </td><td class="bl">9</td><td class="bl">4</td><td class="bl"> </td><td class="bl">3 </td><td class="bl">2</td><td class="bl">3</td><td class="bl">5</td><td class="bl"> </td><td class="bl">2</td><td class="bl">1</td><td class="bl">10</td></tr> +<tr><td class="text_lf">Nicaragua: La Cumplida</td><td class="bl">56</td><td class="bl">44</td><td class="bl">12</td><td class="bl"> </td><td class="bl">54</td><td class="bl">2</td><td class="bl"> </td><td class="bl">13</td><td class="bl">43</td><td class="bl"> </td><td class="bl">11</td><td class="bl">35</td><td class="bl">8</td><td class="bl">2</td><td class="bl"> </td><td class="bl">0</td><td class="bl">19</td><td class="bl">37</td></tr> +<tr><td class="text_lf">Nicaragua: Tipitapa </td><td class="bl">10 </td><td class="bl">10 </td><td class="bl">0</td><td class="bl"> </td><td class="bl">10</td><td class="bl">0</td><td class="bl"> </td><td class="bl">8</td><td class="bl">2</td><td class="bl"> </td><td class="bl">0 </td><td class="bl">5</td><td class="bl">3</td><td class="bl">2</td><td class="bl"> </td><td class="bl">0</td><td class="bl">3</td><td class="bl">7</td></tr> +<tr><td class="text_lf">Nicaragua: Santo Thomás</td><td class="bl">10 </td><td class="bl">10</td><td class="bl">0</td><td class="bl"> </td><td class="bl">10</td><td class="bl">0</td><td class="bl"> </td><td class="bl">8</td><td class="bl">2</td><td class="bl"> </td><td class="bl">3</td><td class="bl">0</td><td class="bl">7</td><td class="bl">0</td><td class="bl"> </td><td class="bl">0</td><td class="bl">5</td><td class="bl">5</td></tr> +<tr><td class="text_lf">Costa Rica: Tenorio-Tilarán</td><td class="bl">12 </td><td class="bl">0 </td><td class="bl">12</td><td class="bl"> </td><td class="bl">6</td><td class="bl">6</td><td class="bl"> </td><td class="bl">7</td><td class="bl">5</td><td class="bl"> </td><td class="bl">0 </td><td class="bl">0</td><td class="bl">12</td><td class="bl">0</td><td class="bl"> </td><td class="bl">0 </td><td class="bl">0</td><td class="bl">12</td></tr> +<tr><td class="text_lf">Costa Rica: Las Cañas-Liberia</td><td class="bl">38</td><td class="bl center">21<a name="FNanchor_A_1"></a><a href="#Footnote_A_1"><span class="fnanchor">[A]</span></a></td><td class="bl">15 </td><td class="bl"> </td><td class="bl">34</td><td class="bl">4</td><td class="bl"> </td><td class="bl">25</td><td class="bl">13</td><td class="bl"> </td><td class="bl">0</td><td class="bl">11</td><td class="bl">19</td><td class="bl">8</td><td class="bl"> </td><td class="bl">0</td><td class="bl">0</td><td class="bl">38</td></tr> +<tr><td class="text_lf bb">Costa Rica: Esparta</td><td class="bb bl">32</td><td class="bb bl">26 </td><td class="bb bl">6</td><td class="bb bl"> </td><td class="bb bl">29</td><td class="bb bl">3</td><td class="bb bl"> </td><td class="bb bl">30</td><td class="bb bl">2</td><td class="bb bl"> </td><td class="bb bl">0 </td><td class="bb bl">0</td><td class="bb bl">14</td><td class="bb bl">18</td><td class="bb bl"> </td><td class="bb bl">0 </td><td class="bb bl">0</td><td class="bb bl">32</td></tr> +</table><br> +<div class="footnote"><a name="Footnote_A_1" id="Footnote_A_1"></a><p><a href="#FNanchor_A_1">[A]</a> Longitudinal stripes present in two specimens.</p></div> +<p> </p> +<p> </p> + +<p><span class="pagenum"><a name="Page_531" id="Page_531">[Pg 531]</a></span> +When this frog is active at night its dorsum is pale yellow; faint +flecks are present in some individuals. The white dorsolateral line +usually is evident in the tympanic region, but in many individuals a +dorsal pattern of lines and other marks is not evident. By day the +dorsum changes to yellowish tan or pale brown with dark brown or +reddish brown markings (<a href="#Pl_13">Pl. 13</a>). The venter is white, and the vocal +sac in breeding males is yellow. The iris is pale bronze with a brown +tint anterior and posterior to the pupil.</p> + +<div class="remarks"> +<p><i>Remarks.</i>—<i>Hyla microcephala underwoodi</i> has had a confused +nomenclatural history. The taxon was first named <i>Hyla microcephala</i> +by Boulenger (1898); this name was preoccupied by <i>Hyla microcephala</i> +Cope (1886). Cole and Barbour (1906) and Kellogg (1932) used the name +<i>Hyla phlebodes</i> Stejneger (1906) for specimens of this frog from +México. Dunn (1931, 1933, 1934) applied the name <i>Hyla underwoodi</i> to +Panamanian specimens that we identify as <i>Hyla phlebodes</i>. Smith +(1951) named <i>Hyla microcephala martini</i> from southern México and +Guatemala and considered the northern populations to represent a +subspecies distinct from the Costa Rican <i>Hyla microcephala +underwoodi</i>, despite the fact the Stuart (1935:39) stated that +comparisons of specimens from El Petén, Guatemala, with the holotype +of <i>Hyla underwoodi</i> showed only trivial differences.</p> + +<p>Much of the confusion regarding the name <i>Hyla underwoodi</i> stems from +the illustration given by Boulenger (1898:pl. 39, fig. 3) and +reproduced by Taylor (1952:892), which shows a frog having a unicolor +dorsum, dorsolateral white lines, and dark flanks. This pattern is in +marked contrast to the pattern seen in most preserved specimens, which +have the dorsum variously marked by dark brown lines or irregular +marks. Smith (1951:185), in his description of <i>Hyla microcephala +martini</i> from southern México, considered <i>H. underwoodi</i> to be a +subspecies of <i>H. microcephala</i> that lacked dorsal dark markings.</p> + +<p>Data accumulated in 1961 through field studies by the senior +author at the type locality, Bebedero, and other localities in Guanacaste +and Puntarenas provinces in Costa Rica provide a reasonable +explanation of the differences in color pattern. As noted in the +preceding description of this subspecies, at night the dorsal markings +<span class="pagenum"><a name="Page_532" id="Page_532">[Pg 532]</a></span> +are not evident in many living individuals, whereas by day +the dorsal markings are prominent. Most collectors prepare their +specimens by day; consequently the majority of specimens have a +pronounced dorsal pattern. Of the frogs collected in Costa Rica +in 1961, some specimens were preserved at night; others from the +same series were preserved by day. The differences are striking. +In those preserved at night, dorsal markings are faint, if present at +all. Some specimens closely match the figure given by Boulenger +(1898).</p> + +<p>It is extremely doubtful if the frog described and illustrated by +Boulenger could be associated with either <i>Hyla phlebodes</i> or <i>H. +microcephala microcephala</i>. Individuals of the former species lack +a dorsolateral white line and always have some dorsal markings +evident at night; furthermore, <i>H. phlebodes</i> is not known to occur +on the Pacific lowlands. <i>Hyla microcephala microcephala</i> occurs +farther southeast. Since there is no reason to doubt the type locality +of <i>H. underwoodi</i>, since specimens from the area around the type +locality that have been preserved at night are like the holotype in +pattern, and since the characteristics of the populations of the frogs +in Guanacaste are the same as, or gradually blend into those of, +populations in northern Central America and southern México, the +frogs from throughout the entire range can be referred to one taxon, +the earliest name for which is <i>Hyla underwoodi</i> Boulenger, which +herein is considered to be a subspecies of <i>H. microcephala</i> Cope.</p> +</div> + +<p><i>Distribution.</i>—<i>Hyla microcephala underwoodi</i> inhabits the +Atlantic slopes and lowlands from southern Veracruz and extreme +northern Oaxaca eastward across the base of the Yucatan Peninsula +(possibly the species is extant in the northern part of the +peninsula) to British Honduras and thence southeastward through +the Caribbean lowlands and interior valleys in Honduras to central +Nicaragua, where it apparently avoids the forested Caribbean +lowlands and the dry Pacific lowlands of northwestern Nicaragua, +but in the vicinity of Managua invades the Pacific lowlands and +continues southward into northwestern Costa Rica as far as the +Puntarenas Peninsula (<a href="#Fig_1">Fig. 1</a>). In México and Guatemala the species +has not been taken at elevations of more than 350 meters, whereas +farther south it occurs at higher elevations—780 meters at +Silencio, Costa Rica, 830 meters on Montaña de Guaimaca, Honduras, +960 meters at Finca Tepeyac, Nicaragua, and 1200 meters at Finca +Venecia, Nicaragua.</p> + +<p><i>Specimens examined.</i>—1270, as follows: <b>Mexico</b>: +<span class="smcap">Campeche</span>: Balchacaj, FMNH 100406, UIMNH 20944-6; +Encarnación, FMNH 27069-70, 75784, MCZ 28360, 29637, UIMNH +20948-58, 20965, USNM 134264-5; Escárcega, UMMZ 122999; *7.5 km. W +Escárcega, KU 71229-43; Laguna Alvarado, 65 km. S Xpujil, KU +75084-9; Pacaitún, Río Candelaria, FMNH 83118-20; *Tres Brazos, +FMNH 113101-22, UIMNH 20947; 10 km. W Xpujil, KU 75082-3. <span class="smcap">Chiapas</span>: +Palenque, UIMNH 47984, 49139-50, USNM 114973-8. <span class="smcap">Oaxaca</span>: *5 km. N +Chiltepec, KU 87015-23; 3 km. N Donají UMMZ 115249 (9); *3.7 km. N +Donají, UMMZ 115250 (5); *43 km. N Matías Romero, UIMNH 42550-68; +*3.5 km. N Palomares, TNHC 25185, 25321-31, 25341-68; 4.6 km. +<span class="pagenum"><a name="Page_533" id="Page_533">[Pg 533]</a></span> +N Sarabia, UMMZ 115247 (2); *6.1 km. N Sarabia, UMMZ 115248 (11), *3 +km. N Tolocita, KU 39655; Tuxtepec, KU 87024-40. <span class="smcap">Tabasco</span>: 24 km. N +Frontera, MCZ 35665-70; 0.8 km. E Río Tonolá, TNHC 25189; Teapa, UMMZ +119218 (4); *2.7 km. N Teapa, UMMZ 119216 (4); *10 km. N Teapa, UMMZ +119217 (6); *11.5 km. N Teapa, UMMZ 119219; *15.2 km. N Teapa, UMMZ +119220 (4); *17.6 km. N Teapa, UMMZ 119221 (12), 3.3 km. S +Villahermosa, UMMZ 119215 (12), *17.6 km. S Villahermosa, UMMZ 119214 +(12). <span class="smcap">Veracruz</span>: 2.1 km. N Acayucan, UIMNH 42547-9; *6.4 km. NW +Acayucan, UMMZ 115254 (14); 1.6 km. ESE Alvarado, UMMZ 115258 (39); +*2.4 km. ESE Alvarado, UMMZ 115251 (2); *4.5 km. S Aquilera, +<a name="UMMZ"></a><a href="#typos">UMMZ</a>115252 (21); *8 km. SW Coatzacoalcos, UMMZ 119213 (10); +2.2 km. E Cosoleacaque, UMMZ 119222 (26); 10 km. SE Hueyapan, UMMZ +115255; 0.8 km. S Lerdo de Tejada, UMMZ 122778; *3.6 km. NE +Minatítlán, TNHC 25150-2; 1.9 km. S Naranja, UMMZ 115253 (3); 4.5 km. +NE Novillero, UMMZ 115256; San Andrés Tuxtla, FMNH 113124-8, UIMNH +20942-3. <span class="smcap">Yucatán</span>: Chichén-Itzá, FMNH 36570, MCZ 2463 (2).</p> + +<p><b>British Honduras</b>: <span class="smcap">Cayo</span>: 6.2 km. S El Cayo, MCZ 37885-92. <span class="smcap">Stann Creek</span>: +Stann Creek, FMNH 49068.</p> + +<p><b>Guatemala</b>: <span class="smcap">Alta Verapaz</span>: 28.3 km. N Campur, KU 64578-90; Chinajá, KU +57425; Cubilquitz, UMMZ 90887, 90888 (4); Finca Chamá, UMMZ 90879 +(13), 90880 (4), 90881, 90882 (28), 90883 (12), 90884 (46), 90885 +(39), 90886 (20); *Finca Tinaja, BYU 16032; Panzós, UMMZ 90889 (2). +<span class="smcap">Chiquimula</span>: Chiquimula, UMMZ 98113; 2 km. N Esquipulas, UMMZ 106844. +<span class="smcap">El petén</span>: La Libertad, KU 57447-97, 59907-11 (skeletons), MCZ 21461, +UMMZ 75332 (13), 75333 (11), 75334 (14), 75335 (10); Piedras Negras, +FMNH 113123, UIMNH 20966; *5 km. S Piedras Negras, USNM 114951-72; +Tikal, UMMZ 117981 (2); Toocog, 15 km. SE La Libertad, KU 57426-46. <span class="smcap">El +Quiché</span>: Finca Tesoro, UMMZ 89165 (5). <span class="smcap">Huehuetenango</span>: Finca San Rafael, +16 km. SE Barillas, FMNH 40917-9. <span class="smcap">Izabal</span>: Puerto Barrios, FMNH +20004-7; 8 km. S Puerto Barrios, KU 57507-37, 59991 (eggs), 59992 +(tadpoles); Quirigua, CAS 69657-701; 2.5 km. NE Río Blanco, KU 57539; +San Felípe, FMNH 35065. <span class="smcap">Zacapa</span>: 14 km. ENE Mayuelas, KU 57502-6; 8 km. +ENE Río Hondo, KU 57498-501.</p> + +<p><b>Honduras</b>: <span class="smcap">Atlantidad</span>: La Ceiba, UMMZ 91948 (2), USNM 117593-600; +Lancetilla, MCZ 17981. <span class="smcap">Cortes</span>: Lago Yojoa, AMNH 54917-9, 54957, 55134, +KU 64563-77. <span class="smcap">El Paraiso</span>: Valle de Jamastran, AMNH 54807-12. +<span class="smcap">Francisco-Moranza</span>: El Zamorano, AMNH 54873-81, KU 103223, UMMZ 123101; +Montaña de Guaimaca, AMNH 54900-4 (8); Ranch San Diego, 19 km. SW +Guaimaca, AMNH 53939. <span class="smcap">Itibucá</span>: Vieja Itibucá, AMNH 54912-3.</p> + +<p><b>Nicaragua</b>: <span class="smcap">Chontales</span>: 3 km. SW Santo Tomás, KU 64770-9, 68308 +(skeleton). <span class="smcap">Esteli</span>: Finca Venecia, 7 km. N, 16 km. E Condega, KU +85296; 2.4 km. N Estelí, MCZ 28933-7. MANAGUA: 12-13 km. E +Managua, KU 85297-301; *10 km. SW Tipitapa, UMMZ 119977 (10). +<span class="smcap">Matagalpa</span>: *Finca Tepeyac, 10.5 km. N, 9 km. E Matagalpa, KU 85302-3; +Hacienda La Cumplida, KU 64780-96, 68309-11 (skeletons), UMMZ 116482 +(8), 116483 (23), 116484 (3), 116485 (5), 119984 (3). <span class="smcap">Rivas</span>: *Finca +Amayo, 13 km. S, 14 km. E Rivas, KU 85304-7; 16 km. S Rivas, MCZ +29011-7; *20.5 km. SE Rivas, KU 85308-10; 5 km. SE San Pablo, KU +43111-4.</p> + +<p><b>Costa Rica</b>: <span class="smcap">Guanacaste</span>: Arenal, USC 6254 (2); *3 km. W Bagaces, USC +7019 (10); *3 km. NE Boca del Barranca, USC 8017 (21), *Finca San +Bosco, USC 6272 (6), 6276 (3); *Guayabo de Bagaces, USC 7022 (4), 7023 +(3), 7025; 12 km. S La Cruz, USC 8091 (2); *Laguna Arenal, USC 6262; +*27 km. N Las Cañas, USC 8171 (6); *16 km. E Las Cañas, KU 102252-8; +16 km. SSE Las Cañas, KU 65090-5; *20 km. SE Las Cañas, KU 102251; +Liberia, KU 30827-39; *7.3 km. N Liberia, USC 8096 (4); *10 km. N +Liberia, USC 8085 (9); *7.5 km. SE Liberia, KU 65102-8, 68621-2 +(skeletons); *14.7 km. S Liberia, USC 8238 (3); *4 km. W Liberia, KU +36847-57; 2 km. S Nicoya, USC 8230; *3-10 km. ESE Playa del Coco, USC +8012 (16), 8137 (14); *21.6 km. ESE Playa del Coco, USC 8138 (13); +*Peñas Blancas, KU 102247-50; *Río Bebedero, 5 km. S Bebedero, KU +65089; *Río Higuerón, USC 7168 (2); Santa Cruz, USC 8232 (2); +*Silencio, USC 6248; *Tenorio, KU 32313; Tilarán, +<span class="pagenum"><a name="Page_534" id="Page_534">[Pg 534]</a></span> +KU 36858-60; *2 km. E Tilarán, KU 86403, *5 km. NE Tilarán, KU +36840-6 USC 6269. <span class="smcap">Puntarenas</span>: Barranca, KU 32305-12, *5 km. WNW +Barranca, UMMZ 119976 (2); *10 km. E Esparta, KU 86400-2; 1 km. +WNW Esparta, KU 65101; *4 km. WNW Esparta, KU 65088; *10 km. WNW +Esparta, KU 65063-87, 68616-20 (skeletons); *12 km. WNW Esparta, +KU 65096-100, USC 8251; 21.8 km. W San Ramón, USC 8242 (15).</p> +<p> </p> +<p> </p> + +<div class="caption3nb"><b>Hyla robertmertensi</b> Taylor</div> + +<div class="species"> +<p><i>Hyla robertmertensi</i> Taylor, Proc. Biol. Soc. Washington, 50:43, April 21, +1937 [Holotype.—CNHM 100096 (formerly EHT-HMS 2270) from +Tapachula, Chiapas, México; H. M. Smith and E. H. Taylor collectors]. +Smith and Taylor, Bull. U. S. Natl. Mus., 194:84, June 17, 1948; Univ. +Kansas Sci. Bull., 33:326, March 20, 1950. Mertens. Senckenbergiana, +33:170, June 15, 1952; Senckenbergischen Naturf. Gesell., 487:30, December +1, 1952. Stuart, Contr. Lab. Vert. Biol., Univ. Michigan, 68:47, +November, 1954. Duellman, Univ. Kansas Publ., Mus. Nat. Hist., 13:63, +August 16, 1960. Duellman and Hoyt, Copeia, 1961 (2): 417, December +19, 1961. Porter, Herpetologica, 18:168, October 17, 1962. Stuart, +Misc. Publ. Mus. Zool., Univ. Michigan, 122:36, April 2, 1963. Duellman +and Trueb, Univ. Kansas Publ., Mus. Nat. Hist., 17:348, July 14, 1966.</p> +</div> + +<p><a name="Diagnosis"></a><a href="#typos"><i>Diagnosis.</i></a>—Brown lateral stripe wide, including loreal region and entire +tympanum, extending to groin, bordered above by narrow white line; dorsum +unicolor or with pair of dark lines (or rows of dashes) usually extending only +to the sacral region; shanks having dark flecks, no transverse bars; interorbital +bar lacking.</p> + +<p><i>Description and Variation.</i>—Males attain a maximum snout-vent length of +26.4 mm. in Oaxaca, whereas in a sample from Acacoyagua, Chiapas, the +largest male has a snout-vent length of 25.7 mm., and from La Trinidad, +Guatemala, 24-6 mm. Specimens from the western part of the range (eastern +Oaxaca) have slightly smaller heads and proportionately larger tympani than +the more eastern populations (<a href="#Table_1">Table 1</a>).</p> + +<p>The color pattern shows little variation, except in the nature of the dorsal +markings. In a few specimens from throughout the range, but especially in +the eastern part of the range, the dorsum lacks markings between the dorsolateral +white lines. In most specimens the dorsal pattern consists of flecks or dashes +arranged in two parallel longitudinal rows, and in some specimens the marks +are fused into parallel lines. Small brown flecks are present on the dorsal surfaces +of the shanks; in some specimens these flecks tend to form a longitudinal +stripe on the shank. An interorbital dark mark is invariably absent.</p> + +<p>When active at night <i>Hyla robertmertensi</i> is pale yellow above with a white +dorsolateral line and pale brown lateral stripe; the dorsal markings are faint. +By day the dorsum is yellowish tan with brown markings. The dorsolateral +stripe is creamy white, and the lateral stripe is dark brown (<a href="#Pl_14">Pl. 14</a>). The +venter is white, and the iris is dull bronze. In breeding males the vocal sac +is yellow.</p> + +<div class="remarks"> +<p><i>Remarks.</i>—Although this species superficially resembles <i>Hyla +microcephala microcephala</i>, the latter is easily distinguished by the +narrow brown lateral stripe, as compared with the much wider stripe +in <i>H. robertmertensi</i>. No other hylids in northern Central America +and southern México can be confused with this species.</p> +</div> + +<p><i>Distribution.</i>—<i>Hyla robertmertensi</i> inhabits the Pacific slopes (to elevations +of 700 meters) and lowlands from eastern Oaxaca (east of the Plains of Tehuantepec) +<span class="pagenum"><a name="Page_535" id="Page_535">[Pg 535]</a></span> +southeastward to central El Salvador. The species also occurs in the +Cintalapa Valley (Atlantic drainage) in southwestern Chiapas (<a href="#Fig_2">Fig. 2.</a>) The +distribution seems to be limited on the northwest and southeast by arid environments. +The region in which <i>Hyla robertmertensi</i> lives is characterized by +higher rainfall and more luxuriant vegetation than occur on the Plains of +Tehuantepec or on the Pacific lowlands of eastern El Salvador and southern +Honduras. In addition to the localities listed below, Mertens (1952:30) +recorded the species from Hacienda Cuyan-Cuya, Depto. Sonsonate, El +Salvador.</p> +</div> +<p> </p> +<p> </p> + +<a name="Fig_2"></a> +<div class="center"> +<img src="images/fig_2.png" width="600" height="490" title="Map Locality - Hyla robertmertensi" alt="Map Locality - Hyla robertmertensi"><br><br> +<span class="smcap">Fig. 2.</span> Map showing locality records for <i>Hyla robertmertensi</i>.<br> +</div> +<p> </p> +<p> </p> + +<p><i>Specimens examined.</i>—490, as follows: <b>Mexico</b>: <span class="smcap">Chiapas</span>: Acacoyagua, +USNM 114754-61; *2 km. W Acacoyagua, UMMZ 87843 (28), 87844 (50), +87845 (50), 87846 (45), 87847 (27), 87848 (3); 32 km. N Arriaga, KU +57619-24, 59917-8 (skeletons); Asunción, FMNH 100413, 100501-4, UIMNH +26989-90, USNM 134267; *La Esperanza, USNM 114737-48, 114750-3, 17 km. +S Las Cruces, KU 57625-49, 59997 (eggs); 8.5 km. N Puerto Madero, UMMZ +119981 (2); *11.7 km. N Puerto Madero, UMMZ 119982; Tapachula, FMNH +100096, UIMNH 26987; *11 km. S Tapachula, KU 57605-18, 59916 (skeleton); +Tonolá, FMNH 27073, 100505-10, UIMNH 26988. <span class="smcap">Oaxaca</span>: Tapanatepec, +UMMZ 115245 (2), *1.6 km. E Tapanatepec, UMMZ 115244 (14); *4.3 km. +E Tapanatepec, UIMNH 38368-9; *7.5 km. W Tapanatepec, UMMZ 115246 +(39); 12.8 km. W Tapanatepec, KU 65007-14; 7.2 km. WNW Zanatepec, +UMMZ 115243 (77); *13.6 km. WNW Zanatepec, TNHC 25213-22; 22.7 km. +WNW Zanatepec, TNHC 25203-9.</p> + +<p><b>Guatemala</b>: <span class="smcap">Jutiapa</span>: Jutiapa, UMMZ 106848; La Trinidad, UMMZ +107733 (23). <span class="smcap">Retalhueleu</span>: Casa Blanca, UMMZ 107732.</p> + +<p><b>El Salvador</b>: <span class="smcap">La Libertad</span>: 16 km. NW Santa Tecla, KU 44112. <span class="smcap">San +Salvador</span>: 21.9 km. N San Salvador, UMMZ 119983 (6).</p> + +<p><span class="pagenum"><a name="Page_536" id="Page_536">[Pg 536]</a></span></p> + +<div class="caption3nb"><b>Hyla phlebodes</b> Stejneger</div> + +<div class="species"> +<p><i>Hyla phlebodes</i> Stejneger, Proc. U. S. Natl. Mus., 30:817, June 4, 1906 +[Holotype.—USNM 2997 from "San Carlos," Costa Rica; Burgdorf and +Schild collectors]. Taylor, Proc. Biol. Soc. Washington, 50:44, April 21, +1937; Univ. Kansas Sci. Bull., 35:888, July 1, 1952; Univ. Kansas Sci. +Bull., 39:25, November 18, 1958. Fouquette, Evolution, 14:484, December +16, 1960. Duellman and Trueb, Univ. Kansas Publ., Mus. Nat. +Hist., 17:348, July 14, 1966.</p> + +<p><i>Hyla underwoodi</i>, Dunn, Occas. Papers Boston Soc. Nat. Hist., 5:413, +October 10, 1931; Occas. Papers Boston Soc. Nat. Hist. 8:72, June 7, 1933; +Amer. Mus. Novitiates, 747.2, September 17, 1934, Gaige, Hartweg, and +Stuart, Occas. Papers Mus. Zool., Univ. Michigan, 357:5, October 26, +1937. Breder, 1946, Bull. Amer. Mus. Nat. Hist., 86:416, August 22, +1946.</p> +</div> + +<p><i>Diagnosis.</i>—Dark brown lateral stripe, if present, usually extending only to +insertion of forearm, never posteriorly to sacral region; white line above brown +stripe absent or faint; dorsal pattern weak, usually consisting of irregular dashes +or interconnected lines; interorbital dark mark present; shanks having weakly +defined transverse bars.</p> + +<p><i>Description and variation.</i>—In the majority of specimens (70%) the lateral +dark stripe extends from the nostril to the eye and thence above the tympanum +to a point above the insertion of the arm; in 17 per cent the stripe extends to +the mid-flank, whereas in 13 per cent the stripe is absent. A narrow and faint +white line is present on the canthus in some specimens, but no distinct white +stripe is present above the lateral dark line posterior to the eye. An interorbital +bar and transverse marks on the shanks are invariably present. The +dorsal markings are variable, but in most specimens (92%) consist of either an +X- or )(-shaped mark in the scapular region; in the other specimens the markings +are irregular short lines or absent. Approximately equal numbers of +specimens have a transverse bar, chevron, or broken lines in the sacral region, +whereas about eight per cent of the specimens lack markings in the sacral +region.</p> + +<p>When active at night, individuals are pale yellowish tan with faint brown +dorsal markings. By day they are tan with more distinct brown markings +(<a href="#Pl_14">Pl. 14</a>). The thighs are pale yellow; the belly is white. The iris is pale creamy +tan with brown flecks. In breeding males the vocal sac is yellow.</p> + +<p><i>Tadpoles.</i>—Tadpoles of this species have been found in an extensive grassy +pond at Puerto Viejo, Costa Rica. The following description is based on KU +104099, a specimen in development stage 36 (Gosner, 1960).</p> + +<p>Total length, 21.0 mm.; body length, 6.7 mm.; body slightly wider than +deep, snout pointed; nostrils large, directed anteriorly, situated near end of +snout; eyes small, situated dorsolaterally, directed laterally; spiracle sinistral, +located just posteroventral to eye; anal tube dextral. Tail xiphicercal; caudal +musculature moderately deep, extending far beyond posterior edge of fins; fins +deepest at about midlength; dorsal fin extending onto body, slightly deeper than +caudal musculature; ventral fin slightly shallower than musculature. Mouth +small, terminal, lacking teeth and fringing papillae, but having finely serrate +beaks. In preservative top of head olive-tan with brown flecks; dark stripe +from snout through eye to posterior edge of body; belly white, flecked with +brown anteriorly; tail creamy tan with grayish brown blotches. In life, dorsum +of body reddish tan mottled with darker brown; lateral stripe dark brown; belly +white, mottled with brown and black; caudal musculature heavily pigmented +<span class="pagenum"><a name="Page_537" id="Page_537">[Pg 537]</a></span> +with grayish tan; posterior tip of tail marked with dark gray; caudal fins heavily +blotched with grayish tan; iris orange-tan peripherally, red centrally (<a href="#Pl_15">Pl. 15</a>).</p> + +<div class="remarks"> +<p><i>Remarks.</i>—This species has been confused with <i>Hyla microcephala +underwoodi</i> by many workers. Dunn (1931, 1933, 1934) +and Breder (1946) referred Panamanian specimens of <i>H. phlebodes</i> +to <i>H. underwoodi</i>; likewise, Gaige, Hartweg, and Stuart (1937) +made the same error. Cole and Barbour (1906) and Kellog (1932) +used the name <i>H. phlebodes</i> for Mexican specimens of <i>H. microcephala +underwoodi</i>. The similarity in color pattern of <i>H. microcephala +underwoodi</i> and <i>H. phlebodes</i> easily accounts for the misapplication +of names. Although both species have nearly identical +dorsal color patterns, that of <i>H. microcephala underwoodi</i> usually +is bolder. Furthermore, in that species a narrow white line usually +is present above the well-defined lateral dark stripe, whereas the +lateral dark stripe is short and posterior to the eye is not bordered +above by a white line in <i>H. phlebodes</i>.</p> + +<p>The type locality "San Carlos, Costa Rica" given by Stejneger +(1906:817) apparently refers to a region, the Llanuras de San Carlos, +in the northern part of Alajuela Province, Costa Rica.</p> +</div> +<p> </p> +<p> </p> + +<a name="Fig_3"></a> +<div class="center"> +<img src="images/fig_3.png" width="600" height="433" title="Map Locality - Hyla phlebodes" alt="Map Locality - Hyla phlebodes"><br><br> +<span class="smcap">Fig. 3.</span> Map showing locality records for <i>Hyla phlebodes</i>.<br> +</div> +<p> </p> +<p> </p> + +<div class="blockquot"> +<p><i>Distribution.</i>—<i>Hyla phlebodes</i> inhabits humid tropical forests from southeastern +Nicaragua southeastward on the Caribbean slopes and lowlands to the +Canal Zone in Panamá, thence eastward in the Chucunaque Basin of eastern +Panamá and onto the Pacific lowlands of Colombia (<a href="#Fig_3">Fig. 3</a>). The species also +<span class="pagenum"><a name="Page_538" id="Page_538">[Pg 538]</a></span> +reaches the Pacific slopes in the Arenal Depression in northwestern Costa Rica +and in the Panamanian isthmus, where it occurs in humid forests on the Pacific +slope of El Valle and Cerro La Campana. Mostly the species is found at low +elevations, but it occurs at 600 meters at Turrialba and at 700 meters at Finca +San Bosco in Costa Rica.</p> + +<p><i>Specimens examined.</i>—410, as follows: <b>Nicaragua</b>: <span class="smcap">Zelaya</span>: Isla Grande +del Maíz, MCZ 14848; Río Mico, El Recrero, UMMZ 79720 (6).</p> + +<p><b>Costa Rica</b>: <span class="smcap">Alajuela</span>: 12.4 km. N Florencia, MVZ 76108-10, USC 2628; +*Las Playuelas, 11 km. S Los Chiles, USC 7216; Los Chiles, USC 7217, 7219; +3 km. NE Muelle de Arenal, USC 2644 (2); *"San Carlos," USNM 29970. +<span class="smcap">Cartago</span>: Chitaría, KU 103690; *1.6 km. E Río Reventazón Bridge, east of +Turrialba, UMMZ 119978 (2); *Tunnel Camp, near Peralta, KU 32456, +32458-69, 41098 (skeleton); Turrialba, FMNH 101794, 103188-9, KU 25725-9, +32439-48, 41095-7 (skeletons), 64797-827, 68300-2 (skeletons), 68403 (eggs), +68404 (tadpoles), MCZ 29224-5, 29310-2, UMMZ 119979 (6), USC 31, 256 +(2), 458 (2), 580, 594, 599 (7), 7074 (2), USNM 29933. <span class="smcap">Guanacaste</span>: +Arenal, USC 6254; *Finca San Bosco, USC 62724, 6276 (3), Guayabo de +Bagaces, USC 7022 (3), 7023; *Laguna Arenal, USC 6262 (4); 3 km. NE +Tilarán, USC 524; *5 km. NE Tilarán, USC 6269; *6 km. NE Tilarán, UMMZ +122653 (6), S-2680 (skeleton), USC 523 (8). <span class="smcap">Heredia</span>: Puerto Viejo, KU +64828-63, 68303-7 (skeletons), 68405-6 (tadpoles), 104099-100 (tadpoles); +*1.5 km. N Puerto Viejo, KU 64871; *1 km. S Puerto Viejo, KU 86432-40; +*4.2 km. W Puerto Viejo, KU 64864-5; *5.9 km. W Puerto Viejo, KU 64866-70; +*7.5 km. W Puerto Viejo, KU 86431. <span class="smcap">Limón</span>: Batán, UMMZ 119980 (2); La +Castilla, ANSP 23707; Puerto <b>Limón</b>, KU 32449-55.</p> + +<p><b>Panama</b>: <span class="smcap">Bocas del Toro</span>: 3.2 km. NW Almirante, KU 96026; Cayo de +Agua, KU 96027-31; Fish Creek, KU 96032-4. <span class="smcap">Canal Zone</span>: Barro Colorado +Island, AMNH 69790, ANSP 23244-50; FMNH 13380, 22972-4; Juan Mina, +AMNH 55429, UU 3899; *8.6-13.8 km. N Miraflores Locks, TNHC 23439, +23477, 23484-8, 23491, 23494-9, 23501-2, 23504-8, 23510-17, 23519-30, +23532-8, 23541-54, 23561. *Rio Chagres, AMNH 55431-4; Río Cocolí, 3.5 km. +N Miraflores Locks, TNHC 23461, 23489-90, 23493, 23500, 23503, 23509, +23518, 23531, 23539-40; *Summit, ANSP 23361, KU 97788; *Three Rivers +Plantation, SU 2130. <span class="smcap">Coclé</span>: El Valle de Antón, AMNH 55435, 69786-9, +ANSP 23506-9. <span class="smcap">Colón</span>: Achiote, KU 77215-78; Ciricito, CAS 71499-500, +71505-6. <span class="smcap">Darién</span>: Río Canclon at Río Chucunaque, UMMZ 126733; Río +Chucunaque, near Yavisa, AMNH 51783. <span class="smcap">Panamá</span>: Cero La Campana, FMNH +67847-50.</p> + +<p><b>Colombia</b>: <span class="smcap">Chocó</span>: Andagoya, FMNH 81856; Boca de Raspadura, AMNH +13570-8.</p> +</div> +<p> </p> +<p> </p> + +<div class="caption3nb"><b>Hyla sartori</b> Smith</div> + +<div class="species"><p> +<i>Hyla underwoodi</i> (in part), Smith and Taylor, Bull. U. S. Natl. Mus., 194:85, +June 17, 1948.</p> + +<p><i>Hyla microcephala sartori</i> Smith, Herpetologica, 7:186, December 31, 1951 +[Holotype.—UIMNH 20934 from 1 mile north of Organos, south of El +Treinte, Guerrero, México; H. M. Smith and E. H. Taylor collectors]. +Duellman, Univ. Kansas Publ., Mus. Nat. Hist., 15:124, December 20, +1961. Porter, Herpetologica, 18:168, October 17, 1962. Davis and Dixon, +Herpetologica, 20:230, January 25, 1965. Duellman, Univ. Kansas Publ. +Mus. Nat. Hist., 15:652, December 30, 1965.</p></div> + +<p><i>Diagnosis.</i>—Dorsum tan with broad dark brown chevrons or transverse +bars; shanks marked with two or three broad transverse bars; dorsolateral +stripes absent.</p> + +<p><i>Description and variation.</i>—No noticeable geographic variation is apparent +in either structural features or coloration in this species. All specimens lack +a dorsolateral dark stripe and white line, although a dark line is present on the +<span class="pagenum"><a name="Page_539" id="Page_539">[Pg 539]</a></span> +canthus and dissipates in the loreal region. A broad interorbital brown bar is +present in all specimens. The color pattern on the dorsum invariably consists +of a broad, dark, chevron-shaped mark in the scapular region and a broad +dark chevron or transverse bar in the sacral region. The shanks invariably +have two or three dark brown transverse bars.</p> + +<p>When active at night individuals are yellowish tan above with chocolate +brown markings (<a href="#Pl_14">Pl. 14</a>). The belly is white, and the thighs are pale yellowish +tan. The iris is dark bronze-color. In breeding males the vocal sac is yellow. +By day some individuals were observed to change to creamy gray with distinct +darker markings.</p> + +<div class="remarks"> +<p><i>Remarks.</i>—Although tadpoles of this species have not been found, +observations on the breeding sites indicate that the tadpoles probably +develop in ponds. Except for calling males observed around a +pool in a stream-bed 11.8 kilometers west-northwest of Tierra Colorada, +Guerrero, all breeding congregations have been found at +temporary ponds.</p> + +<p>Smith (1951:186) named <i>Hyla sartori</i> as a subspecies of <i>Hyla +microcephala</i>. This subspecific relationship seemed reasonable until +analysis of the mating calls showed that the call of <i>H. sartori</i> is more +nearly like that of <i>H. phlebodes</i> than that of <i>H. microcephala</i>. The +broad hiatus separating the ranges of <i>H. microcephala</i> and <i>H. sartori</i> +is additional evidence for considering <i>H. sartori</i> as a distinct species.</p> +</div> +<p> </p> +<p> </p> + +<a name="Fig_4"></a> +<div class="center"> +<img src="images/fig_4.png" width="600" height="297" title="Map Locality - Hyla sartori" alt="Map Locality - Hyla sartori"><br><br> +<span class="smcap">Fig. 4.</span> Map showing locality records for <i>Hyla sartori</i>.<br> +</div> +<p> </p> +<p> </p> + +<p><i>Distribution.</i>—<i>Hyla sartori</i> occurs in mesophytic forests to elevations of +about 300 meters on the Pacific slopes of southern México from southwestern +Jalisco to south-central Oaxaca (<a href="#Fig_4">Fig. 4</a>). The lack of specimens from Colima +and Michoacán probably reflects inadequate collecting instead of the absence +of the species there. On the basis of available habitat the species would be +expected to occur in Nayarit, but extensive collecting there has failed to +reveal its presence. The semi-arid Plains of Tehuantepec apparently limit the +distribution to the east.</p> + +<p><span class="pagenum"><a name="Page_540" id="Page_540">[Pg 540]</a></span> +<i>Specimens examined.</i>—190, as follows: <b>México</b>: <span class="smcap">Guerrero</span>: 5 km. E Acapulco, +AMNH 54611-2; 23.2 km. N Acapulco, UIMNH 26404-7; Colonia Buenas +Aires, 23 km. E Tecpán de Galeana, UMMZ 119223 (7); *El Limoncito, +FMNH 75785, 100390-402, 104631, 104633, UMMZ 117250, USNM 134266; +El Treinte, FMNH 100403, UIMNH 20935-7; Laguna Coyuca, AMNH +59686; La Venta, MCZ 29635; *Morjonares, UIMNH 26392-402; 1.6 km. +N Organos, FMNH 100404-5, UIMNH 20933-4; 19.2 km. S Petaquillas, +UIMNH 26408; 6.1 km. E. Tecpán de Galeana, TNHC 23396-408; *11.2 km. +N Tierra Colorada, UIMNH 26403; 11.8 km. WNW Tierra Colorada, UMMZ +119225 (51), S-2677-9 (skeletons); Zacualpán, UMMZ 119224 (6). <span class="smcap">Jalisco</span>: +6.4 km. NE La Resolana, KU 67853-69; 24 km NE La Resolana, KU 67870-3. +<span class="smcap">Oaxaca</span>: 3 km. N Pochutla, KU 57539; 13.4 km. N Pochutla, UMMZ +123495 (40).</p> +<p> </p> +<p> </p> + +<a name="CRANIAL_OSTEOLOGY" id="CRANIAL_OSTEOLOGY"></a> +<div class="caption2">CRANIAL OSTEOLOGY</div> + +<p>The frogs of the <i>Hyla microcephala</i> group have a minimal amount +of cranial ossification as compared to more generalized hylid skulls, +such as <i>Smilisca</i> (Duellman and Trueb, 1966). In the <i>Hyla microcephala</i> +group the sphenethmoid is small and short, and a large +frontoparietal fontanelle is present. The quadratojugal exists only +as a small spur and is not in contact with the maxillary. The +proötics are poorly developed. The anterior and posterior arms +of the squamosal are short; the anterior arm extends no more than +one-fourth of the distance to the maxillary, and the posterior arm +does not have a bony connection with the proötic. The nasal lacks +a maxillary process, and the medial ramus of the pterygoid lacks a +bony connection to the proötic.</p> + +<p>Teeth are absent on the parasphenoid and palatines, but present +on the maxillaries, premaxillaries, and prevomers. The teeth are +simple, pointed, and slightly curved. Although the number of teeth +varies (<a href="#Table_3">Table 3</a>), no consistent differences between the species are +apparent.</p> +<p> </p> +<p> </p> + +<a name="Table_3"></a> +<span class="smcap">Table 3.</span>—Variation in the Number of Teeth in the Species of the Hyla Microcephala Group. (N=Number of Jaws, or Twice the Number of Individuals; Means are Given in Parentheses After the Observed Ranges).<br> + +<div class="center"> +<table width="75%" class="data" summary="Teeth Numer Variation Hyla microcephala Group"> +<tr><td colspan=5 class="bb"> </td></tr> +<tr><td class="bt bb"><span class="smcap">Species</span></td><td class="bt bl bb">N</td><td class="bt bl bb">Maxillary</td><td class="bt bl bb">Premaxillary</td><td class="bt bl bb">Prevomer</td></tr> +<tr><td class="text_lf"><i>H. microcephala</i></td><td class="bl">32</td><td class="bl">31-47(37.8)</td><td class="bl">4-13(8.9)</td><td class="bl">2-4(3.2)</td></tr> +<tr><td class="text_lf"><i>H. phlebodes</i></td><td class="bl">10</td><td class="bl">38-45(40.1)</td><td class="bl">8-13(10.3)</td><td class="bl">2-5(3.9)</td></tr> +<tr><td class="text_lf"><i>H. robertmertensi</i></td><td class="bl">6</td><td class="bl">23-43(32.8)</td><td class="bl">7-12(10.5)</td><td class="bl">2-3(2.7)</td></tr> +<tr><td class="text_lf"><i>H. sartori</i></td><td class="bl">6</td><td class="bl">27-43(38.2)</td><td class="bl">9-10(9.3)</td><td class="bl">3-4(3.7)</td></tr> +</table> +</div> +<p> </p> +<p> </p> + +<div class="center"> +<a name="Pl_13"></a> +PLATE 13<br><br> +<img src="images/pl_13.png" width="354" height="600" title="Adult Frogs" alt="Adult Frogs"><br><br> +Upper figure, <i>Hyla microcephala microcephala</i> (KU 64593);<br> +middle figure, <i>H. microcephala underwoodi</i> (KU 64565);<br> +lower figure, <i>H. microcephala underwoodi</i> (UMMZ 115247).<br> +All approximately ×3.<br> +</div> +<p> </p> +<p> </p> + +<div class="center"> +<a name="Pl_14"></a> +PLATE 14<br><br> +<img src="images/pl_14.png" width="352" height="595" title="Adult Frogs" alt="Adult Frogs"><br><br> +Upper figure, <i>Hyla robertmertensi</i> (UMMZ 115243);<br> +middle figure, <i>H. phlebodes</i> (KU 64798);<br> +lower figure, <i>H. sartori</i> (UMMZ 119225).<br> +All approximately ×3.<br> +</div> +<p> </p> +<p> </p> + +<div class="center"> +<a name="Pl_15"></a> +PLATE 15<br><br> +<img src="images/pl_15.png" width="600" height="342" title="Tadpolrs" alt="Tadpolrs"><br><br><br> +Tadpoles of <i>Hyla microcephala</i> group:<br> +upper figure, <i>H. m. microcephala</i> (KU 104097);<br> +lower figure, <i>H. phlebodes</i> (KU 104099).<br> +Both ×4.<br> +</div> +<p> </p> +<p> </p> + +<div class="center"> +<a name="Pl_16"></a> +PLATE 16<br><br> +<img src="images/pl_16.png" width="420" height="593" title="Audiospectrograms" alt="Audiospectrograms"><br><br> +Audiospectrograms and sections of mating calls of <i>Hyla microcephala</i> group:<br> +(a) <i>H. m. microcephala</i> (KU Tape No. 19);<br> +(b) <i>H. robertmertensi</i> (KU Tape No. 41);<br> +(c) <i>H. phlebodes</i> (KU Tape No. 6);<br> +(d) <i>H. sartori</i> (KU Tape No. 190).<br> +</div> +<p> </p> +<p> </p> + +<span class="pagenum"><a name="Page_541" id="Page_541">[Pg 541]</a></span> +<a name="Table_4"></a> +<span class="smcap">Table 4.</span>—Comparative Cranial Osteology of Hyla microcephala Group<br> + +<table width="100%" class="data" summary="Comparative Cranial Osteology of Hyla microcephala Group"> +<tr><td colspan=5 class="bb"> </td></tr> +<tr> +<td class="center bt bb"><span class="smcap">Character</span></td> +<td class="center bt bl bb"><i>H. microcephala</i></td> +<td class="center bt bl bb"><i>H. robertmertensi</i></td> +<td class="center bt bl bb"><i>H. phlebodes</i></td> +<td class="center bt bl bb"><i>H. sartori</i></td> +</tr> +<tr> +<td class="vtop text_lf">Frontoparietal</td> +<td class="vtop bl">Minimally ossified with large fontanelle extending from sphenethmoid to occipital ridge.</td> +<td class="vtop bl">Ossification extensive anteriorly with narrow medial separation; fontanelle largest in parietal region.</td> +<td class="vtop bl">Ossification extensive anteriorly with narrow medial separation; fontanelle largest in parietal region.</td> +<td class="vtop bl">Ossification moderately extensive anteriorly; medial separation of about uniform width throughout length of fontanelle.</td> +</tr> +<tr> +<td class="vtop text_lf">Nasals</td> +<td class="vtop bl">Moderately long and slender; arcuate in dorsal view.</td> +<td class="vtop bl">Moderate in size; slightly wider anteriorly than posteriorly in dorsal view.</td> +<td class="vtop bl">Moderate in size; slightly wider anteriorly than posteriorly in dorsal view.</td> +<td class="vtop bl">Long and broad; arcuate in dorsal view.</td> +</tr> +<tr> +<td class="vtop text_lf">Sphenethmoid</td> +<td class="vtop bl">Extremely short in dorsal view.</td> +<td class="vtop bl">Moderately short in dorsal view.</td> +<td class="vtop bl">Moderately short in dorsal view.</td> +<td class="vtop bl">Moderately short in dorsal view; ossified anteriorly between nasals.</td> +</tr> +<tr> +<td class="vtop text_lf bb">Columella</td> +<td class="vtop bl bb">Distal and greatly expanded.</td> +<td class="vtop bl bb">Distal and slightly expanded or not.</td> +<td class="vtop bl bb">Distal and not expanded.</td> +<td class="vtop bl bb">Distal and not expanded.</td> +</tr> +</table> +<p> </p> +<p> </p> + +<a name="Fig_5"></a> +<p><span class="pagenum"><a name="Page_542" id="Page_542">[Pg 542]</a></span></p> +<div class="center"> +<img src="images/fig_5.png" width="345" height="600" title="Skulls Dorsal View" alt="Skulls Dorsal View"><br><br> +<span class="smcap">Fig. 5.</span>Dorsal views of the skulls of (a) <i>Hyla m. microcephala</i> +(KU 68293) and (b) <i>H. sartori</i>(UMMZ S-2677). Both × 12.<br> +</div> +<p> </p> +<p> </p> + +<a name="Fig_6"></a> +<p><span class="pagenum"><a name="Page_543" id="Page_543">[Pg 543]</a></span></p> +<div class="center"> +<img src="images/fig_6.png" width="323" height="600" title="Skulls Dorsal View" alt="Skulls Dorsal View"><br><br> +<span class="smcap">Fig. 6.</span> Dorsal views of skulls of (a) <i>Hyla phlebodes</i> (KU 68303) +and (b) <i>H. robertmertensi</i> (KU 59917). Both × 12.<br><br> +</div> +<p> </p> +<p> </p> + +<p>Despite the great reduction in the ossification of the cranial +elements, certain apparently consistent differences exist between +<span class="pagenum"><a name="Page_544" id="Page_544">[Pg 544]</a></span> +the species seem to be consistent. The most notable differences +are: 1) amount of ossification of the frontoparietals and consequent +shape and size of the frontoparietal fontanelle, 2) shape of the +nasals, 3) shape and extent of the sphenethmoid, and 4) shape of +the columella (<a href="#Table_4">Table 4</a>, <a href="#Fig_5">Figs. 5-6</a>). On the basis of these characters, +<i>Hyla microcephala</i> can be set apart from the other species and +characterized as having a poorly ossified frontoparietal and correspondingly +large frontoparietal <a name="fontanelle"></a><a href="#typos">fontanelle</a>; long, slender, arcuate +nasals; extremely short sphenethmoid; and expanded distal end of +the columella. The other species in the group (<i>phlebodes</i>, <i>robertmertensi</i>, +and <i>sartori</i>) have more ossification of the frontoparietals, +broader nasals, only a moderately short sphenethmoid, and an unexpanded +distal end of the columella. Among these three species, +the skulls of <i>phlebodes</i> and <i>robertmertensi</i> are most nearly alike, +whereas the skull of <i>sartori</i> differs by having a differently shaped +frontoparietal fontanelle, broader nasals, and an ossified anterior +extension of the sphenethmoid between the nasals (compare <a href="#Fig_5">Fig. 5b</a> with <a href="#Fig_6">Fig. 6 a-b</a>).</p> + +<p>Although all skulls examined belong to breeding adults, the +extent of the ossification of the frontoparietals and the resulting +shape of the frontoparietal fontanelle might be correlated with the +age of the frog. Nevertheless, in the 24 skulls of <i>Hyla microcephala</i> +examined, the frontoparietals are less extensively ossified than in +the skulls of the other species. The trivial differences among the +other three species certainly are suggestive of close relationship, +but on the basis of present knowledge of the evolutionary trends +in hylid cranial osteology, the differences offer little evidence for +determining phylogenetic lineage.</p> +<p> </p> +<p> </p> + +<a name="ANALYSIS_OF_MATING_CALLS" id="ANALYSIS_OF_MATING_CALLS"></a> +<div class="caption2">ANALYSIS OF MATING CALLS</div> + +<p>Calls of all five taxa were compared in several characteristics, of +which three are deemed most significant systematically. These +are 1) the pattern and duration of the notes of a call-group, 2) the +fundamental frequency, and 3) the dominant frequency. Air temperatures +were noted at the time the calls were recorded, but no +valid correlation could be determined between this factor and any +of the parameters of the calls; consequently recordings made at all +temperatures (21-29° C.) were grouped together.</p> + +<p><i>Pattern and duration of notes.</i>—In all five taxa the basic pattern +consists of a call-group made up of one primary note followed by +a series of shorter secondary notes. In some species the secondary +<span class="pagenum"><a name="Page_545" id="Page_545">[Pg 545]</a></span> +notes differ from the primary in other characteristics. Both subspecies +of <i>Hyla microcephala</i> have a long, unpaired primary note +followed by 0 to 18 (usually about 4) somewhat shorter paired +secondary notes. In calls of <i>Hyla m. microcephala</i> the mean duration +of the primary is 0.131 (0.10-0.16) second and that of the +secondaries is 0.101 (0.05-0.14) second, whereas in <i>H. m. underwoodi</i> +the mean duration of the primary is 0.018 (0.05-0.15) second +and that of the secondaries is 0.086 (0.06-0.11) second.</p> + +<p><i>Hyla robertmertensi</i> has a reverse of this pattern in that the +primary note is paired and the secondaries are unpaired. In the +sample studied a call-group contains 0-28 secondary notes (generally +about 3). The mean duration of the primary is 0.091 (0.07-0.11) +second and that of the secondaries is 0.040 (0.025-0.06) second.</p> + +<p><i>Hyla phlebodes</i> and <i>sartori</i> have call-groups composed of a rather +short, unpaired primary and several short, unpaired secondaries +(0-28 in <i>phlebodes</i>, 0-23 in <i>sartori</i>). The mean duration of the +primary of <i>phlebodes</i> is 0.105 (0.07-0.16) second and that of the +secondaries is 0.067 (0.035-0.12) second. The mean duration of the +primary of <i>sartori</i> is 0.080 (0.07-0.09) second and that of the +secondaries is 0.053 (0.035-0.07) second.</p> + +<p>The two subspecies of <i>H. microcephala</i> are identical in call pattern +and agree closely in duration of notes, although those of the nominate +subspecies tend to be slightly longer. <i>Hyla robertmertensi</i> is +distinctive in call pattern in that it is the only species having a paired +primary; the duration of the primary is completely overlapped by +that in the other species, but the secondaries tend to be the shortest +in the group. The call patterns of <i>H. phlebodes</i> and <i>H. sartori</i> are +identical and the range of duration of notes of <i>phlebodes</i> completely +overlaps that of <i>sartori</i>, although both the primary and secondary +notes of the latter tend to be somewhat shorter (<a href="#Table_5">Table 5</a>, <a href="#Pl_16">Pl. 16</a>).</p> + +<p><i>Fundamental frequency.</i>—This parameter was analyzed for the +primary notes. It was measured for the secondaries as well and +was found to differ in magnitude in the same way as the primary +note. In a few examples of both subspecies of <i>H. microcephala</i> a +high <a name="primary"></a><a href="#typos">primary</a> note, in which the fundamental frequency is exceptionally +high, is sometimes emitted (Fouquette, 1960b). None of +these notes was used in this analysis; only the fundamental frequencies +of normal primary notes are compared (<a href="#Table_5">Table 5</a>, <a href="#Fig_7">Fig. 7</a>).</p> +<p> </p> +<p> </p> + + +<span class="pagenum"><a name="Page_546" id="Page_546">[Pg 546]</a></span> +<a name="Table_5"></a> +<span class="smcap">Table 5.</span>—Comparison of Normal Mating Calls in the Hyla microcephala Group. (Observed Range Given in Parentheses Below +Mean; Unless Otherwise Noted Data Are for Primary Notes.).<br> + +<div class="center"> +<table width="100%" class="data" summary="Teeth Numer Variation Hyla microcephala Group"> +<tr><td colspan=7 class="bb"> </td></tr> +<tr><td rowspan=2 class="bt bb">Species</td><td rowspan=2 class="bt bl bb">N</td><td rowspan=2 class="bt bl bb">Dominant frequency (cps)</td><td rowspan=2 class="bt bl bb">Fundamental frequency (cps)</td><td colspan=2 class="bt bl bb">Duration of notes (seconds)</td><td rowspan=2 class="bt bl bb">Repetition rate of secondaries (notes/minute)</td></tr> +<tr><td class="bl bb">Primary</td><td class="bl bb">Secondary</td></tr> +<tr><td><i>H. m. microcephala</i></td><td class="bl">44</td><td class="bl">5637</td><td class="bl">205</td><td class="bl">0.13</td><td class="bl">0.10</td><td class="bl">268</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">(5150-5962)</td><td class="bl"> (184-244) </td><td class="bl"> (0.11-0.16) </td><td class="bl"> (0.05-0.14) </td><td class="bl"> (192-353)</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td></tr> +<tr><td><i>H. m. underwoodi</i></td><td class="bl">47</td><td class="bl">5772</td><td class="bl">220</td><td class="bl">0.11</td><td class="bl">0.09</td><td class="bl">283</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">(5177-6200)</td><td class="bl">(192-275)</td><td class="bl">(0.05-0.15)</td><td class="bl"> (0.06-0.11) </td><td class="bl"> (197-384)</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td></tr> +<tr><td><i>H. robertmertensi</i></td><td class="bl">25</td><td class="bl">5388</td><td class="bl">162</td><td class="bl">0.09</td><td class="bl">0.04</td><td class="bl">418</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">(5150-5785)</td><td class="bl"> (140-178) </td><td class="bl"> (0.07-0.11) </td><td class="bl"> (0.03-0.06) </td><td class="bl">(368-570)</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td></tr> +<tr><td><i>H. phlebodes</i></td><td class="bl">34</td><td class="bl">3578</td><td class="bl">148</td><td class="bl">0.11</td><td class="bl">0.07</td><td class="bl">284</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">(3220-4067)</td><td class="bl">(125-158)</td><td class="bl">(0.07-0.16)</td><td class="bl">(0.04-0.12)</td><td class="bl">(210-350)</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td><td class="bl"> </td></tr> +<tr><td><i>H. sartori</i></td><td class="bl">10</td><td class="bl">3217</td><td class="bl">126</td><td class="bl">0.08</td><td class="bl">0.05</td><td class="bl">434</td></tr> +<tr><td> </td><td class="bl"> </td><td class="bl">(2950-3600)</td><td class="bl"> (116-135) </td><td class="bl"> (0.07-0.09) </td><td class="bl"> (0.04-0.07) </td><td class="bl"> (396-477)</td></tr> +</table> +</div> +<p> </p> +<p> </p> + +<p>The two subspecies of <i>H. microcephala</i> agree closely in fundamental +frequency. There is considerable overlap, but the difference +between the means is significant at the 0.001 level of probability +(t = 4.2406). The call of <i>H. robertmertensi</i> does not overlap that +<span class="pagenum"><a name="Page_547" id="Page_547">[Pg 547]</a></span> +of <i>H. sartori</i> or either subspecies of <i>H. microcephala</i> in this parameter; +but it does overlap that of <i>H. phlebodes</i>, although again the +difference between the means is significant at the 0.001 level +(t = 9.360). <i>Hyla phlebodes</i> and <i>sartori</i> have the lowest fundamental +frequencies, and there is some overlap, but here too the +difference between the means is significant at the 0.001 level +(t = 4.923).</p> + +<p><i>Dominant frequency.</i>—A dominant <a name="of_of"></a><a href="#typos">band of of frequencies</a> cuts +across the harmonics of the fundamental, obscuring the harmonic +pattern and generally shifting upward in frequency. The midpoint +of this band is measured at the terminal border as the dominant +frequency. As with the fundamental frequency, only the normal +primary notes were utilized in the comparisons (<a href="#Table_5">Table 5</a>, <a href="#Fig_8">Fig 8</a>).</p> +<p> </p> +<p> </p> + +<a name="Fig_7"></a> +<div class="center"> +<img src="images/fig_7.png" width="600" height="355" title="Variation in fundamental frequency" alt="Variation in fundamental frequency"><br><br> +<span class="smcap">Fig. 7.</span> Variation in the fundamental frequency of the normal primary notes +in the <i>Hyla microcephala</i> group. The horizontal lines = range of variation, +vertical lines = mean, solid bars = twice the standard error of the mean, and +open bars = one standard deviation. The number of specimens in each +sample is indicated in parentheses after the name of the taxon.<br> +</div> +<p> </p> +<p> </p> + +<p>The two subspecies of <i>H. microcephala</i> agree more closely in this +parameter than in fundamental frequency. The overlap is great, +but the difference between the means is significant at the 0.001 level +(t = 3.658). The calls of both subspecies completely overlap that +of <i>robertmertensi</i> in this parameter, but the difference between the +means is significant at the 0.001 level. The calls of <i>H. phlebodes</i> +and <i>H. sartori</i> overlap considerably in this characteristic, although +the difference between the means is significant at the 0.001 level +(t = 7.504) (<a href="#Fig_9">Fig. 9</a>). The call of neither species overlaps those +of <i>H. microcephala</i> and <i>robertmertensi</i>.</p> +<p> </p> +<p> </p> + +<p><span class="pagenum"><a name="Page_548" id="Page_548">[Pg 548]</a></span></p> +<div class="center"> +<a name="Fig_8"></a> +<img src="images/fig_8.png" width="600" height="353" title="Variation in the mid-point frequency" alt="Variation in the mid-point frequency"><br><br> +<span class="smcap">Fig. 8.</span> Variation in the mid-point of the dominant frequency band of the +normal primary notes in the <i>Hyla microcephala</i> group. The horizontal lines += range of variation, vertical lines = mean, solid bars = twice the standard +error of the mean, and open bars = one standard deviation. The number of +specimens in each sample is indicated in parentheses after the name of the +taxon.<br> +<p> </p> +<p> </p> + +<a name="Fig_9"></a> +<img src="images/fig_9.png" width="600" height="327" title="Scattergram" alt="Scattergram"><br><br> +<span class="smcap">Fig. 9.</span> Scatter diagram relating the dominant and fundamental frequencies +of the normal primary notes in the <i>Hyla microcephala</i> group. Each symbol +represents a different individual.<br> +<p> </p> +<p> </p> +</div> + +<p><i>Repetition rate.</i>—The repetition rate of the secondary notes, in +calls consisting of more than one secondary, was measured for each +form. A considerable amount of variation in this parameter was +found in all of the taxa (<a href="#Table_5">Table 5</a>). This variation probably is due +in part to the effect of temperature differences. Repetition rate is +<span class="pagenum"><a name="Page_549" id="Page_549">[Pg 549]</a></span> +the only parameter analyzed for which there is a correlation with +the air-temperature, but even here the correlation is weak, probably +due to the microenvironmental effects of humidity, air-movement, +and other factors in addition to the ambient air temperature that +influences the body temperature of the frogs. These rates are +nearly alike in both subspecies of <i>H. microcephala</i> and in <i>phlebodes</i>. +The repetition rates in <i>H. robertmertensi</i> and <i>H. sartori</i> are considerably +faster than in the other three taxa. <i>Hyla sartori</i> has the +fastest repetition rate of the group.</p> + +<p>In all characteristics of the mating calls the two subspecies of +<i>H. microcephala</i> agree closely, as might be expected, although the +differences are statistically significant. <i>Hyla robertmertensi</i> is distinctive +in call pattern and seems to be closer to <i>microcephala</i> in +dominant frequency but closer to <i>H. phlebodes</i> in fundamental frequency. +Thus, it is somewhat intermediate between <i>microcephala</i> +and <i>phlebodes</i>. The identical pattern and similarity in fundamental +and dominant frequencies of the calls of <i>H. phlebodes</i> and <i>H. sartori</i> +possibly indicate close relationship.</p> + +<p><i>Geographic variation in call.</i>—<i>Hyla m. microcephala</i> has higher +fundamental and dominant frequencies in Costa Rica than in Panamá. +In Costa Rican <i>H. m. underwoodi</i> the fundamental and dominant +frequencies are lower than in other parts of the range. Frogs of +this subspecies recorded in Nicaragua and Honduras have slightly +lower dominant frequencies and higher fundamental frequencies +than those recorded in Guatemala or Oaxaca. The duration of both +primary and secondary notes decreases to the south; samples from +Nicaragua and Costa Rica have the shortest notes. Comparison +of duration of notes in the two subspecies shows that the Panamanian +<i>H. m. microcephala</i> have slightly longer notes than do any +<i>H. m. underwoodi</i>; the more northern populations of <i>H. m. underwoodi</i> +from México most closely approach <i>H. m. microcephala</i> in +this characteristic.</p> + +<p>The calls of <i>H. robertmertensi</i> in Oaxaca have higher dominant +and fundamental frequencies and longer secondary notes than do +those in Chiapas.</p> + +<p>The calls of <i>H. phlebodes</i> recorded at Puerto Viejo, Costa Rica, +have slightly lower dominant frequencies than do those recorded +at Turrialba, Costa Rica, and in Panamá, whereas those recorded at +Turrialba have lower fundamental frequencies than in other samples. +The duration of notes is slightly shorter in both Costa Rican samples +than in those recorded in Panamá.</p> +<p> </p> +<p> </p> + +<p><span class="pagenum"><a name="Page_550" id="Page_550">[Pg 550]</a></span></p> +<a name="LIFE_HISTORY" id="LIFE_HISTORY"></a> +<div class="caption2">LIFE HISTORY</div> + +<p>The frogs of the <i>Hyla microcephala</i> group breed in shallow grassy +ponds. In some places they breed in permanent ponds, but usually +congregate around temporary pools, such as depressions in forests, +flooded fields, and roadside ditches. At the height of their breeding +season, usually in the early part of the rainy season, the congregations +are made up of large numbers of individuals. In April, 1961, +and in June, 1966, the senior author noted nearly continuous +choruses of <i>H. m. microcephala</i> in roadside ditches along the 75 +kilometers of road between Villa Neily and Palmar Sur, Puntarenas +Province, Cost Rica; on June 20, 1966, at Puerto Viejo, Heredia +Province, Costa Rica, he estimated approximately 900 <i>Hyla phlebodes</i> +in one pond, and two nights later noticed that the number +of individuals <a name="had"></a><a href="#typos">had</a> increased substantially. Other observations by +the first author on size of breeding congregations include nearly +continuous choruses of <i>H. m. underwoodi</i> between Villahermosa and +Teapa, Tabasco, in July of 1958, an estimated 400 <i>Hyla robertmertensi</i> +in a road side ditch 7.2 kilometers west-northwest of Zanatepec, +Oaxaca, on July 13, 1956, and approximately 150 <i>Hyla sartori</i> around +a rocky pool in a riverbed, 11.8 kilometers west-northwest of Tierra +Colorada, Guerrero, on June 28, 1958.</p> + +<p>The length of the breeding season seemingly is more dependent +on climatic conditions in various parts of Middle America than on +behavioral differences in the various species. Thus, Fouquette +(1960b) found in the Canal Zone that <i>H. m. microcephala</i> formed +breeding choruses from May through January, the entire rainy +season in that area. In the wetter coastal region of Puntarenas +Province, Costa Rica, the species breeds as early as mid-March, +whereas in the drier region encompassing Guanacaste Province, +Costa Rica, and southwestern Nicaragua breeding activity is initiated +by the first heavy rains of the season, usually in June.</p> + +<p><i>Hyla phlebodes</i> inhabits regions having rainfall throughout the +year. Although large breeding congregations are most common in +the early parts of the rainy season, males probably call throughout +the year. At Puerto Viejo in Costa Rica the senior author has heard +<i>Hyla phlebodes</i> in February, April, June, July, and August. Charles +W. Myers noted calling males of this species in the area around +Almirante, Bocas del Toro Province, Panamá, in September, October, +and February. An exception to the correlation between rainfall and +breeding activity was noted by the junior author in <i>Hyla phlebodes</i> +in the Canal Zone, where he noticed a decrease in activity of that +species in October and November, when the rains are heaviest and +<span class="pagenum"><a name="Page_551" id="Page_551">[Pg 551]</a></span> +most frequent. Furthermore, independent observations made by +both of us indicate that <i>H. phlebodes</i> does not reach peaks of +activity during or immediately after heavy rains, but instead builds +up to peaks of activity two or three days after a heavy rain. This +is in contrast to the other species, all of which characteristically inhabit +drier environments than does <i>H. phlebodes</i>. Peaks of breeding +activity in the other species occur immediately after, or even +during, heavy rains.</p> + +<p>The calling location of the males generally is on vegetation above, +or at the edge of, the water. <i>Hyla microcephala</i> and <i>H. phlebodes</i> +call almost exclusively from grasses and sedges; <i>phlebodes</i> usually +calls from taller and more dense grasses than does <i>microcephala</i>. +Except for some minor differences in calling location observed by +the junior author (Fouquette, 1960b) in the Canal Zone, the differences +in density and height of grasses utilized for calling-locations +probably is dependent primarily on the nature of the available +vegetation. Although bushes and broad-leafed herbs are usually +present at the breeding sites, males of these species seldom utilize +them for calling locations. Both <i>H. robertmertensi</i> and <i>H. sartori</i> +have been observed calling from grasses, herbs, bushes, and low +trees. Calling males of <i>robertmertensi</i> have been found two meters +above the ground in small trees.</p> + +<p>Daytime retreats in the breeding season sometimes are no more +than shaded <a name="clumps"></a><a href="#typos">clumps</a> of vegetation adjacent to a pond or in clumps +of grass in a pond. Individuals of <i>H. m. underwoodi</i> were found by +day under the outer sheaths of banana plants next to a water-filled +ditch. Dry season refuges are unknown.</p> + +<p>Amplexus is axillary in all four species. Egg deposition has been +observed in <i>H. m. microcephala</i>, <i>m. underwoodi</i>, and <i>phlebodes</i>. +In all three the eggs are deposited in small masses that float near +the surface of the water and usually are at least partly attached to +emergent vegetation. Each clutch does not represent the entire egg +complement of the female.</p> + +<p>Tadpoles are definitely known of only <i>H. m. microcephala</i> and +<i>phlebodes</i>; these have been described in the preceding accounts of +the species. The tadpoles of these two species can be distinguished +readily (<a href="#Pl_15">Pl. 15</a>). The tadpole of <i>H. microcephala</i> has a uniformly +white venter and nearly transparent tail, whereas in <i>H. phlebodes</i> +the venter is flecked anteriorly and the tail is mottled. In life, <i>H. +microcephala</i> is easily recognized by the orange posterior half of +the tail, whereas the tail in <i>H. phlebodes</i> is mottled tan and grayish +brown.</p> +<p> </p> +<p> </p> + +<p><span class="pagenum"><a name="Page_552" id="Page_552">[Pg 552]</a></span> +<a name="PHYLOGENETIC_RELATIONSHIPS" id="PHYLOGENETIC_RELATIONSHIPS"></a> +<div class="caption2">PHYLOGENETIC RELATIONSHIPS</div> + +<p>The evidence already presented on osteology, external structure, +coloration, mating call, and life history emphatically show that the +four species under consideration are a closely related assemblage. +Now the question arises: To what other groups in the genus is the +<i>Hyla microcephala</i> group related? Furthermore, it is pertinent +to this discussion to attempt a reconstruction of the phylogeny of +the group as a whole and of the individual species in the <i>Hyla +microcephala</i> group. With regard to the relationships of the group +we must take into account certain species in South America. Our +endeavors there are hampered by the absence of data on the mating +calls and life histories of most of the relevant species.</p> + +<p>As mentioned in the <a name="account"></a><a href="#typos">account</a> of <i>Hyla m. microcephala</i>, the species +<i>microcephala</i> possibly is subspecifically related to <i>Hyla misera</i>, a +frog widespread in the Amazon Basin. <i>Hyla misera</i> resembles +<i>microcephala</i> in coloration, external structure, and cranial characters. +The frontoparietals are equally poorly ossified, and the frontoparietal +fontanelle is extensive. Our principal reason for not considering +the two taxa conspecific at this time is our lack of knowledge +concerning the color of living <i>H. misera</i>, the structure of the tadpoles, +and the characteristics of the mating call. Even with the +absence of such data that we think essential to establish the nomenclature +status of the taxa, we are confident that the two are sufficiently +closely related that any discussion of the phylogenetic relationships +of one species certainly must involve consideration of the +other.</p> + +<p><i>Hyla misera</i> possibly is allied to other small yellowish tan South +American <i>Hyla</i> that lack dark pigmentation on the thighs. Probable +relatives are <i>Hyla elongata</i>, <i>minuta</i> (with <i>goughi</i>, <i>pallens</i>, <i>suturata</i>, +<i>velata</i>, and possibly others as synonyms), <i>nana</i>, and <i>werneri</i>. The +consideration of the interspecific relationships of these taxa is beyond +the scope of this paper, but we can say that each of these +species has a pale yellowish tan dorsum, relatively broad dorsolateral +brown stripe, and narrow longitudinal brown lines or irregular marks +on the dorsum. Furthermore, examination of the skulls of <i>elongata</i>, +<i>nana</i>, and <i>werneri</i> reveals that they are like <i>misera</i> and <i>microcephala</i> +in the nature of the frontoparietal fontanelle and in having a greatly +reduced quadratojugal. Thus, on the basis of cranial and external +characters the <i>Hyla microcephala</i> group can be associated with <i>Hyla +misera</i> and its apparent allies in South America. This association +can be only tentative until the mating calls, tadpoles, and chromosome +numbers of the South American species are known.</p> + +<p><span class="pagenum"><a name="Page_553" id="Page_553">[Pg 553]</a></span> +Among the Middle American hylids, only the <i>Hyla microcephala</i> +group and <i>H. ebraccata</i> have a haploid number of 15 chromosomes +(Duellman and Cole, 1965). All other New World <i>Hyla</i>, for which +the number is known, have a haploid number of 12; the only other +<i>Hyla</i> having 15 is a Papuan <i>Hyla angiana</i> (Duellman, 1967).</p> + +<p><i>Hyla ebraccata</i> occurs in the humid tropical lowlands of Middle +America and the Pacific lowlands of northwestern South America. +It is the northernmost, and only Central American, representative +of the <i>Hyla leucophyllata</i> group, which is diverse (about 10 species +currently recognized) and widespread in tropical South America +east of the Andes. This group is characterized by having broad, +flat skulls with larger nasals and more ossification of the frontoparietals +than in the <i>Hyla microcephala</i> group. The quadratojugal +is present as a small anteriorly projecting spur that does not connect +with the maxillary. Externally, the <i>Hyla leucophyllata</i> group is +characterized by having a well-developed axillary membrane, uniformly +yellow thighs, and a dorsal color pattern in many species +consisting of a dark lateral band, a pale dorsolateral band or dorsal +ground color, and a large middorsal dark mark. In some species, +the dorsal pattern consists of small dark markings or is nearly uniformly +pale. At least in the Central American <i>Hyla ebraccata</i>, the +mating call consists of a single primary note followed by a series of +shorter secondary notes, the tadpoles have xiphicercal tails and lack +teeth, and the haploid number of chromosomes is 15. On the +strength of these observations it seems imperative to consider the +<i>Hyla leucophyllata</i> group as a close ally to the <i>Hyla microcephala</i> +group. Successful artificial hybridization supports the close relationship +of <i>H. m. microcephala</i> and <i>phlebodes</i>; partial success of +artificial hybridization of these two with <i>ebraccata</i> (Fouquette, +1960b) provides further evidence for close relationship between the +<i>Hyla leucophyllata</i> and <i>Hyla microcephala</i> groups.</p> + +<p>In México and northern Central America two small species, <i>Hyla +picta</i> and <i>Hyla smithi</i>, comprise the <i>Hyla picta</i> group. These frogs +resemble members of the <i>Hyla microcephala</i> group by having a +yellowish tan dorsum with a dorsolateral white stripe and uniformly +yellow thighs. Furthermore the mating call is not unlike those of +the species in the <i>Hyla microcephala</i> group. Despite these similarities, +the <i>Hyla picta</i> group differs from the <i>Hyla microcephala</i> +group by having a well-developed quadratojugal that connects to +the maxillary, tadpoles with teeth present and caudal fins completely +enclosing the caudal musculature, and a haploid number +of 12 chromosomes. In all of these characteristics the frogs of the +<span class="pagenum"><a name="Page_554" id="Page_554">[Pg 554]</a></span> +<i>Hyla picta</i> group more closely resemble other Middle American +<i>Hyla</i> than they do the <i>Hyla microcephala</i> group. Therefore, it +can best be presumed that the superficial resemblances of coloration +and the mating call are the result of convergence.</p> + +<p>Since the <i>Hyla microcephala</i> and <i>leucophyllata</i> groups apparently +are related and since the greatest diversity of these frogs is in +South America (if <i>Hyla misera</i> and its relatives are placed with the +<i>Hyla microcephala</i> group), it seems appropriate to place the +centers of origins of these groups in South America. Therefore, +the <i>Hyla microcephala</i> group and <i>Hyla ebraccata</i> of the <i>Hyla leucophyllata</i> +group either have immigrated into Central America, or +they are representatives of those groups that were isolated in +Central America during most of the Cenozoic when South America +was separated from Central America.</p> + +<p>The interspecific relationships of the species in the <i>Hyla microcephala</i> +group are not clear. On the basis of coloration, <i>H. m. microcephala</i> +and <i>H. robertmertensi</i> are close, and <i>H. m. underwoodi</i> and +<i>H. phlebodes</i> are nearly identical. The mating calls of <i>H. phlebodes</i> +and <i>sartori</i> closely resemble one another, whereas the call of <i>robertmertensi</i> +is intermediate between these and <i>microcephala</i>.</p> + +<p>In most respects <i>Hyla microcephala</i> is distinct from the other +species, and with the exception of the amount of ossification of the +frontoparietals, the other species can be easily derived from a +<i>microcephala</i>-like ancestor. Possibly the slightly increased ossification +of the frontoparietals in <i>robertmertensi</i>, <i>phlebodes</i>, and <i>sartori</i> +is secondary, or possibly after differentiation of the species the +amount of ossification was further reduced in <i>microcephala</i>. If so, +the species fall into a reasonable phylogenetic scheme that has +<i>microcephala</i> as the extant species most like the ancestral stock.</p> + +<p>We visualize the evolutionary history of the group to have followed +a course that began with the invasion of Central America by +a <i>microcephala</i> ancestral stock that differentiated into two populations +in lower Central America—a <i>microcephala</i>-like frog on the +Pacific lowlands and a <i>phlebodes</i>-like frog on the Caribbean lowlands. +Differentiation could have been brought about by isolation +by montaine or marine barriers. The population on the Pacific +lowlands either was preadapted for subhumid conditions or became +so adapted and dispersed northward onto the Pacific lowlands of +northern Central America. Simultaneously the frogs on the Caribbean +lowlands, which were adapted to humid environments, dispersed +northward in the humid forested regions to southern México +and crossed the Isthmus of Tehuantepec onto the Pacific slopes of +<span class="pagenum"><a name="Page_555" id="Page_555">[Pg 555]</a></span> +Oaxaca and Guerrero northward to Jalisco. Subsequent development +of arid conditions, possibly in the Pliocene, Pleistocene, or even as +late as the Thermal Maximum in post-Wisconsin time, resulted in +a restriction of the ranges in northern Central America, thereby +isolating part of the <i>phlebodes</i>-stock on the Pacific slopes of México, +where it adapted to drier conditions and evolved into <i>sartori</i>. The +rest of the <i>phlebodes</i>-stock was restricted to the humid forests on +the Caribbean lowlands of lower Central America. The increased +aridity on the Pacific lowlands eliminated the <i>microcephala</i>-stock +from southern Honduras and northwestern Nicaragua and in so +doing left an isolated population on the lowlands of Chiapas and +Guatemala, which differentiated into <i>robertmertensi</i>. The original +stock on the Pacific lowlands of Panamá and southeastern Costa +Rica became <i>microcephala</i>.</p> + +<p>If the <i>microcephala</i>-stock was, as we believe, better adapted for +existence under subhumid conditions than was the <i>phlebodes</i>-stock, +the development of subhumid conditions in much of the lowland +region of northern Central America and southern México would +have permitted the expansion of the range of <i>microcephala</i> into the +area now inhabited by <i>H. m. underwoodi</i>, while <i>phlebodes</i> was +being eliminated from this area by climatic conditions that were +unsuited to its survival there. Perhaps the similarity in coloration +of <i>H. m. underwoodi</i> and <i>phlebodes</i> is the result of convergence or +possibly hybridization occurred at the time the former was expanding +its range and the latter's range was being restricted. If hybridization +did occur, the differences in mating call subsequently +were enhanced, thereby providing a valid isolating mechanism in +sympatric populations.</p> + +<p><i>Hyla microcephala</i> and <i>phlebodes</i> range into northern South +America. Probably both species entered South America in relatively +recent times after they had differentiated from one another in +Central America.</p> +<p> </p> +<p> </p> + +<p><span class="pagenum"><a name="Page_556" id="Page_556">[Pg 556]</a></span></p> +<a name="LITERATURE_CITED" id="LITERATURE_CITED"></a> +<div class="caption2">LITERATURE CITED</div> + +<span class="smcap">Boulenger, G. A.</span><br> +<div class="references"><p>1898. Fourth report on additions to the batrachian collection in the Natural-History Museum. Proc. Zool. Soc. London, 1898, pp. 373-482, pls. 38-39. October 1.</p></div> +<div class="references"><p>1899. Descriptions of new batrachians in the collection of the British Museum (Natural History). Ann. Mag. Nat. Hist, ser. 7, 3:273-277, pls. 11-12.</p></div> +<br> +<span class="smcap">Breder, C. M. Jr.</span><br> +<div class="references"><p>1946. Amphibians and reptiles of the Rio Chucunaque Drainage, Darien, Panama, with notes on their life histories and habits. Bull. Amer. Mus. Nat. Hist, 86:375-436, pls. 42-60, August 26.</p></div> +<br> +<span class="smcap">Cole, L. J. and Barbour, T.</span><br> +<div class="references"><p>1906. Vertebrata from Yucatan: Reptilia; Amphibia; Pisces. Bull. Mus. Comp. Zool., 50:146-159. November.</p></div> +<br> +<span class="smcap">Cope, E. D.</span><br> +<div class="references"><p>1886. Thirteenth contribution to the herpetology of tropical America. Proc. Amer. Philos. Soc, 23:271-287. February 11.</p></div> +<div class="references"><p>1894. Third addition to a knowledge of the Batrachia and Reptilia of +Costa Rica. Proc. Acad. Nat. Sci. Philadelphia, 1894, pp. 194-206.</p></div> +<br> +<span class="smcap">Duellman, W. E.</span><br> +<div class="references"><p>1956. The frogs of the hylid genus <i>Phrynohyas</i> Fitzinger, 1843. Misc. Publ. Mus. Zool., Univ. Michigan, 96:1-47, pls. 1-6. February 21.<br></div> +<div class="references"><p>1967. Additional studies of chromosomes of anuran amphibians. Syst. Zool., 16:38-43, March 17.</p></div> +<br> +<span class="smcap">Duellman, W. E. and Cole, C. J.</span><br> +<div class="references"><p>1965. Studies of chromosomes of some anuran amphibians (Hylidae and Centrolenidae). Syst. Zool., 14:139-143. July 9.</p></div> +<br> +<span class="smcap">Duellman, W. E. and Trueb, L.</span><br> +<div class="references"><p>1966. Neotropical hylid frogs, genus Smilisca. Univ. Kansas Publ., Mus. Nat. Hist., 17:281-375, pls. 1-12. July 14.</p></div> +<br> +<span class="smcap">Dunn, E. R.</span><br> +<div class="references"><p>1931. The amphibians of Barro Colorado Island. Occas. Papers Boston Soc. Nat. Hist., 5:403-421. October 10.</p></div> +<div class="references"><p>1933. Amphibians and reptiles from El Valle de Anton, Panamá. <i>Ibid.</i>, 8:65-79. June 7.</p></div> +<div class="references"><p>1934. Two new frogs from Darien. Amer. Mus. Novit., 747:1-2. September 17.</p></div> +<br> +<span class="smcap">Fouquette, M. J. Jr.</span><br> +<div class="references"><p>1960a. Call structure in frogs of the family Leptodactylidae. Texas Jour. Sci., 12:201-215. October.</p></div> +<div class="references"><p>1960b. Isolating mechanisms in three sympatric tree frogs in the Canal Zone. Evolution, 14:484-497. December 16.</p></div> +<br> +<span class="smcap">Gaige, H. T., Hartweg, N. and Stuart, L. C.</span><br> +<div class="references"><p>1937. Notes on a collection of amphibians and reptiles from eastern Nicaragua. Occas. Papers Mus. Zool., Univ. Michigan, 357:1-18. October 26.</p></div> +<br> +<span class="smcap">Gosner, K. L.</span><br> +<div class="references"><p>1960. A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica, 16:183-190. September 23.</p></div> +<br> +<p><span class="pagenum"><a name="Page_557" id="Page_557">[Pg 557]</a></span></p> +<span class="smcap">Kellogg, R.</span><br> +<div class="references"><p>1932. Mexican tailless amphibians in the United States National Museum. Bull. U.S. Natl. Mus., 160:1-224. March 31.</p></div> +<br> +<span class="smcap">Rivero, J. A.</span><br> +<div class="references"><p>1961. Salientia of Venezuela. Bull. Mus. Comp. Zool., 126:1-207. November.</p></div> +<br> +<span class="smcap">Smith, H. M.</span><br> +<div class="references"><p>1951. The identity of <i>Hyla underwoodi</i> Auctorum of Mexico. Herpetologica, 7:184-190. December 31.</p></div> +<br> +<span class="smcap">Stejneger, L.</span><br> +<div class="references"><p>1906. A new tree toad from Costa Rica. Proc. U. S. Natl. Mus., 30:817-818. June 4.</p></div> +<br> +<span class="smcap">Stuart, L. C.</span><br> +<div class="references"><p>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. <a name="Michigan"></a><a href="#typos">Michigan</a>, 29:1-56, pls. 1-4. October 4.</p></div> +<br> +<span class="smcap">Taylor, E. H.</span><br> +<div class="references"><p>1952. The frogs and toads of Costa Rica. Univ. Kansas Sci. Bull., 35-577-942. July 1.</p></div> +<p> </p> +<p><i>Transmitted July 11, 1967.</i></p> +<p> </p> +<p> </p> + + + + + + + + + + +<pre> + + + + + +End of the Project Gutenberg EBook of Middle American Frogs of the Hyla +microcephala Group, by William E. Duellman and M. J. 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Duellman and M. J. Fouquette + +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: Middle American Frogs of the Hyla microcephala Group + +Author: William E. Duellman + M. J. Fouquette + +Release Date: December 9, 2010 [EBook #34604] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK MIDDLE AMERICAN FROGS *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +https://www.pgdp.net + + + + + + + + + +University of Kansas Publications + +Museum of Natural History + +Volume 17, No. 12, pp. 517-557, pls. 13-16, 9 figs. +March 20, 1968 + +Middle American Frogs +of the Hyla microcephala Group + +BY + +WILLIAM E. DUELLMAN AND M. J. FOUQUETTE, JR. + +University of Kansas +Lawrence +1968 + + + + +University of Kansas Publications, Museum of Natural History + +Editors: E. Raymond Hall, Chairman, Henry S. Fitch, +Frank B. Cross + +Volume 17, No. 12, pp. 517-557, 4 pls. 9 figs. +Published March 20, 1968 + +University of Kansas +Lawrence, Kansas + +PRINTED BY +ROBERT R. (BOB) SANDERS, STATE PRINTER +TOPEKA, KANSAS +1968 + +31-9419 + + + + +Middle American Frogs +of the Hyla microcephala Group + +BY +WILLIAM E. DUELLMAN AND M. J. FOUQUETTE, JR. + + +CONTENTS + + + PAGE + +Introduction 519 + Acknowledgments 520 + Materials and Methods 520 + +Hyla microcephala Group 521 + Key to Species and Subspecies 522 + +Accounts of Species and Subspecies 523 + +Cranial Osteology 540 + +Analysis of Mating Calls 544 + +Life History 550 + +Phylogenetic Relationships 552 + +Literature Cited 556 + + + + + +INTRODUCTION + + +The small yellow tree frogs, _Hyla microcephala_ and its relatives, +are among the most frequently heard and commonly collected frogs in +the lowlands of southern Mexico and Central America. The similarities +in size, proportions, and coloration of the different species have +resulted not so much in a multiplicity of specific names, but in +differences of opinion on the application of existing names to the +various taxa. For example, the populations on the Atlantic lowlands +have been known by three names, two of which have been applied to +other taxa. Much of the confusion has been the result of previous +workers' unfamiliarity with the animals in life and unawareness of the +intraspecific geographic variation in the most widespread species. + +Independently we undertook studies of these frogs in the field. The +second author worked on the interspecific relationships and isolating +mechanisms in Panama (Fouquette, 1960b) and later studied the species +in southern Mexico. As part of his survey of the hylids of Middle +America, the first author accumulated field and laboratory data on the +frogs throughout their ranges in Mexico and Central America. The +purpose of this report is to present our findings on the four species +of Middle American frogs that we place in the _Hyla microcephala_ +group. In addition to conventional taxonomic characters, we have +utilized the features of the cranial osteology and have relied heavily +on the data obtained from an analysis of the mating calls. +Furthermore, we have included ecological and distributional data in +our synthesis of interspecific relationships. + + +ACKNOWLEDGMENTS + +Examination of specimens was made possible by the provision of working +space at various institutions or through the loan of specimens. For +their generosity in this manner we are grateful to Richard J. Baldauf, +Charles M. Bogert, James E. Boehlke, Doris M. Cochran, Robert F. Inger, +John M. Legler, Alan E. Leviton, Gerald Raun, Jay M. Savage, Hobart M. +Smith, Robert C. Stebbins, Wilmer W. Tanner, Charles F. Walker, Ernest +E. Williams, and Richard G. Zweifel. + +Duellman is especially grateful to Charles W. Myers, Linda Trueb, +Jerome B. Tulecke, and John Wellman for their assistance in the field +and to Linda Trueb for her work on the cranial osteology that is +incorporated in this report. Fouquette is indebted to H. Morgan Smith +and A. C. Collins for assistance in the field, to A. J. Delahoussaye +for assistance in the laboratory, and to W. Frank Blair for use of the +facilities of the sound laboratory at the University of Texas and for +much help in the early stages of this study. + +The research reported herein was accomplished mainly through support +by the National Science Foundation (grants NSF G-9827 and GB-1441 to +Duellman and GB-599 to Fouquette). The latter's field work in Mexico +was assisted in part by NSF Grant G-4956 to W. Frank Blair. Some of +the field studies carried out in Panama by Duellman were supported by +a grant from the National Institutes of Health (NIH GM-12020). + +We are grateful to many persons, too numerous to mention, who in +various ways aided our field work in Middle America. We are especially +indebted to Dr. Rodolfo Hernandez Corzo and the late Ing. Luis Macias +Arellano of the Direccion General de la Fauna Silvestre of the Mexican +government for providing permits to collect in Mexico. + + +Materials and Methods + +For this report, data has been obtained from 2829 preserved frogs, 42 +skeletal preparations, 8 lots of tadpoles and young, and 4 lots of +eggs. Much of the material was collected in our independent field +work, which has extended over a period of 11 years. + +Measurements were taken in the manner described by Duellman (1956). +Osteological data were obtained from specimens that were cleared in +potassium hydroxide, stained with alizarin red, and stored in +glycerine. Recordings were made by means of Magnemite portable tape +recorders (Amplifier Corp. America). The calls recorded by Fouquette +were analyzed on a Sonagraph (Kay Electric Co.) at the University of +Texas; those recorded by Duellman were analyzed mainly on a Vibralyzer +(Kay Electric Co.) at the University of Kansas and in part on a +Sonagraph at the University of Southwestern Louisiana. Sample calls +were analyzed on all three instruments; the slight differences in +results were found to be less than the error in measurement, so the +data from all sources were combined without correction. The techniques +and terminology of the calls are those defined by Fouquette (1960a, +1960b). + +In the accounts of the species we have attempted to give a complete +synonymy. At the end of each species account the localities from which +specimens were examined are listed alphabetically within each state, +province, or department, which in turn are listed alphabetically +within each country. The countries are arranged from north to south. +Localities preceded by an asterisk (*) are not plotted on the +accompanying maps due to the crowding of symbols that would have +resulted. Abbreviations for museum specimens are listed below: + + AMNH --American Museum of Natural History + ANSP --Academy of Natural Sciences of Philadelphia + BMNH --British Museum (Natural History) + BYU --Brigham Young University + CAS --California Academy of Sciences + FMNH --Field Museum of Natural History + KU --University of Kansas Museum of Natural History + MCZ --Museum of Comparative Zoology + MVZ --Museum of Vertebrate Zoology + SU --Stanford University + UIMNH--University of Illinois Museum of Natural History + UMMZ --University of Michigan Museum of Zoology + USC --University of Southern California + USNM --United States National Museum + UU --University of Utah + TCWC --Texas Cooperative Wildlife Collection + TNHM --Texas Natural History Museum + + + + + +HYLA MICROCEPHALA GROUP + + +_Definition._--Small hylids attaining a maximum snout-vent length of +27 mm. in males and 32 mm. in females; dorsum yellowish tan with brown +markings; thighs uniformly yellow, vocal sac in breeding males yellow; +snout truncate in lateral profile; tympanum distinct, usually slightly +smaller than one-half diameter of eye; vocal sac single, median, +subgular; fingers about one-third webbed; toes webbed nearly to bases +of discs, except only to middle of antepenultimate or base of +penultimate phalanx of fourth toe; tarsal fold weak; inner metatarsal +tubercle low, flat, elliptical; axillary membrane present; pupil +horizontally elliptical; palpebral membrane unmarked; cranial elements +reduced in ossification; sphenethmoid small, short; frontoparietal +fontanelle large; tegmen tympani not extensive; quadratojugal greatly +reduced; anterior arm of squamosal extending only about one-fourth +distance to maxillary; posterior arm of squamosal not having bony +connection with prootic; nasals lacking maxillary processes; medial +ramus of pterygoid not having bony attachment to prootic; maxillary, +premaxilary, and prevomerine teeth present; palatine and parasphenoid +teeth absent; Mentomeckelians ossified; tadpoles having xiphicercal +tails with deep caudal fins and terminal mouth lacking teeth; mating +call consisting of one primary note followed by a series of shorter +secondary notes; haploid number of chromosomes, 15 (known only in _H. +microcephala_ and _H. phlebodes_.) + +_Content._--As recognized here the _Hyla microcephala_ group contains +four species, one having two subspecies. An alphabetical list of the +specific and subspecific names that we consider to be applicable to +the _Hyla microcephala_ group are listed below. + + + Names Proposed Valid Names + +_Hyla cherrei_ Cope, 1894 ? = _H. m. microcephala_ +_Hyla microcephala_ Cope, 1886 = _H. m. microcephala_ +_Hyla microcephala_ Boulenger, + 1898 (_nec_ Cope, 1886) = _H. microcephala underwoodi_ +_Hyla microcephala martini_ Smith, 1951 = _H. microcephala underwoodi_ +_Hyla microcephala sartori_ Smith, 1951 = _H. sartori_ +_Hyla phlebodes_ Stejneger, 1906 = _H. phlebodes_ +_Hyla robertmertensi_ Taylor, 1937 = _H. robertmertensi_ +_Hyla underwoodi_ Boulenger, 1899 = _H. microcephala underwoodi_ + + +_Discussion._--The color pattern is the most useful character in +distinguishing the species of the _Hyla microcephala_ group from one +another. Except in _Hyla microcephala_, little geographic variation in +color pattern is noticeable. The features of color pattern that are +helpful in identifying the species are: 1) presence or absence of +lateral dark brown stripe; 2) longitudinal extent and width of lateral +stripe, if present; 3) presence or absence of a narrow white line just +dorsal to the lateral dark stripe; 4) presence or absence of an +interorbital dark mark; 5) the arrangement of dark markings on the +back, either as longitudinal lines or series of dashes, or in the form +of various kinds of transverse markings; 6) presence of dark flecks, +longitudinal line, or transverse marks on shanks. + +Few consistent differences in measurements and proportions exist among +the species (Table 1). The most obvious morphological difference is +that the head is noticeably narrower in _H. robertmertensi_ than in +the other species. _Hyla phlebodes_ is the smallest species; adult +males attain snout-vent lengths of only 23.6 mm. The body is slender +in _H. microcephala_ and _robertmertensi_, slightly wider in +_phlebodes_, and noticeably broader in _sartori_. + +_Distribution._--The composite range of the Middle American frogs of +the _Hyla microcephala_ group includes the lowlands of southern Mexico +and Central America, in some places to elevations of 1200 meters, +southeastward from southern Jalisco and southern Veracruz, excluding +arid regions (northern Yucatan Peninsula, Balsas-Tepalcatepec Basin, +Plains of Tehuantepec, Grijalva Valley, Salama Basin, and upper +Motagua Valley) to the Pacific lowlands and the Cauca and Magdalena +valleys in Colombia. + + +Key to Species and Subspecies + + +1. Lateral dark stripe, bordered above by narrow white line, + extending from snout at least to sacral region 2 + + Lateral dark stripe, if present, not extending posteriorly to + sacral region and not bordered above by narrow white line 4 + +2. Lateral dark stripe continuous to groin; dark flecks or + longitudinal line on shanks; interorbital dark bar absent; + dorsal pattern usually consisting of pair of longitudinal dark + lines or series of dashes 3 + + Lateral dark stripe usually extending only to sacral region; + dark transverse bars on shanks; interorbital bar usually + present; dorsal pattern usually consisting of interconnecting + dark lines, sometimes forming transverse marks + _H. microcephala underwoodi_ + +3. Lateral dark stripe narrow, covering only upper edge of + tympanum; dorsal longitudinal stripes continuous, extending to + vent _H. microcephala microcephala_ + + Lateral dark stripe wide, encompassing entire tympanum; dorsal + markings consisting of longitudinal series of flecks or dashes, + or of two lines, usually not extending to vent _H. robertmertensi_ + +4. Lateral dark stripe indistinct, present only above tympanum and + insertion of arm; dorsal markings consisting of narrow lines + and dashes, sometimes interconnected; transverse bars on shanks + narrow relative to interspaces _H. phlebodes_ + + Lateral dark stripe absent; dorsal markings consisting of two broad + chevron-shaped marks; transverse bars on shanks wide relative to + interspaces _H. sartori_ + + + + +ACCOUNTS OF SPECIES AND SUBSPECIES + + +_Hyla microcephala_ Cope + + +_Diagnosis._--Lateral dark stripe narrow, covering only upper edge of +tympanum, bordered above by narrow white stripe; dorsal pattern +consisting of pair of longitudinal brown lines and no interorbital bar +(eastern populations), or of irregular dark markings forming an X- or +)(-shaped mark in scapular region and an interorbital bar (western +populations). + +_Content._--The populations inhabiting the Pacific lowlands of +southeastern Costa Rica eastward to Colombia are recognized herein as +_Hyla microcephala microcephala_ Cope; the populations in western +Costa Rica northward to Mexico are assigned to _Hyla microcephala +underwoodi_ Boulenger. + +_Distribution._--Southern Veracruz and northern Oaxaca southeastward +through the Atlantic lowlands of Central America to north-central +Nicaragua, thence southeastward on the Pacific lowlands to eastern +Panama, and thence into the Cauca and Magdalena valleys (Caribbean +drainage) of Colombia (Fig. 1). + + + [Illustration: Fig. 1. Map showing locality records for _Hyla + microcephala_.] + + +Table 1.--Variation in Certain Measurements and Properties in the + Hyla microcephala Group. (All Data Based on Adult Males; + Mean and Standard Error of Mean Below Observed Range.) + +======================================================================== + Locality | N | Snout-vent | Tibia length |Foot length| + | | length | ------------ | --------- | + | | (S-V L) | S-V L | S-V L | +------------------------------------------------------------------------ + | _H. m. microcephala_ + | +Panama: Canal Zone | 25 | 21.5-24.1 | 50.2-56.0 | 40.9-46.6 | + | | 22.8+-0.20 | 52.9+-0.37 | 43.5+-0.28 | + | | | | | +Costa Rica: Golfito | 25 | 18.5-24.5 | 49.1-54.4 | 41.8-48.0 | + | | 22.4+-0.27 | 51.6+-0.26 | 45.1+-0.32 | + | + | _H. m. underwoodi_ + | | +Nicaragua: La Cumplida | 25 | 23.0-25.6 | 51.0-55.7 | 41.3-46.5 | + | | 24.1+-0.19 | 52.9+-0.25 | 43.7+-0.25 | + | | | | | +Guatemala: Finca Chama | 25 | 21.8-25.0 | 51.0-57.2 | 41.2-47.8 | + | | 23.5+-0.16 | 54.3+-0.39 | 44.4+-0.30 | + | | | | | +Tabasco: Teapa | 25 | 22.7-25.8 | 48.0-54.5 | 40.7-46.8 | + | | 24.3+-0.14 | 51.5+-0.29 | 43.3+-0.25 | + | | | | | +Oaxaca: Donaji-Sarabia | 25 | 22.1-25.9 | 49.8-55.6 | 40.5-46.6 | + | | 23.8+-0.19 | 52.8+-0.33 | 43.4+-0.27 | + | | | | | +Veracruz: Alvarado | 25 | 21.9-25.4 | 49.6-54.4 | 40.7-47.5 | + | | 24.1+-0.17 | 51.1+-0.28 | 42.6+-0.34 | + | + | _H. robertmertensi_ + | +Guatemala: La Trinidad | 21 | 21.8-24.6 | 47.1-52.8 | 40.9-51.3 | + | | 23.4+-0.15 | 49.9+-0.34 | 43.5+-0.17 | + | | | | | +Chiapas: Acacoyagua | 25 | 21.4-25.7 | 47.8-52.4 | 41.7-46.3 | + | | 24.1+-0.20 | 50.4+-0.45 | 43.9+-0.23 | + | | | | | +Oaxaca: Tapanatepec | 25 | 22.4-26.4 | 44.1-48.3 | 39.1-44.5 | + | | 24.7+-0.18 | 46.4+-0.23 | 41.7+-0.23 | + | + | _H. phlebodes_ + | +Panama: Canal Zone | 25 | 19.6-23.2 | 49.1-56.9 | 41.9-47.1 | + | | 22.2+-0.16 | 52.8+-0.35 | 45.4+-0.26 | + | | | | | +Costa Rica: Turrialba | 25 | 19.7-23.6 | 47.4-55.7 | 38.1-46.4 | + | | 22.0+-0.18 | 51.1+-0.35 | 42.8+-0.38 | + | + | _H. sartori_ + | +Guerrero: Tierra Colorada| 25 | 23.7-26.0 | 47.2-51.4 | 42.4-47.8 | + | | 24.8+-0.13 | 49.6+-0.23 | 45.2+-0.27 | +------------------------------------------------------------------------ +Table 1. (continued) +=============================================================== + Locality | Head length | Head width | Tympanum + | ----------- | ---------- | -------- + | S-V L | S-V L | Eye +--------------------------------------------------------------- + | _H. m. microcephala_ + | +Panama: Canal Zone | 28.5-32.8 | 28.1-30.9 | 44.0-54.1 + | 31.0+-0.22 | 29.4+-0.11 | 49.0+-0.55 + | +Costa Rica: Golfito | 30.2-35.5 | 29.0-32.7 | 40.0-57.8 + | 33.1+-0.25 | 30.8+-0.16 | 48.4+-1.10 + | + | _H. m. underwoodi_ + | +Nicaragua: La Cumplida | 29.7-33.5 | 28.9-31.8 | 42.3-60.0 + | 31.6+-0.19 | 30.4+-0.17 | 49.3+-0.97 + | +Guatemala: Finca Chama | 30.8-35.3 | 29.6-33.6 | 37.5-56.4 + | 33.0+-0.16 | 31.3+-0.36 | 45.2+-0.89 + | +Tabasco: Teapa | 29.5-33.0 | 28.7-31.8 | 40.7-53.8 + | 31.7+-0.17 | 30.3+-0.16 | 45.5+-0.38 + | +Oaxaca: Donaji-Sarabia | 30.4-34.8 | 28.9-32.6 | 37.0-54.1 + | 32.8+-0.19 | 30.8+-0.17 | 45.1+-0.76 + | +Veracruz: Alvarado | 29.9-33.8 | 29.1-32.9 | 40.7-53.8 + | 31.4+-0.18 | 30.5+-0.17 | 46.6+-0.65 + | + | _H. robertmertensi_ + | +Guatemala: La Trinidad | 30.0-33.3 | 27.3-29.8 | 44.4-50.0 + | 31.3+-0.20 | 28.5+-0.23 | 47.4+-0.46 + | | | +Chiapas: Acacoyagua | 29.1-32.7 | 26.0-30.3 | 42.8-53.8 + | 31.2+-0.29 | 28.1+-0.20 | 46.5+-0.50 + | | | +Oaxaca: Tapanatepec | 26.1-30.4 | 25.4-28.1 | 45.8-58.3 + | 28.4+-0.16 | 26.8+-0.14 | 52.9+-0.77 + | + | _H. phlebodes_ + | +Panama: Canal Zone | 33.6-37.4 | 32.3-36.0 | 37.9-46.4 + | 34.8+-0.18 | 33.8+-0.18 | 41.6+-0.49 + | | | +Costa Rica: Turrialba | 32.6-35.9 | 30.5-35.0 | 35.7-48.2 + | 34.1+-0.16 | 32.9+-0.17 | 40.1+-0.53 + | + | _H. sartori_ + | +Guerrero: Tierra Colorada| 29.4-31.8 | 28.9-31.0 | 42.3-52.0 + | 30.6+-0.13 | 30.0+-0.12 | 47.4+-0.59 +--------------------------------------------------------------- + + + + +_Hyla microcephala microcephala_ Cope + + + _Hyla microcephala_ Cope, Proc. Amer. Philos. Soc., 23:281, February + 11, 1886 [Syntypes.--USNM 13473 (2 specimens, now lost) from + Chiriqui, Panama; Mr. MacNeil collector]; Bull. U.S. Natl. Mus., + 32:14, 1887. Guenther, Biologia-Centrali Americana, Reptilia and + Batrachia, p. 265, June, 1901. Dunn, Occas. Papers Boston Soc. + Nat. Hist., 5:413, October 10, 1931; Occas. Papers Boston Soc. + Nat. Hist., 8:72, June 7, 1933. Stebbins and Hendrickson, Univ. + California Publ. Zool., 56:524, February 17, 1959. Fouquette, + Evolution, 14:484, December 16, 1960. Busack, Copeia, 2:371, + June 21, 1966. + + ? _Hyla cherrei_ Cope, Proc. Acad. Nat. Sci. Philadelphia, 1894, + p. 195, 1894 [Holotype.--location unknown, apparently lost; + type-locality: "Alajuela, Costa Rica;" R. Alfaro collector]. + Guenther, Biologia Centrali-Americana: Reptilia and Batrachia, + p. 264, June, 1901. Taylor, Univ. Kansas Sci. Bull., 35:846, + July 1, 1952. + + _Hyla underwoodi_, Ruthven, Misc. Publ. Mus. Zool., Univ. Michigan, + 8:55, September 15, 1922. Barbour, Proc. New England Zool. Club, + 10:31, March 2, 1928. + + _Hyla microcephala microcephala_, Smith, Herpetologica, 7:185, + December 31, 1951. Taylor, Univ. Kansas Sci. Bull., 39:23, + November 18, 1958. + + +_Diagnosis._--Brown lateral stripe narrow, extending from nostril +along canthus, along upper edge of tympanum to groin, bordered above +by narrow white line; pair of dark brown longitudinal lines on dorsum +extending to vent; shanks having dark longitudinal line or flecks, no +transverse bars; interorbital dark mark lacking. + +_Description and Variation._--The color pattern is nearly constant. Of +103 males from the Canal Zone, all lack an interorbital dark bar, and +all have a dark longitudinal line on the dorsal surface of the shank +and a narrow lateral dark stripe, bordered above by a narrow white +line, extending to the groin. The longitudinal dark lines on the +dorsum are continuous to the groin in 95 specimens and fragmented in +two specimens. In two others the lines converge and fuse in the +scapular region, and in four specimens auxiliary, fragmented lines are +present dorsolaterally. + +In all specimens from southeastern Costa Rica (Golfito, Palmar Sur, +and Villa Neilly) the pattern is constant, except that in about 10 per +cent of the specimens the longitudinal line on the dorsal surface of +the shank is replaced by a row of brown flecks. + +Of the limited number of Colombian specimens examined, all are +patterned normally, except three from Sautata, Choco, three from +Curumani, and three from Arcataca, Magdalena, which have flecks on the +dorsal surfaces of the shanks, and one from Espinal, Tolima, which has +no markings on the shanks. + +When active at night most individuals are pale yellowish tan dorsally; +the white dorsolateral line is noticeable, but the brown lateral +stripe, dorsal brown lines, and lines on shanks are so pale that often +they are barely discernible. By day the dorsum changes to tan or pale +reddish brown; the stripes are dark brown, and the dorsolateral stripe +that is white at night becomes creamy yellow (Pl. 13). Small brown +flecks are present on the dorsum of most individuals. The venter +always is white, and the iris is pale bronze with a brown tint +immediately anterior and posterior to the pupil. In breeding males the +vocal sac is pale yellow. + +_Tadpoles._--Tadpoles of this species have been found in weed-choked +ponds in eastern Panama Province. The following description is based +on KU 104097, a specimen in developmental stage 34 (Gosner, 1960). + +Total length, 20.5 mm.; body length, 8.2 mm.; body slightly wider +than deep; snout pointed; nostrils large, situated dorsally, much +closer to snout than eyes, directed anteriorly; eyes moderately +small, situated dorsolaterally and directed laterally; spiracle +sinistral, located just posteroventral to eye; anal tube dextral. +Tail xiphicercal; caudal musculature moderately deep, becoming +slender posteriorly, extending beyond caudal fin; fins deepest at +about one-third distance from body to tip of tail; dorsal fin +extending onto body, deeper than deepest part of caudal +musculature; ventral fin slightly shallower than musculature. +Mouth small, terminal, lacking teeth and fringing papillae, but +having finely serrate beaks. In preservative, top of head pale +brown; dark stripe from tip of snout through eye to posterior edge +of body, narrowing to thin line on proximal one-fourth of tail; +venter white; tail creamy tan with fine black flecks most numerous +posteriorly; posterior two-thirds of fins edged with black. In +life, top of head yellowish tan; lateral stripe brown; belly +white; anterior half of tail lacking pigment; posterior half deep +orange; iris pale bronze (Pl. 15). + +_Remarks._--Evidence for intergradation of _Hyla microcephala_ with +_H. underwoodi_ is provided by four specimens [USC 818 (2), 6081-2] +from 6.1 kilometers northeast of the mouth of the Rio Tarcoles, and +nine specimens [USC 8254 (2), 8255, 8256 (4), 8258 (2)] from Parrita, +both in Puntarenas Province, Costa Rica. These localities lie about +two-thirds the distance from the northwesternmost locality for _H. +m. microcephala_ (Palmar Sur) to the southeasternmost locality for +_H. m. underwoodi_ (Barranca). Although in most aspects of coloration +the frogs are more nearly like _H. m. underwoodi_ than _H. m. +microcephala_, some specimens have longitudinal lines on their shanks, +such as are characteristic of _H. m. microcephala_. The dorsal pattern +varies from nearly complete longitudinal lines to broken lines, fused +into an X-shaped scapular mark or not. + +As noted by Rivero (1961:135), _Hyla microcephala_ seems to be closely +related to _Hyla misera_ Werner, a species having a wide distribution +east of the Andes in South America. Despite the similarity in color +pattern, size, and structure, we are reluctant to place the two taxa +in the same species until data on coloration in life, mating calls, +and life history are available for _Hyla misera_ and compared with +those of _Hyla microcephala_. + +The status of Cope's _Hyla cherrei_ is questionable. Since the type, +the only specimen ever referred to the species, apparently is lost, +the only extant information regarding the taxon is contained in the +original description (Cope, 1894). There the species was characterized +as having a narrow dorsolateral white stripe and lacking pigment on +the upper arms and thighs. These characteristics of the color pattern +combined with the statements "vomerine teeth few, opposite the middle +of the very large choanae" and "tympanic drum distinct, one half the +area of eye" serve to distinguish _H. cherrei_ from all other Costa +Rican hylids, except _H. m. microcephala_ and _H. m. underwoodi_. No +statements in the type description will definitely associate _cherrei_ +with one or the other of these subspecies. Since it seems obvious that +_H. cherrei_ can be associated with _H. microcephala_, we prefer to +place the name in the synonymy of the nominate subspecies, thereby +preserving the commonly used name _H. underwoodi_ (Boulenger, 1899) as +a subspecies of _H. microcephala_. + +_Distribution._--_Hyla microcephala microcephala_ inhabits coastal +lowlands from the area of Golfo Dulce (apparently absent from the +Osa Peninsula) in southeastern Costa Rica eastward in Panama, +including the Azuero Peninsula to northern Colombia and thence +southward in the valleys of the Rio Cauca and Rio Magdalena in +Colombia (Fig. 1). Except for the central area of the Canal Zone +the subspecies is unknown from the Caribbean drainage in Central +America, but in Colombia the subspecies occurs only in the +Caribbean drainage. In Central America this frog occurs mostly on +the coastal lowlands; the highest recorded elevation is 560 meters +at El Valle, Cocle, Panama. Throughout most of its range _Hyla +microcephala microcephala_ occurs in disturbed habitats--cut-over +forests, secondary growth, and pastureland. It does not seem to be +an inhabitant of either primary forest or of _Curatella_-savanna. + +_Specimens examined._--522, as follows: +Costa Rica+: Puntarenas: +Golfito, KU 32172-207; 3 km. E Golfito, KU 86399, USC 2757-8; +Palmar Sur, KU 64591-608, USC 2650 (14), UU 3907-32; *1.5-2.5 km. +ESE Palmar Sur, KU 68293-7 (skeletons), 93957-62; Parrita, USC +8254 (2), 8255, 8256 (4), 8258 (2) [intergrades with _H. m. +underwoodi_]; 3 km. NW Piedras Blancas, KU 103689; 6.1 km. NE +mouth of Rio Tarcoles, USC 818 (2), 6081-2 [intergrades with _H. +m. underwoodi_]; Villa Neilly, USC 2651; *1-5 km. WNW Villa +Neilly, USC 6182-4, 8003 (4), 8031 (3), 8032; *10.5 km. WNW Villa +Neilly, KU 64609-27, 68398 (eggs). + ++Panama+: Canal Zone: Albrook Air Base, TNHC 23389, 23497; Balboa, +ANSP 19555-6; *Fort Clayton, UIMNH 42008-12; *2.8 km. SW Fort +Kobbe, KU 96015-25; *Frijoles, MCZ 19208; *Bamboa, MCZ 21507; *8.3 +km. N Gatun Locks, TNHC 23441; *Juan Diaz, MCZ 13747; *Juan Mina, +AMNH 55436-7, ANSP 21811-2, UMMZ 126734, 126735 (6), UU 3900-6; +*8-14 km. N Miraflores Locks, TNHC 23374-88, 23390-409, 23411-38, +23440, 23442-60, 23462-76; 23478-83, 23492, 23555-60, 23562-76; +*Rio Chagres, AMNH 55430, 55439; *Rio Cocoli, 3.5 km. N Miraflores +Locks, TNHC 23410; *Summit, ANSP 23365-71, FMNH 22966-9, KU +97783-87. Chiriqui: 5.5 km. E Concepcion, AMNH 69772; *14.4 km. E +Concepcion, AMNH 69773-8; 2 km. S David, AMNH 69779; *Progreso, +UMMZ 58252, 58253 (2), 58254, 58436; Rio Gariche, 8.3 km. ESE Paso +Canoas, KU 103065-8. Cocle: 1 km. SE El Cano, KU 103042-51; El +Valle de Anton, AMNH 59614-18 (10), 69785, ANSP 23502-5, KU +77201-14, MVZ 66578-83, UIMNH 46532. Colon: Cement Plant, +Transisthmian Highway, FMNH 60394-5. Darien: El Real, KU 80454-5, +103052-64, UMMZ 125036 (10), USNM 140567-8; Rio Canclon at Rio +Chucunaque, UMMZ 125035; *Rio Chucunaque, near Yavisa, AMNH 59523. +Los Santos: Tonosi, KU 101606-9. Panama: 5 km. S Bejuco, AMNH +69782; 3 km. W Chepo, KU 77172-4, 104097-8 (tadpoles); *6 km. WSW +Chepo, KU 77175; *Chico, Rio La Jagua, USNM 129070; *La Joya, +Cacora, ANSP 25129-33; Madden Dam, FMNH 67819; Nueva Gorgona, AMNH +69780-1; *1.6 km. W Nueva Gorgona, AMNH 69783-4; 1.5 km. W Pacora, +77176-200; *Rio La Laja, near Chame, ANSP 21845; *Rio Tapia, MCZ +10048; *Tapia, AMNH 18930, 18950, 18952-3; *18 km. E Tocumen, MVZ +78662. + ++Colombia+: Choco: Sautata, Atrato, FMNH 74918 (2), 74919. +Magdalena: Aracataca, ANSP 19755-7; Curumani, MCZ 21465-74, UIMNH +28855; UMMZ 90168, USNM 118247; El Banco, Rio Magdalena, ANSP +25061; Fundacion, UMMZ 48281-2. Tolima: Espinal, MCZ 15068; +Mariquita, FMNH 81822-3. Valle: Sevilla, MCZ 13751-3. + + + + +_Hyla microcephala underwoodi_ Boulenger + + + _Hyla microcephala_ Boulenger, Proc. Zool. Soc. London, p. 481, + October 1, 1898 [Syntypes.--BMNH 94. 11. 1532-33 from Bebedero, + Guanacaste Province, Costa Rica; C. F. Underwood collector] (not + _Hyla microcephala_ Cope, Proc. Amer. Philos. Soc., 23:281, + February 11, 1886, from Chiriqui, Panama). + + _Hyla underwoodi_ Boulenger, Ann. Mag. Nat. Hist., ser. 7, 3:277, + April, 1899 (substitute name for _Hyla microcephala_ Boulenger, + preoccupied). Guenther, Biologia-Centrali Americana, Reptilia and + Batrachia, p. 278, September, 1901. Dunn and Emlen, Proc. Acad. + Nat. Sci. Philadelphia, 84:25, March 22, 1932. Stuart, Misc. Publ. + Mus. Zool., Univ. Michigan, 29:39, October 1, 1935. Taylor, Proc. + Biol. Soc. Washington, 50:44, April 21, 1937. Stuart, Occas. + Papers Mus. Zool., Univ. Michigan, 471:15, May 17, 1943. Taylor + and Smith, Proc. U. S. Natl. Mus., 95:586, June 30, 1945. Stuart, + Misc. Publ. Mus. Zool., Univ. Michigan, 69:35, June 12, 1948. + Smith and Taylor, Bull. U. S. Natl. Mus., 194:85, June 17, 1948; + Univ. Kansas Sci. Bull., 33:316, March 20, 1950. Stuart, Contr. + Lab. Vert. Biol., Univ. Michigan, 45:48, May, 1950. Taylor, Univ. + Kansas Sci. Bull., 35:891, July 1, 1952; Univ. Kansas Sci. Bull., + 39:25, November 18, 1958. + + _Hyla phlebodes_, Cole and Barbour, Bull. Mus. Comp. Zool., 50:154, + November, 1906. Kellogg, Bull. U. S. Natl. Mus., 160:172, + March 31, 1932. + + _Hyla microcephala martini_ Smith, Herpetologica, 7:187, December + 31, 1951 [Holotype.--UIMNH 20965 from Encarnacion, Campeche, + Mexico; H. M. Smith collector]. Stuart, Contr. Lab. Vert. Biol., + Univ. Michigan, 68:46, November, 1954. Fugler and Webb, + Herpetologica, 13:105, July 10, 1957. Stuart, Contr. Lab. Vert. + Biol., Univ. Michigan, 75:17, June, 1958. Neill and Allen, Publ. + Research Div., Ross Allen's Reptile Inst., 2:26, November 10, + 1959. Duellman, Univ. Kansas Publ., Mus. Nat. Hist., 13:62, + August 16, 1960. Stuart, Herpetologica, 17:74, July 11, 1961. + Hensley and Smith, Herpetologica, 18:70, April 9, 1962. Stuart, + Misc. Publ. Mus. Zool., Univ. Michigan, 122:36, April 2, 1963. + Holman and Birkenholz, Herpetologica, 19:144, July 3, 1963. + Duellman, Univ. Kansas Publ., Mus. Nat. Hist., 15:225, October 4, + 1963; Univ. Kansas Publ., Mus. Nat. Hist., 15:588, June 22, 1965. + + _Hyla microcephala underwoodi_, Smith, Herpetologica, 7:188, + December 31, 1951. + + +_Diagnosis._--Brown lateral stripe narrow, extending to groin or +only to sacral region, bordered above by narrow white line; dorsal +pattern bold, consisting of X- or )(-shaped mark in scapular +region or pair of interconnected dark lines on back; interorbital +dark mark usually present; shanks usually having dark transverse +bars. + +_Description and Variation._--The dorsal color pattern is highly +variable. The various permutations of the X-shaped scapular mark +and dark sacral marks differ proportionately in different samples. +The variation in color pattern in 12 samples is summarized in +Table 2. In samples from the southern part of the range (southern +Nicaragua and Guanacaste Province, Costa Rica) more (40-93%) +individuals have the lateral stripes extending to the groin than +in northern samples (0-42%) from southern Mexico and Guatemala. +Likewise, the percentage of specimens lacking bars on the shanks and +a dark interorbital bar is higher in the Costa Rican samples than +elsewhere in the range. The X- or )(-shaped scapular markings and +/\- or / \-shaped sacral markings are most prevalent in northern +samples, whereas to the south the dorsal markings are more commonly +arranged in a pattern of paired lines, which usually are discontinuous +and usually extend posteriorly only to the sacral region. Thus, the +color pattern in _H. m. underwoodi_ in the southern part of its range +shows trends towards the pattern characteristic of _H. m. +microcephala_. Intergrades between these two subspecies have +been discussed in the account of the nominate subspecies. + + + + + Table 2.--Variation in Color Pattern in Hyla microcephala underwoodi +========================================================================== + Population | N | Shanks || Interorbital || Dorsolateral | + | | || bar || stripe | + | |-------------||----------------||--------------| + | | Bars |Flecks|| Present| Absent|| Groin| Sacrum| +-------------------------------------------------------------------------- +Oaxaca: | 27 | 22 | 5 || 27 | 0 || 0 | 27 | + Donaji-Sarabia | | | || | || | | + | | | || | || | | +Tabasco: | 55 | 46 | 9 || 55 | 0 || 0 | 55 | + Teapa-Villahermosa| | | || | || | | + | | | || | || | | +Guatemala: | 51 | 51 | 0 || 51 | 0 || 17 | 34 | + La Libertad | | | || | || | | + | | | || | || | | +Guatemala: | 32 | 32 | 0 || 32 | 0 || 0 | 32 | + Finca Chama | | | || | || | | + | | | || | || | | +Guatemala: | 31 | 31 | 0 || 31 | 0 || 14 | 17 | + Puerto Barrios | | | || | || | | + | | | || | || | | +Honduras: | 13 | 13 | 0 || 13 | 0 || 9 | 4 | + Lago Yojoa | | | || | || | | + | | | || | || | | +Nicaragua: | 56 | 44 | 12 || 54 | 2 || 13 | 43 | + La Cumplida | | | || | || | | + | | | || | || | | +Nicaragua: | 10 | 10 | 0 || 10 | 0 || 8 | 2 | + Tipitapa | | | || | || | | + | | | || | || | | +Nicaragua: | 10 | 10 | 0 || 10 | 0 || 8 | 2 | + Santo Thomas | | | || | || | | + | | | || | || | | +Costa Rica: | 12 | 0 | 12 || 6 | 6 || 7 | 5 | + Tenorio-Tilaran | | | || | || | | + | | | || | || | | +Costa Rica: | 38 | 21[A]| 15 || 34 | 4 || 25 | 13 | + Las Canas-Liberia | | | || | || | | + | | | || | || | | +Costa Rica: | 32 | 26 | 6 || 29 | 3 || 30 | 2 | + Esparta | | | || | || | | +-------------------------------------------------------------------------- + +========================================================================== + Population | Scapular markings || Sacral | + | || markings | + |----------------------------||----------------------| + | X | )( | ][ | Other || /\ | / \ | Other | +-------------------------------------------------------------------------- +Oaxaca: | 23 | 4 | 0 | 0 || 7 | 6 | 14 | + Donaji-Sarabia | | | | || | | | + | | | | || | | | +Tabasco: | 53 | 2 | 0 | 0 || 19 | 11 | 23 | + Teapa-Villahermosa| | | | || | | | + | | | | || | | | +Guatemala: | 45 | 6 | 0 | 0 || 16 | 14 | 21 | + La Libertad | | | | || | | | + | | | | || | | | +Guatemala: | 32 | 0 | 0 | 0 || 26 | 2 | 4 | + Finca Chama | | | | || | | | + | | | | || | | | +Guatemala: | 23 | 0 | 4 | 4 || 6 | 4 | 21 | + Puerto Barrios | | | | || | | | + | | | | || | | | +Honduras: | 3 | 2 | 3 | 5 || 2 | 1 | 10 | + Lago Yojoa | | | | || | | | + | | | | || | | | +Nicaragua: | 11 | 35 | 8 | 2 || 0 | 19 | 37 | + La Cumplida | | | | || | | | + | | | | || | | | +Nicaragua: | 0 | 5 | 3 | 2 || 0 | 3 | 7 | + Tipitapa | | | | || | | | + | | | | || | | | +Nicaragua: | 3 | 0 | 7 | 0 || 0 | 5 | 5 | + Santo Thomas | | | | || | | | + | | | | || | | | +Costa Rica: | 0 | 0 | 12 | 0 || 0 | 0 | 12 | + Tenorio-Tilaran | | | | || | | | + | | | | || | | | +Costa Rica: | 0 | 11 | 19 | 8 || 0 | 0 | 38 | + Las Canas-Liberia | | | | || | | | + | | | | || | | | +Costa Rica: | 0 | 0 | 14 | 18 || 0 | 0 | 32 | + Esparta | | | | || | | | +-------------------------------------------------------------------------- + +[Footnote A: Longitudinal stripes present in two specimens.] + + + +When this frog is active at night its dorsum is pale yellow; faint +flecks are present in some individuals. The white dorsolateral line +usually is evident in the tympanic region, but in many individuals a +dorsal pattern of lines and other marks is not evident. By day the +dorsum changes to yellowish tan or pale brown with dark brown or +reddish brown markings (Pl. 13). The venter is white, and the vocal +sac in breeding males is yellow. The iris is pale bronze with a brown +tint anterior and posterior to the pupil. + +_Remarks._--_Hyla microcephala underwoodi_ has had a confused +nomenclatural history. The taxon was first named _Hyla microcephala_ +by Boulenger (1898); this name was preoccupied by _Hyla microcephala_ +Cope (1886). Cole and Barbour (1906) and Kellogg (1932) used the name +_Hyla phlebodes_ Stejneger (1906) for specimens of this frog from +Mexico. Dunn (1931, 1933, 1934) applied the name _Hyla underwoodi_ to +Panamanian specimens that we identify as _Hyla phlebodes_. Smith +(1951) named _Hyla microcephala martini_ from southern Mexico and +Guatemala and considered the northern populations to represent a +subspecies distinct from the Costa Rican _Hyla microcephala +underwoodi_, despite the fact the Stuart (1935:39) stated that +comparisons of specimens from El Peten, Guatemala, with the holotype +of _Hyla underwoodi_ showed only trivial differences. + +Much of the confusion regarding the name _Hyla underwoodi_ stems from +the illustration given by Boulenger (1898:pl. 39, fig. 3) and +reproduced by Taylor (1952:892), which shows a frog having a unicolor +dorsum, dorsolateral white lines, and dark flanks. This pattern is in +marked contrast to the pattern seen in most preserved specimens, which +have the dorsum variously marked by dark brown lines or irregular +marks. Smith (1951:185), in his description of _Hyla microcephala +martini_ from southern Mexico, considered _H. underwoodi_ to be a +subspecies of _H. microcephala_ that lacked dorsal dark markings. + +Data accumulated in 1961 through field studies by the senior author at +the type locality, Bebedero, and other localities in Guanacaste and +Puntarenas provinces in Costa Rica provide a reasonable explanation of +the differences in color pattern. As noted in the preceding description +of this subspecies, at night the dorsal markings are not evident in +many living individuals, whereas by day the dorsal markings are +prominent. Most collectors prepare their specimens by day; consequently +the majority of specimens have a pronounced dorsal pattern. Of the +frogs collected in Costa Rica in 1961, some specimens were preserved +at night; others from the same series were preserved by day. The +differences are striking. In those preserved at night, dorsal markings +are faint, if present at all. Some specimens closely match the figure +given by Boulenger (1898). + +It is extremely doubtful if the frog described and illustrated by +Boulenger could be associated with either _Hyla phlebodes_ or _H. +microcephala microcephala_. Individuals of the former species lack +a dorsolateral white line and always have some dorsal markings +evident at night; furthermore, _H. phlebodes_ is not known to occur +on the Pacific lowlands. _Hyla microcephala microcephala_ occurs +farther southeast. Since there is no reason to doubt the type locality +of _H. underwoodi_, since specimens from the area around the type +locality that have been preserved at night are like the holotype in +pattern, and since the characteristics of the populations of the frogs +in Guanacaste are the same as, or gradually blend into those of, +populations in northern Central America and southern Mexico, the +frogs from throughout the entire range can be referred to one taxon, +the earliest name for which is _Hyla underwoodi_ Boulenger, which +herein is considered to be a subspecies of _H. microcephala_ Cope. + +_Distribution._--_Hyla microcephala underwoodi_ inhabits the +Atlantic slopes and lowlands from southern Veracruz and extreme +northern Oaxaca eastward across the base of the Yucatan Peninsula +(possibly the species is extant in the northern part of the +peninsula) to British Honduras and thence southeastward through +the Caribbean lowlands and interior valleys in Honduras to central +Nicaragua, where it apparently avoids the forested Caribbean +lowlands and the dry Pacific lowlands of northwestern Nicaragua, +but in the vicinity of Managua invades the Pacific lowlands and +continues southward into northwestern Costa Rica as far as the +Puntarenas Peninsula (Fig. 1). In Mexico and Guatemala the species +has not been taken at elevations of more than 350 meters, whereas +farther south it occurs at higher elevations--780 meters at +Silencio, Costa Rica, 830 meters on Montana de Guaimaca, Honduras, +960 meters at Finca Tepeyac, Nicaragua, and 1200 meters at Finca +Venecia, Nicaragua. + +_Specimens examined._--1270, as follows: +Mexico+: Campeche: Balchacaj, +FMNH 100406, UIMNH 20944-6; Encarnacion, FMNH 27069-70, 75784, +MCZ 28360, 29637, UIMNH 20948-58, 20965, USNM 134264-5; Escarcega, +UMMZ 122999; *7.5 km. W Escarcega, KU 71229-43; Laguna Alvarado, 65 km. +S Xpujil, KU 75084-9; Pacaitun, Rio Candelaria, FMNH 83118-20; +*Tres Brazos, FMNH 113101-22, UIMNH 20947; 10 km. W Xpujil, KU 75082-3. +Chiapas: Palenque, UIMNH 47984, 49139-50, USNM 114973-8. Oaxaca: *5 km. +N Chiltepec, KU 87015-23; 3 km. N Donaji UMMZ 115249 (9); *3.7 km. +N Donaji, UMMZ 115250 (5); *43 km. N Matias Romero, UIMNH 42550-68; +*3.5 km. N Palomares, TNHC 25185, 25321-31, 25341-68; 4.6 km. N +Sarabia, UMMZ 115247 (2); *6.1 km. N Sarabia, UMMZ 115248 (11), *3 km. +N Tolocita, KU 39655; Tuxtepec, KU 87024-40. Tabasco: 24 km. N +Frontera, MCZ 35665-70; 0.8 km. E Rio Tonola, TNHC 25189; Teapa, UMMZ +119218 (4); *2.7 km. N Teapa, UMMZ 119216 (4); *10 km. N Teapa, UMMZ +119217 (6); *11.5 km. N Teapa, UMMZ 119219; *15.2 km. N Teapa, UMMZ +119220 (4); *17.6 km. N Teapa, UMMZ 119221 (12), 3.3 km. S Villahermosa, +UMMZ 119215 (12), *17.6 km. S Villahermosa, UMMZ 119214 (12). +Veracruz: 2.1 km. N Acayucan, UIMNH 42547-9; *6.4 km. NW Acayucan, +UMMZ 115254 (14); 1.6 km. ESE Alvarado, UMMZ 115258 (39); *2.4 km. ESE +Alvarado, UMMZ 115251 (2); *4.5 km. S Aquilera, UMMZ 115252 (21); +*8 km. SW Coatzacoalcos, UMMZ 119213 (10); 2.2 km. E Cosoleacaque, +UMMZ 119222 (26); 10 km. SE Hueyapan, UMMZ 115255; 0.8 km. S Lerdo de +Tejada, UMMZ 122778; *3.6 km. NE Minatitlan, TNHC 25150-2; 1.9 km. S +Naranja, UMMZ 115253 (3); 4.5 km. NE Novillero, UMMZ 115256; San Andres +Tuxtla, FMNH 113124-8, UIMNH 20942-3. Yucatan: Chichen-Itza, FMNH 36570, +MCZ 2463 (2). + ++British Honduras+: Cayo: 6.2 km. S El Cayo, MCZ 37885-92. Stann Creek: +Stann Creek, FMNH 49068. + ++Guatemala+: Alta Verapaz: 28.3 km. N Campur, KU 64578-90; Chinaja, +KU 57425; Cubilquitz, UMMZ 90887, 90888 (4); Finca Chama, UMMZ 90879 +(13), 90880 (4), 90881, 90882 (28), 90883 (12), 90884 (46), 90885 (39), +90886 (20); *Finca Tinaja, BYU 16032; Panzos, UMMZ 90889 (2). +Chiquimula: Chiquimula, UMMZ 98113; 2 km. N Esquipulas, UMMZ 106844. +El peten: La Libertad, KU 57447-97, 59907-11 (skeletons), MCZ 21461, +UMMZ 75332 (13), 75333 (11), 75334 (14), 75335 (10); Piedras Negras, +FMNH 113123, UIMNH 20966; *5 km. S Piedras Negras, USNM 114951-72; +Tikal, UMMZ 117981 (2); Toocog, 15 km. SE La Libertad, KU 57426-46. El +Quiche: Finca Tesoro, UMMZ 89165 (5). Huehuetenango: Finca San Rafael, +16 km. SE Barillas, FMNH 40917-9. Izabal: Puerto Barrios, FMNH +20004-7; 8 km. S Puerto Barrios, KU 57507-37, 59991 (eggs), 59992 +(tadpoles); Quirigua, CAS 69657-701; 2.5 km. NE Rio Blanco, KU 57539; +San Felipe, FMNH 35065. Zacapa: 14 km. ENE Mayuelas, KU 57502-6; 8 km. +ENE Rio Hondo, KU 57498-501. + ++Honduras+: Atlantidad: La Ceiba, UMMZ 91948 (2), USNM 117593-600; +Lancetilla, MCZ 17981. Cortes: Lago Yojoa, AMNH 54917-9, 54957, 55134, +KU 64563-77. El Paraiso: Valle de Jamastran, AMNH 54807-12. +Francisco-Moranza: El Zamorano, AMNH 54873-81, KU 103223, UMMZ 123101; +Montana de Guaimaca, AMNH 54900-4 (8); Ranch San Diego, 19 km. SW +Guaimaca, AMNH 53939. Itibuca: Vieja Itibuca, AMNH 54912-3. + ++Nicaragua+: Chontales: 3 km. SW Santo Tomas, KU 64770-9, 68308 +(skeleton). Esteli: Finca Venecia, 7 km. N, 16 km. E Condega, KU +85296; 2.4 km. N Esteli, MCZ 28933-7. MANAGUA: 12-13 km. E Managua, +KU 85297-301; *10 km. SW Tipitapa, UMMZ 119977 (10). Matagalpa: *Finca +Tepeyac, 10.5 km. N, 9 km. E Matagalpa, KU 85302-3; Hacienda La +Cumplida, KU 64780-96, 68309-11 (skeletons), UMMZ 116482 (8), 116483 +(23), 116484 (3), 116485 (5), 119984 (3). Rivas: *Finca Amayo, 13 km. +S, 14 km. E Rivas, KU 85304-7; 16 km. S Rivas, MCZ 29011-7; *20.5 km. +SE Rivas, KU 85308-10; 5 km. SE San Pablo, KU 43111-4. + ++Costa Rica+: Guanacaste: Arenal, USC 6254 (2); *3 km. W Bagaces, USC +7019 (10); *3 km. NE Boca del Barranca, USC 8017 (21), *Finca San +Bosco, USC 6272 (6), 6276 (3); *Guayabo de Bagaces, USC 7022 (4), 7023 +(3), 7025; 12 km. S La Cruz, USC 8091 (2); *Laguna Arenal, USC 6262; +*27 km. N Las Canas, USC 8171 (6); *16 km. E Las Canas, KU 102252-8; +16 km. SSE Las Canas, KU 65090-5; *20 km. SE Las Canas, KU 102251; +Liberia, KU 30827-39; *7.3 km. N Liberia, USC 8096 (4); *10 km. N +Liberia, USC 8085 (9); *7.5 km. SE Liberia, KU 65102-8, 68621-2 +(skeletons); *14.7 km. S Liberia, USC 8238 (3); *4 km. W Liberia, KU +36847-57; 2 km. S Nicoya, USC 8230; *3-10 km. ESE Playa del Coco, USC +8012 (16), 8137 (14); *21.6 km. ESE Playa del Coco, USC 8138 (13); +*Penas Blancas, KU 102247-50; *Rio Bebedero, 5 km. S Bebedero, KU +65089; *Rio Higueron, USC 7168 (2); Santa Cruz, USC 8232 (2); +*Silencio, USC 6248; *Tenorio, KU 32313; Tilaran, KU 36858-60; *2 km. +E Tilaran, KU 86403, *5 km. NE Tilaran, KU 36840-6 USC 6269. +Puntarenas: Barranca, KU 32305-12, *5 km. WNW Barranca, UMMZ 119976 +(2); *10 km. E Esparta, KU 86400-2; 1 km. WNW Esparta, KU 65101; *4 km. +WNW Esparta, KU 65088; *10 km. WNW Esparta, KU 65063-87, 68616-20 +(skeletons); *12 km. WNW Esparta, KU 65096-100, USC 8251; 21.8 km. +W San Ramon, USC 8242 (15). + + + + ++Hyla robertmertensi+ Taylor + + + _Hyla robertmertensi_ Taylor, Proc. Biol. Soc. Washington, 50:43, + April 21, 1937 [Holotype.--CNHM 100096 (formerly EHT-HMS 2270) + from Tapachula, Chiapas, Mexico; H. M. Smith and E. H. Taylor + collectors]. Smith and Taylor, Bull. U. S. Natl. Mus., 194:84, + June 17, 1948; Univ. Kansas Sci. Bull., 33:326, March 20, 1950. + Mertens. Senckenbergiana, 33:170, June 15, 1952; + Senckenbergischen Naturf. Gesell., 487:30, December 1, 1952. + Stuart, Contr. Lab. Vert. Biol., Univ. Michigan, 68:47, + November, 1954. Duellman, Univ. Kansas Publ., Mus. Nat. Hist., + 13:63, August 16, 1960. Duellman and Hoyt, Copeia, 1961 (2): 417, + December 19, 1961. Porter, Herpetologica, 18:168, October 17, + 1962. Stuart, Misc. Publ. Mus. Zool., Univ. Michigan, 122:36, + April 2, 1963. Duellman and Trueb, Univ. Kansas Publ., Mus. Nat. + Hist., 17:348, July 14, 1966. + + +_Diagnosis._--Brown lateral stripe wide, including loreal region and +entire tympanum, extending to groin, bordered above by narrow white +line; dorsum unicolor or with pair of dark lines (or rows of dashes) +usually extending only to the sacral region; shanks having dark flecks, +no transverse bars; interorbital bar lacking. + +_Description and Variation._--Males attain a maximum snout-vent length +of 26.4 mm. in Oaxaca, whereas in a sample from Acacoyagua, Chiapas, +the largest male has a snout-vent length of 25.7 mm., and from La +Trinidad, Guatemala, 24-6 mm. Specimens from the western part of the +range (eastern Oaxaca) have slightly smaller heads and proportionately +larger tympani than the more eastern populations (Table 1). + +The color pattern shows little variation, except in the nature of the +dorsal markings. In a few specimens from throughout the range, but +especially in the eastern part of the range, the dorsum lacks markings +between the dorsolateral white lines. In most specimens the dorsal +pattern consists of flecks or dashes arranged in two parallel +longitudinal rows, and in some specimens the marks are fused into +parallel lines. Small brown flecks are present on the dorsal surfaces +of the shanks; in some specimens these flecks tend to form a +longitudinal stripe on the shank. An interorbital dark mark is +invariably absent. + +When active at night _Hyla robertmertensi_ is pale yellow above with a +white dorsolateral line and pale brown lateral stripe; the dorsal +markings are faint. By day the dorsum is yellowish tan with brown +markings. The dorsolateral stripe is creamy white, and the lateral +stripe is dark brown (Pl. 14). The venter is white, and the iris is +dull bronze. In breeding males the vocal sac is yellow. + +_Remarks._--Although this species superficially resembles _Hyla +microcephala microcephala_, the latter is easily distinguished by the +narrow brown lateral stripe, as compared with the much wider stripe +in _H. robertmertensi_. No other hylids in northern Central America +and southern Mexico can be confused with this species. + +_Distribution._--_Hyla robertmertensi_ inhabits the Pacific slopes (to +elevations of 700 meters) and lowlands from eastern Oaxaca (east of +the Plains of Tehuantepec) southeastward to central El Salvador. The +species also occurs in the Cintalapa Valley (Atlantic drainage) in +southwestern Chiapas (Fig. 2.) The distribution seems to be limited on +the northwest and southeast by arid environments. The region in which +_Hyla robertmertensi_ lives is characterized by higher rainfall and +more luxuriant vegetation than occur on the Plains of Tehuantepec or +on the Pacific lowlands of eastern El Salvador and southern Honduras. +In addition to the localities listed below, Mertens (1952:30) +recorded the species from Hacienda Cuyan-Cuya, Depto. Sonsonate, El +Salvador. + + + [Illustration: Fig. 2. Map showing locality records for + _Hyla robertmertensi_.] + + +_Specimens examined._--490, as follows: +Mexico+: Chiapas: Acacoyagua, +USNM 114754-61; *2 km. W Acacoyagua, UMMZ 87843 (28), 87844 (50), +87845 (50), 87846 (45), 87847 (27), 87848 (3); 32 km. N Arriaga, KU +57619-24, 59917-8 (skeletons); Asuncion, FMNH 100413, 100501-4, UIMNH +26989-90, USNM 134267; *La Esperanza, USNM 114737-48, 114750-3, 17 km. +S Las Cruces, KU 57625-49, 59997 (eggs); 8.5 km. N Puerto Madero, UMMZ +119981 (2); *11.7 km. N Puerto Madero, UMMZ 119982; Tapachula, FMNH +100096, UIMNH 26987; *11 km. S Tapachula, KU 57605-18, 59916 (skeleton); +Tonola, FMNH 27073, 100505-10, UIMNH 26988. Oaxaca: Tapanatepec, +UMMZ 115245 (2), *1.6 km. E Tapanatepec, UMMZ 115244 (14); *4.3 km. +E Tapanatepec, UIMNH 38368-9; *7.5 km. W Tapanatepec, UMMZ 115246 +(39); 12.8 km. W Tapanatepec, KU 65007-14; 7.2 km. WNW Zanatepec, +UMMZ 115243 (77); *13.6 km. WNW Zanatepec, TNHC 25213-22; 22.7 km. +WNW Zanatepec, TNHC 25203-9. + ++Guatemala+: Jutiapa: Jutiapa, UMMZ 106848; La Trinidad, UMMZ +107733 (23). Retalhueleu: Casa Blanca, UMMZ 107732. + ++El Salvador+: La Libertad: 16 km. NW Santa Tecla, KU 44112. San +Salvador: 21.9 km. N San Salvador, UMMZ 119983 (6). + + + + ++Hyla phlebodes+ Stejneger + + + _Hyla phlebodes_ Stejneger, Proc. U. S. Natl. Mus., 30:817, June 4, + 1906 [Holotype.--USNM 2997 from "San Carlos," Costa Rica; + Burgdorf and Schild collectors]. Taylor, Proc. Biol. Soc. + Washington, 50:44, April 21, 1937; Univ. Kansas Sci. Bull., + 35:888, July 1, 1952; Univ. Kansas Sci. Bull., 39:25, November + 18, 1958. Fouquette, Evolution, 14:484, December 16, 1960. + Duellman and Trueb, Univ. Kansas Publ., Mus. Nat. Hist., 17:348, + July 14, 1966. + + _Hyla underwoodi_, Dunn, Occas. Papers Boston Soc. Nat. Hist., 5:413, + October 10, 1931; Occas. Papers Boston Soc. Nat. Hist. 8:72, + June 7, 1933; Amer. Mus. Novitiates, 747.2, September 17, 1934, + Gaige, Hartweg, and Stuart, Occas. Papers Mus. Zool., Univ. + Michigan, 357:5, October 26, 1937. Breder, 1946, Bull. Amer. Mus. + Nat. Hist., 86:416, August 22, 1946. + + +_Diagnosis._--Dark brown lateral stripe, if present, usually extending +only to insertion of forearm, never posteriorly to sacral region; +white line above brown stripe absent or faint; dorsal pattern weak, +usually consisting of irregular dashes or interconnected lines; +interorbital dark mark present; shanks having weakly defined +transverse bars. + +_Description and variation._--In the majority of specimens (70%) the +lateral dark stripe extends from the nostril to the eye and thence +above the tympanum to a point above the insertion of the arm; in 17 +per cent the stripe extends to the mid-flank, whereas in 13 per cent +the stripe is absent. A narrow and faint white line is present on the +canthus in some specimens, but no distinct white stripe is present +above the lateral dark line posterior to the eye. An interorbital bar +and transverse marks on the shanks are invariably present. The dorsal +markings are variable, but in most specimens (92%) consist of either +an X- or )(-shaped mark in the scapular region; in the other specimens +the markings are irregular short lines or absent. Approximately equal +numbers of specimens have a transverse bar, chevron, or broken lines +in the sacral region, whereas about eight per cent of the specimens +lack markings in the sacral region. + +When active at night, individuals are pale yellowish tan with faint +brown dorsal markings. By day they are tan with more distinct brown +markings (Pl. 14). The thighs are pale yellow; the belly is white. The +iris is pale creamy tan with brown flecks. In breeding males the vocal +sac is yellow. + +_Tadpoles._--Tadpoles of this species have been found in an extensive +grassy pond at Puerto Viejo, Costa Rica. The following description is +based on KU 104099, a specimen in development stage 36 (Gosner, 1960). + +Total length, 21.0 mm.; body length, 6.7 mm.; body slightly wider than +deep, snout pointed; nostrils large, directed anteriorly, situated +near end of snout; eyes small, situated dorsolaterally, directed +laterally; spiracle sinistral, located just posteroventral to eye; +anal tube dextral. Tail xiphicercal; caudal musculature moderately +deep, extending far beyond posterior edge of fins; fins deepest at +about midlength; dorsal fin extending onto body, slightly deeper than +caudal musculature; ventral fin slightly shallower than musculature. +Mouth small, terminal, lacking teeth and fringing papillae, but having +finely serrate beaks. In preservative top of head olive-tan with brown +flecks; dark stripe from snout through eye to posterior edge of body; +belly white, flecked with brown anteriorly; tail creamy tan with +grayish brown blotches. In life, dorsum of body reddish tan mottled +with darker brown; lateral stripe dark brown; belly white, mottled +with brown and black; caudal musculature heavily pigmented with +grayish tan; posterior tip of tail marked with dark gray; caudal fins +heavily blotched with grayish tan; iris orange-tan peripherally, red +centrally (Pl. 15). + +_Remarks._--This species has been confused with _Hyla microcephala +underwoodi_ by many workers. Dunn (1931, 1933, 1934) and Breder (1946) +referred Panamanian specimens of _H. phlebodes_ to _H. underwoodi_; +likewise, Gaige, Hartweg, and Stuart (1937) made the same error. Cole +and Barbour (1906) and Kellog (1932) used the name _H. phlebodes_ for +Mexican specimens of _H. microcephala underwoodi_. The similarity in +color pattern of _H. microcephala underwoodi_ and _H. phlebodes_ +easily accounts for the misapplication of names. Although both species +have nearly identical dorsal color patterns, that of _H. microcephala +underwoodi_ usually is bolder. Furthermore, in that species a narrow +white line usually is present above the well-defined lateral dark +stripe, whereas the lateral dark stripe is short and posterior to the +eye is not bordered above by a white line in _H. phlebodes_. + +The type locality "San Carlos, Costa Rica" given by Stejneger +(1906:817) apparently refers to a region, the Llanuras de San Carlos, +in the northern part of Alajuela Province, Costa Rica. + + + [Illustration: Fig. 3. Map showing locality records for + _Hyla phlebodes_.] + + +_Distribution._--_Hyla phlebodes_ inhabits humid tropical forests from +southeastern Nicaragua southeastward on the Caribbean slopes and +lowlands to the Canal Zone in Panama, thence eastward in the Chucunaque +Basin of eastern Panama and onto the Pacific lowlands of Colombia +(Fig. 3). The species also reaches the Pacific slopes in the Arenal +Depression in northwestern Costa Rica and in the Panamanian isthmus, +where it occurs in humid forests on the Pacific slope of El Valle and +Cerro La Campana. Mostly the species is found at low elevations, but +it occurs at 600 meters at Turrialba and at 700 meters at Finca San +Bosco in Costa Rica. + +_Specimens examined._--410, as follows: +Nicaragua+: Zelaya: Isla +Grande del Maiz, MCZ 14848; Rio Mico, El Recrero, UMMZ 79720 (6). + ++Costa Rica+: Alajuela: 12.4 km. N Florencia, MVZ 76108-10, USC 2628; +*Las Playuelas, 11 km. S Los Chiles, USC 7216; Los Chiles, USC 7217, +7219; 3 km. NE Muelle de Arenal, USC 2644 (2); *"San Carlos," USNM +29970. Cartago: Chitaria, KU 103690; *1.6 km. E Rio Reventazon Bridge, +east of Turrialba, UMMZ 119978 (2); *Tunnel Camp, near Peralta, +KU 32456, 32458-69, 41098 (skeleton); Turrialba, FMNH 101794, 103188-9, +KU 25725-9, 32439-48, 41095-7 (skeletons), 64797-827, 68300-2 +(skeletons), 68403 (eggs), 68404 (tadpoles), MCZ 29224-5, 29310-2, +UMMZ 119979 (6), USC 31, 256 (2), 458 (2), 580, 594, 599 (7), 7074 (2), +USNM 29933. Guanacaste: Arenal, USC 6254; *Finca San Bosco, USC 62724, +6276 (3), Guayabo de Bagaces, USC 7022 (3), 7023; *Laguna Arenal, +USC 6262 (4); 3 km. NE Tilaran, USC 524; *5 km. NE Tilaran, USC 6269; +*6 km. NE Tilaran, UMMZ 122653 (6), S-2680 (skeleton), USC 523 (8). +Heredia: Puerto Viejo, KU 64828-63, 68303-7 (skeletons), 68405-6 +(tadpoles), 104099-100 (tadpoles); *1.5 km. N Puerto Viejo, KU 64871; +*1 km. S Puerto Viejo, KU 86432-40; *4.2 km. W Puerto Viejo, KU +64864-5; *5.9 km. W Puerto Viejo, KU 64866-70; *7.5 km. W Puerto Viejo, +KU 86431. Limon: Batan, UMMZ 119980 (2); La Castilla, ANSP 23707; +Puerto +Limon+, KU 32449-55. + ++Panama+: Bocas del Toro: 3.2 km. NW Almirante, KU 96026; Cayo de +Agua, KU 96027-31; Fish Creek, KU 96032-4. Canal Zone: Barro Colorado +Island, AMNH 69790, ANSP 23244-50; FMNH 13380, 22972-4; Juan Mina, +AMNH 55429, UU 3899; *8.6-13.8 km. N Miraflores Locks, TNHC 23439, +23477, 23484-8, 23491, 23494-9, 23501-2, 23504-8, 23510-17, 23519-30, +23532-8, 23541-54, 23561. *Rio Chagres, AMNH 55431-4; Rio Cocoli, +3.5 km. N Miraflores Locks, TNHC 23461, 23489-90, 23493, 23500, 23503, +23509, 23518, 23531, 23539-40; *Summit, ANSP 23361, KU 97788; *Three +Rivers Plantation, SU 2130. Cocle: El Valle de Anton, AMNH 55435, +69786-9, ANSP 23506-9. Colon: Achiote, KU 77215-78; Ciricito, CAS +71499-500, 71505-6. Darien: Rio Canclon at Rio Chucunaque, UMMZ 126733; +Rio Chucunaque, near Yavisa, AMNH 51783. Panama: Cero La Campana, FMNH +67847-50. + ++Colombia+: Choco: Andagoya, FMNH 81856; Boca de Raspadura, AMNH +13570-8. + + + + ++Hyla sartori+ Smith + + + _Hyla underwoodi_ (in part), Smith and Taylor, Bull. U. S. Natl. + Mus., 194:85, June 17, 1948. + + _Hyla microcephala sartori_ Smith, Herpetologica, 7:186, + December 31, 1951 [Holotype.--UIMNH 20934 from 1 mile north of + Organos, south of El Treinte, Guerrero, Mexico; H. M. Smith and + E. H. Taylor collectors]. Duellman, Univ. Kansas Publ., Mus. Nat. + Hist., 15:124, December 20, 1961. Porter, Herpetologica, 18:168, + October 17, 1962. Davis and Dixon, Herpetologica, 20:230, + January 25, 1965. Duellman, Univ. Kansas Publ. Mus. Nat. Hist., + 15:652, December 30, 1965. + +_Diagnosis._--Dorsum tan with broad dark brown chevrons or transverse +bars; shanks marked with two or three broad transverse bars; +dorsolateral stripes absent. + +_Description and variation._--No noticeable geographic variation is +apparent in either structural features or coloration in this species. +All specimens lack a dorsolateral dark stripe and white line, although +a dark line is present on the canthus and dissipates in the loreal +region. A broad interorbital brown bar is present in all specimens. +The color pattern on the dorsum invariably consists of a broad, dark, +chevron-shaped mark in the scapular region and a broad dark chevron or +transverse bar in the sacral region. The shanks invariably have two or +three dark brown transverse bars. + +When active at night individuals are yellowish tan above with chocolate +brown markings (Pl. 14). The belly is white, and the thighs are pale +yellowish tan. The iris is dark bronze-color. In breeding males the +vocal sac is yellow. By day some individuals were observed to change +to creamy gray with distinct darker markings. + +_Remarks._--Although tadpoles of this species have not been found, +observations on the breeding sites indicate that the tadpoles probably +develop in ponds. Except for calling males observed around a pool in a +stream-bed 11.8 kilometers west-northwest of Tierra Colorada, Guerrero, +all breeding congregations have been found at temporary ponds. + +Smith (1951:186) named _Hyla sartori_ as a subspecies of _Hyla +microcephala_. This subspecific relationship seemed reasonable until +analysis of the mating calls showed that the call of _H. sartori_ is +more nearly like that of _H. phlebodes_ than that of _H. microcephala_. +The broad hiatus separating the ranges of _H. microcephala_ and _H. +sartori_ is additional evidence for considering _H. sartori_ as a +distinct species. + + + [Illustration: Fig. 4. Map showing locality records for + _Hyla sartori_.] + + +_Distribution._--_Hyla sartori_ occurs in mesophytic forests to +elevations of about 300 meters on the Pacific slopes of southern Mexico +from southwestern Jalisco to south-central Oaxaca (Fig. 4). The lack of +specimens from Colima and Michoacan probably reflects inadequate +collecting instead of the absence of the species there. On the basis of +available habitat the species would be expected to occur in Nayarit, +but extensive collecting there has failed to reveal its presence. The +semi-arid Plains of Tehuantepec apparently limit the distribution to +the east. + +_Specimens examined._--190, as follows: +Mexico+: Guerrero: 5 km. E +Acapulco, AMNH 54611-2; 23.2 km. N Acapulco, UIMNH 26404-7; Colonia +Buenas Aires, 23 km. E Tecpan de Galeana, UMMZ 119223 (7); *El +Limoncito, FMNH 75785, 100390-402, 104631, 104633, UMMZ 117250, USNM +134266; El Treinte, FMNH 100403, UIMNH 20935-7; Laguna Coyuca, AMNH +59686; La Venta, MCZ 29635; *Morjonares, UIMNH 26392-402; 1.6 km. +N Organos, FMNH 100404-5, UIMNH 20933-4; 19.2 km. S Petaquillas, +UIMNH 26408; 6.1 km. E. Tecpan de Galeana, TNHC 23396-408; *11.2 km. +N Tierra Colorada, UIMNH 26403; 11.8 km. WNW Tierra Colorada, UMMZ +119225 (51), S-2677-9 (skeletons); Zacualpan, UMMZ 119224 (6). Jalisco: +6.4 km. NE La Resolana, KU 67853-69; 24 km NE La Resolana, KU 67870-3. +Oaxaca: 3 km. N Pochutla, KU 57539; 13.4 km. N Pochutla, UMMZ +123495 (40). + + + + +CRANIAL OSTEOLOGY + + +The frogs of the _Hyla microcephala_ group have a minimal amount +of cranial ossification as compared to more generalized hylid skulls, +such as _Smilisca_ (Duellman and Trueb, 1966). In the _Hyla +microcephala_ group the sphenethmoid is small and short, and a large +frontoparietal fontanelle is present. The quadratojugal exists only +as a small spur and is not in contact with the maxillary. The prootics +are poorly developed. The anterior and posterior arms of the squamosal +are short; the anterior arm extends no more than one-fourth of the +distance to the maxillary, and the posterior arm does not have a bony +connection with the prootic. The nasal lacks a maxillary process, and +the medial ramus of the pterygoid lacks a bony connection to the +prootic. + +Teeth are absent on the parasphenoid and palatines, but present on the +maxillaries, premaxillaries, and prevomers. The teeth are simple, +pointed, and slightly curved. Although the number of teeth varies +(Table 3), no consistent differences between the species are apparent. + + +Table 3.--Variation in the Number of Teeth in the Species of the Hyla + Microcephala Group. (N=Number of Jaws, or Twice the Number of + Individuals; Means are Given in Parentheses After the Observed + Ranges). + +========================+====+=============+==============+========== + Species | N | Maxillary | Premaxillary | Prevomer +------------------------+----+-------------+--------------+---------- +_H. microcephala_ | 32 | 31-47(37.8) | 4-13(8.9) | 2-4(3.2) + | | | | +_H. phlebodes_ | 10 | 38-45(40.1) | 8-13(10.3) | 2-5(3.9) + | | | | +_H. robertmertensi_ | 6 | 23-43(32.8) | 7-12(10.5) | 2-3(2.7) + | | | | +_H. sartori_ | 6 | 27-43(38.2) | 9-10(9.3) | 3-4(3.7) +------------------------+----+-------------+--------------+---------- + + + [Illustration: PLATE 13 + Upper figure, _Hyla microcephala microcephala_ (KU 64593); + middle figure, _H. microcephala underwoodi_ (KU 64565); + lower figure, _H. microcephala underwoodi_ (UMMZ 115247). + All approximately x3.] + + + [Illustration: PLATE 14 + Upper figure, _Hyla robertmertensi_ (UMMZ 115243); + middle figure, _H. phlebodes_ (KU 64798); lower figure, + _H. sartori_ (UMMZ 119225). All approximately x3.] + + + [Illustration: PLATE 15 + Tadpoles of _Hyla microcephala_ group: upper figure, _H. m. + microcephala_ (KU 104097); lower figure, _H. phlebodes_ + (KU 104099). Both x4.] + + + [Illustration: PLATE 16 + Audiospectrograms and sections of mating calls of _Hyla + microcephala_ group: + (a) _H. m. microcephala_ (KU Tape No. 19); + (b) _H. robertmertensi_ (KU Tape No. 41); + (c) _H. phlebodes_ (KU Tape No. 6); + (d) _H. sartori_ (KU Tape No. 190).] + + +Table 4.--Comparative Cranial Osteology of Hyla microcephala Group + +===============+=======================+========================+ + Character | _H. microcephala_ | _H. robertmertensi_ | +---------------+-----------------------+------------------------+ +Frontoparietal | Minimally ossified | Ossification extensive | + | with large fontanelle | anteriorly with narrow | + | extending from | medial separation; | + | sphenethmoid to | fontanelle largest in | + | occipital ridge. | parietal region. | + | | | + | | | +Nasals | Moderately long and | Moderate in size; | + | slender; arcuate in | slightly wider | + | dorsal view. | anteriorly than | + | | posteriorly in dorsal | + | | view. | + | | | +Sphenethmoid | Extremely short in | Moderately short in | + | dorsal view. | dorsal view. | + | | | + | | | + | | | +Columella | Distal and greatly | Distal and slightly | + | expanded. | expanded or not. | +---------------+-----------------------+------------------------+ +Table 4. (Continued) +===============+========================+======================== + Character | _H. phlebodes_ | _H. sartori_ +---------------+------------------------+------------------------ +Frontoparietal | Ossification extensive | Ossification moderately + | anteriorly with narrow | extensive anteriorly; + | medial separation; | medial separation of + | fontanelle largest in | about uniform width + | parietal region. | throughout length of + | | fontanelle. + | | +Nasals | Moderate in size; | Long and broad; + | slightly wider | arcuate in dorsal + | anteriorly than | view. + | posteriorly in dorsal | + | view. | + | | +Sphenethmoid | Moderately short in | Moderately short in + | dorsal view. | dorsal view; ossified + | | anteriorly between + | | nasals. + | | +Columella | Distal and not | Distal and not + | expanded. | expanded. +---------------+------------------------+------------------------ + + + [Illustration: Fig. 5. Dorsal views of the skulls of (a) _Hyla m. + microcephala_ (KU 68293) and (b) _H. sartori_ (UMMZ S-2677). + Both x 12.] + + + [Illustration: Fig. 6. Dorsal views of skulls of (a) _Hyla phlebodes_ + (KU 68303) and (b) _H. robertmertensi_ (KU 59917). Both x 12.] + + +Despite the great reduction in the ossification of the cranial +elements, certain apparently consistent differences exist between the +species seem to be consistent. The most notable differences are: +1) amount of ossification of the frontoparietals and consequent shape +and size of the frontoparietal fontanelle, 2) shape of the nasals, +3) shape and extent of the sphenethmoid, and 4) shape of the columella +(Table 4, Figs. 5-6). On the basis of these characters, _Hyla +microcephala_ can be set apart from the other species and characterized +as having a poorly ossified frontoparietal and correspondingly large +frontoparietal fontanelle; long, slender, arcuate nasals; extremely +short sphenethmoid; and expanded distal end of the columella. The other +species in the group (_phlebodes_, _robertmertensi_, and _sartori_) +have more ossification of the frontoparietals, broader nasals, only a +moderately short sphenethmoid, and an unexpanded distal end of the +columella. Among these three species, the skulls of _phlebodes_ and +_robertmertensi_ are most nearly alike, whereas the skull of _sartori_ +differs by having a differently shaped frontoparietal fontanelle, +broader nasals, and an ossified anterior extension of the sphenethmoid +between the nasals (compare Fig. 5b with Fig. 6 a-b). + +Although all skulls examined belong to breeding adults, the extent of +the ossification of the frontoparietals and the resulting shape of the +frontoparietal fontanelle might be correlated with the age of the frog. +Nevertheless, in the 24 skulls of _Hyla microcephala_ examined, the +frontoparietals are less extensively ossified than in the skulls of the +other species. The trivial differences among the other three species +certainly are suggestive of close relationship, but on the basis of +present knowledge of the evolutionary trends in hylid cranial +osteology, the differences offer little evidence for determining +phylogenetic lineage. + + + + +ANALYSIS OF MATING CALLS + + +Calls of all five taxa were compared in several characteristics, of +which three are deemed most significant systematically. These are +1) the pattern and duration of the notes of a call-group, 2) the +fundamental frequency, and 3) the dominant frequency. Air temperatures +were noted at the time the calls were recorded, but no valid +correlation could be determined between this factor and any of the +parameters of the calls; consequently recordings made at all +temperatures (21-29 deg. C.) were grouped together. + +_Pattern and duration of notes._--In all five taxa the basic pattern +consists of a call-group made up of one primary note followed by a +series of shorter secondary notes. In some species the secondary notes +differ from the primary in other characteristics. Both subspecies of +_Hyla microcephala_ have a long, unpaired primary note followed by 0 +to 18 (usually about 4) somewhat shorter paired secondary notes. In +calls of _Hyla m. microcephala_ the mean duration of the primary is +0.131 (0.10-0.16) second and that of the secondaries is 0.101 +(0.05-0.14) second, whereas in _H. m. underwoodi_ the mean duration of +the primary is 0.018 (0.05-0.15) second and that of the secondaries is +0.086 (0.06-0.11) second. + +_Hyla robertmertensi_ has a reverse of this pattern in that the primary +note is paired and the secondaries are unpaired. In the sample studied +a call-group contains 0-28 secondary notes (generally about 3). The +mean duration of the primary is 0.091 (0.07-0.11) second and that of +the secondaries is 0.040 (0.025-0.06) second. + +_Hyla phlebodes_ and _sartori_ have call-groups composed of a rather +short, unpaired primary and several short, unpaired secondaries +(0-28 in _phlebodes_, 0-23 in _sartori_). The mean duration of the +primary of _phlebodes_ is 0.105 (0.07-0.16) second and that of the +secondaries is 0.067 (0.035-0.12) second. The mean duration of the +primary of _sartori_ is 0.080 (0.07-0.09) second and that of the +secondaries is 0.053 (0.035-0.07) second. + +The two subspecies of _H. microcephala_ are identical in call pattern +and agree closely in duration of notes, although those of the nominate +subspecies tend to be slightly longer. _Hyla robertmertensi_ is +distinctive in call pattern in that it is the only species having a +paired primary; the duration of the primary is completely overlapped by +that in the other species, but the secondaries tend to be the shortest +in the group. The call patterns of _H. phlebodes_ and _H. sartori_ are +identical and the range of duration of notes of _phlebodes_ completely +overlaps that of _sartori_, although both the primary and secondary +notes of the latter tend to be somewhat shorter (Table 5, Pl. 16). + +_Fundamental frequency._--This parameter was analyzed for the primary +notes. It was measured for the secondaries as well and was found to +differ in magnitude in the same way as the primary note. In a few +examples of both subspecies of _H. microcephala_ a high primary note, +in which the fundamental frequency is exceptionally high, is sometimes +emitted (Fouquette, 1960b). None of these notes was used in this +analysis; only the fundamental frequencies of normal primary notes are +compared (Table 5, Fig. 7). + + +Table 5.--Comparison of Normal Mating Calls in the Hyla microcephala + Group. (Observed Range Given in Parentheses Below Mean; + Unless Otherwise Noted Data Are for Primary Notes.). + +----------------+--+---------+---------+-------------------+-------------- + | |Dominant | Funda- |Duration of notes | Repetition + | | | mental| (seconds) | rate of + Species |N |frequency|frequency+---------+---------+ secondaries + | | (cps) | (cps) | Primary |Secondary|(notes/minute) +----------------+--+---------+---------+---------+---------+-------------- +_H. m. |44| 5637 | 205 | 0.13 | 0.10 | 268 + microcephala_ | |(5150 |(184-244)|(0.11 |(0.05 | (192-353) + | | -5962)| | -0.16)| -0.14)| + | | | | | | +_H. m. |47| 5772 | 220 | 0.11 | 0.09 | 283 + underwoodi_ | |(5177 |(192-275)|(0.05 |(0.06 | (197-384) + | | -6200)| | -0.15)| -0.11)| + | | | | | | +_H. |25| 5388 | 162 | 0.09 | 0.04 | 418 + robertmertensi_| |(5150 |(140-178)|(0.07 |(0.03 | (368-570) + | | -5785)| | -0.11)| -0.06)| + | | | | | | +_H. phlebodes_ |34| 3578 | 148 | 0.11 | 0.07 | 284 + | |(3220 |(125-158)|(0.07 |(0.04 | (210-350) + | | -4067)| | -0.16)| -0.12)| + | | | | | | +_H. sartori_ |10| 3217 | 126 | 0.08 | 0.05 | 434 + | |(2950 |(116-135)|(0.07 |(0.04 | (396-477) + | | -3600)| | -0.09)| -0.07)| +----------------+--+---------+---------+---------+---------+-------------- + + +The two subspecies of _H. microcephala_ agree closely in fundamental +frequency. There is considerable overlap, but the difference +between the means is significant at the 0.001 level of probability +(t = 4.2406). The call of _H. robertmertensi_ does not overlap that +of _H. sartori_ or either subspecies of _H. microcephala_ in this +parameter; but it does overlap that of _H. phlebodes_, although again +the difference between the means is significant at the 0.001 level +(t = 9.360). _Hyla phlebodes_ and _sartori_ have the lowest fundamental +frequencies, and there is some overlap, but here too the difference +between the means is significant at the 0.001 level (t = 4.923). + +_Dominant frequency._--A dominant band of frequencies cuts across +the harmonics of the fundamental, obscuring the harmonic pattern and +generally shifting upward in frequency. The midpoint of this band is +measured at the terminal border as the dominant frequency. As with the +fundamental frequency, only the normal primary notes were utilized in +the comparisons (Table 5, Fig 8). + + + [Illustration: Fig. 7. Variation in the fundamental frequency of the + normal primary notes in the _Hyla microcephala_ group. The + horizontal lines = range of variation, vertical lines = mean, + solid bars = twice the standard error of the mean, and open + bars = one standard deviation. The number of specimens in each + sample is indicated in parentheses after the name of the taxon.] + + +The two subspecies of _H. microcephala_ agree more closely in this +parameter than in fundamental frequency. The overlap is great, but the +difference between the means is significant at the 0.001 level +(t = 3.658). The calls of both subspecies completely overlap that of +_robertmertensi_ in this parameter, but the difference between the +means is significant at the 0.001 level. The calls of _H. phlebodes_ +and _H. sartori_ overlap considerably in this characteristic, although +the difference between the means is significant at the 0.001 level +(t = 7.504) (Fig. 9). The call of neither species overlaps those of +_H. microcephala_ and _robertmertensi_. + + + [Illustration: Fig. 8. Variation in the mid-point of the dominant + frequency band of the normal primary notes in the _Hyla + microcephala_ group. The horizontal lines = range of variation, + vertical lines = mean, solid bars = twice the standard error of + the mean, and open bars = one standard deviation. The number of + specimens in each sample is indicated in parentheses after the + name of the taxon.] + + + [Illustration: Fig. 9. Scatter diagram relating the dominant and + fundamental frequencies of the normal primary notes in the + _Hyla microcephala_ group. Each symbol represents a different + individual.] + + +_Repetition rate._--The repetition rate of the secondary notes, in +calls consisting of more than one secondary, was measured for each +form. A considerable amount of variation in this parameter was found +in all of the taxa (Table 5). This variation probably is due in part +to the effect of temperature differences. Repetition rate is the only +parameter analyzed for which there is a correlation with the +air-temperature, but even here the correlation is weak, probably due +to the microenvironmental effects of humidity, air-movement, and other +factors in addition to the ambient air temperature that influences the +body temperature of the frogs. These rates are nearly alike in both +subspecies of _H. microcephala_ and in _phlebodes_. The repetition +rates in _H. robertmertensi_ and _H. sartori_ are considerably faster +than in the other three taxa. _Hyla sartori_ has the fastest +repetition rate of the group. + +In all characteristics of the mating calls the two subspecies of +_H. microcephala_ agree closely, as might be expected, although the +differences are statistically significant. _Hyla robertmertensi_ is +distinctive in call pattern and seems to be closer to _microcephala_ +in dominant frequency but closer to _H. phlebodes_ in fundamental +frequency. Thus, it is somewhat intermediate between _microcephala_ +and _phlebodes_. The identical pattern and similarity in fundamental +and dominant frequencies of the calls of _H. phlebodes_ and _H. sartori_ +possibly indicate close relationship. + +_Geographic variation in call._--_Hyla m. microcephala_ has higher +fundamental and dominant frequencies in Costa Rica than in Panama. In +Costa Rican _H. m. underwoodi_ the fundamental and dominant frequencies +are lower than in other parts of the range. Frogs of this subspecies +recorded in Nicaragua and Honduras have slightly lower dominant +frequencies and higher fundamental frequencies than those recorded in +Guatemala or Oaxaca. The duration of both primary and secondary notes +decreases to the south; samples from Nicaragua and Costa Rica have the +shortest notes. Comparison of duration of notes in the two subspecies +shows that the Panamanian _H. m. microcephala_ have slightly longer +notes than do any _H. m. underwoodi_; the more northern populations of +_H. m. underwoodi_ from Mexico most closely approach _H. m. +microcephala_ in this characteristic. + +The calls of _H. robertmertensi_ in Oaxaca have higher dominant and +fundamental frequencies and longer secondary notes than do those in +Chiapas. + +The calls of _H. phlebodes_ recorded at Puerto Viejo, Costa Rica, +have slightly lower dominant frequencies than do those recorded at +Turrialba, Costa Rica, and in Panama, whereas those recorded at +Turrialba have lower fundamental frequencies than in other samples. +The duration of notes is slightly shorter in both Costa Rican samples +than in those recorded in Panama. + + + + +LIFE HISTORY + + +The frogs of the _Hyla microcephala_ group breed in shallow grassy +ponds. In some places they breed in permanent ponds, but usually +congregate around temporary pools, such as depressions in forests, +flooded fields, and roadside ditches. At the height of their breeding +season, usually in the early part of the rainy season, the +congregations are made up of large numbers of individuals. In April, +1961, and in June, 1966, the senior author noted nearly continuous +choruses of _H. m. microcephala_ in roadside ditches along the 75 +kilometers of road between Villa Neily and Palmar Sur, Puntarenas +Province, Cost Rica; on June 20, 1966, at Puerto Viejo, Heredia +Province, Costa Rica, he estimated approximately 900 _Hyla phlebodes_ +in one pond, and two nights later noticed that the number of +individuals had increased substantially. Other observations by the +first author on size of breeding congregations include nearly +continuous choruses of _H. m. underwoodi_ between Villahermosa and +Teapa, Tabasco, in July of 1958, an estimated 400 _Hyla robertmertensi_ +in a road side ditch 7.2 kilometers west-northwest of Zanatepec, +Oaxaca, on July 13, 1956, and approximately 150 _Hyla sartori_ around +a rocky pool in a riverbed, 11.8 kilometers west-northwest of Tierra +Colorada, Guerrero, on June 28, 1958. + +The length of the breeding season seemingly is more dependent on +climatic conditions in various parts of Middle America than on +behavioral differences in the various species. Thus, Fouquette (1960b) +found in the Canal Zone that _H. m. microcephala_ formed breeding +choruses from May through January, the entire rainy season in that +area. In the wetter coastal region of Puntarenas Province, Costa Rica, +the species breeds as early as mid-March, whereas in the drier region +encompassing Guanacaste Province, Costa Rica, and southwestern +Nicaragua breeding activity is initiated by the first heavy rains of +the season, usually in June. + +_Hyla phlebodes_ inhabits regions having rainfall throughout the year. +Although large breeding congregations are most common in the early +parts of the rainy season, males probably call throughout the year. At +Puerto Viejo in Costa Rica the senior author has heard _Hyla phlebodes_ +in February, April, June, July, and August. Charles W. Myers noted +calling males of this species in the area around Almirante, Bocas del +Toro Province, Panama, in September, October, and February. An +exception to the correlation between rainfall and breeding activity +was noted by the junior author in _Hyla phlebodes_ in the Canal Zone, +where he noticed a decrease in activity of that species in October and +November, when the rains are heaviest and most frequent. Furthermore, +independent observations made by both of us indicate that _H. +phlebodes_ does not reach peaks of activity during or immediately +after heavy rains, but instead builds up to peaks of activity two or +three days after a heavy rain. This is in contrast to the other +species, all of which characteristically inhabit drier environments +than does _H. phlebodes_. Peaks of breeding activity in the other +species occur immediately after, or even during, heavy rains. + +The calling location of the males generally is on vegetation above, +or at the edge of, the water. _Hyla microcephala_ and _H. phlebodes_ +call almost exclusively from grasses and sedges; _phlebodes_ usually +calls from taller and more dense grasses than does _microcephala_. +Except for some minor differences in calling location observed by +the junior author (Fouquette, 1960b) in the Canal Zone, the differences +in density and height of grasses utilized for calling-locations +probably is dependent primarily on the nature of the available +vegetation. Although bushes and broad-leafed herbs are usually present +at the breeding sites, males of these species seldom utilize them for +calling locations. Both _H. robertmertensi_ and _H. sartori_ have been +observed calling from grasses, herbs, bushes, and low trees. Calling +males of _robertmertensi_ have been found two meters above the ground +in small trees. + +Daytime retreats in the breeding season sometimes are no more than +shaded clumps of vegetation adjacent to a pond or in clumps of grass +in a pond. Individuals of _H. m. underwoodi_ were found by day under +the outer sheaths of banana plants next to a water-filled ditch. Dry +season refuges are unknown. + +Amplexus is axillary in all four species. Egg deposition has been +observed in _H. m. microcephala_, _m. underwoodi_, and _phlebodes_. +In all three the eggs are deposited in small masses that float near +the surface of the water and usually are at least partly attached to +emergent vegetation. Each clutch does not represent the entire egg +complement of the female. + +Tadpoles are definitely known of only _H. m. microcephala_ and +_phlebodes_; these have been described in the preceding accounts of +the species. The tadpoles of these two species can be distinguished +readily (Pl. 15). The tadpole of _H. microcephala_ has a uniformly +white venter and nearly transparent tail, whereas in _H. phlebodes_ +the venter is flecked anteriorly and the tail is mottled. In life, _H. +microcephala_ is easily recognized by the orange posterior half of +the tail, whereas the tail in _H. phlebodes_ is mottled tan and grayish +brown. + + + + +PHYLOGENETIC RELATIONSHIPS + + +The evidence already presented on osteology, external structure, +coloration, mating call, and life history emphatically show that the +four species under consideration are a closely related assemblage. +Now the question arises: To what other groups in the genus is the +_Hyla microcephala_ group related? Furthermore, it is pertinent to +this discussion to attempt a reconstruction of the phylogeny of the +group as a whole and of the individual species in the _Hyla +microcephala_ group. With regard to the relationships of the group we +must take into account certain species in South America. Our endeavors +there are hampered by the absence of data on the mating calls and life +histories of most of the relevant species. + +As mentioned in the account of _Hyla m. microcephala_, the species +_microcephala_ possibly is subspecifically related to _Hyla misera_, a +frog widespread in the Amazon Basin. _Hyla misera_ resembles +_microcephala_ in coloration, external structure, and cranial +characters. The frontoparietals are equally poorly ossified, and the +frontoparietal fontanelle is extensive. Our principal reason for not +considering the two taxa conspecific at this time is our lack of +knowledge concerning the color of living _H. misera_, the structure of +the tadpoles, and the characteristics of the mating call. Even with the +absence of such data that we think essential to establish the +nomenclature status of the taxa, we are confident that the two are +sufficiently closely related that any discussion of the phylogenetic +relationships of one species certainly must involve consideration of +the other. + +_Hyla misera_ possibly is allied to other small yellowish tan South +American _Hyla_ that lack dark pigmentation on the thighs. Probable +relatives are _Hyla elongata_, _minuta_ (with _goughi_, _pallens_, +_suturata_, _velata_, and possibly others as synonyms), _nana_, and +_werneri_. The consideration of the interspecific relationships of +these taxa is beyond the scope of this paper, but we can say that each +of these species has a pale yellowish tan dorsum, relatively broad +dorsolateral brown stripe, and narrow longitudinal brown lines or +irregular marks on the dorsum. Furthermore, examination of the skulls +of _elongata_, _nana_, and _werneri_ reveals that they are like +_misera_ and _microcephala_ in the nature of the frontoparietal +fontanelle and in having a greatly reduced quadratojugal. Thus, on the +basis of cranial and external characters the _Hyla microcephala_ group +can be associated with _Hyla misera_ and its apparent allies in South +America. This association can be only tentative until the mating calls, +tadpoles, and chromosome numbers of the South American species are +known. + +Among the Middle American hylids, only the _Hyla microcephala_ group +and _H. ebraccata_ have a haploid number of 15 chromosomes (Duellman +and Cole, 1965). All other New World _Hyla_, for which the number is +known, have a haploid number of 12; the only other _Hyla_ having 15 +is a Papuan _Hyla angiana_ (Duellman, 1967). + +_Hyla ebraccata_ occurs in the humid tropical lowlands of Middle +America and the Pacific lowlands of northwestern South America. It is +the northernmost, and only Central American, representative of the +_Hyla leucophyllata_ group, which is diverse (about 10 species +currently recognized) and widespread in tropical South America east of +the Andes. This group is characterized by having broad, flat skulls +with larger nasals and more ossification of the frontoparietals than +in the _Hyla microcephala_ group. The quadratojugal is present as a +small anteriorly projecting spur that does not connect with the +maxillary. Externally, the _Hyla leucophyllata_ group is characterized +by having a well-developed axillary membrane, uniformly yellow thighs, +and a dorsal color pattern in many species consisting of a dark lateral +band, a pale dorsolateral band or dorsal ground color, and a large +middorsal dark mark. In some species, the dorsal pattern consists of +small dark markings or is nearly uniformly pale. At least in the +Central American _Hyla ebraccata_, the mating call consists of a +single primary note followed by a series of shorter secondary notes, +the tadpoles have xiphicercal tails and lack teeth, and the haploid +number of chromosomes is 15. On the strength of these observations it +seems imperative to consider the _Hyla leucophyllata_ group as a close +ally to the _Hyla microcephala_ group. Successful artificial +hybridization supports the close relationship of _H. m. microcephala_ +and _phlebodes_; partial success of artificial hybridization of these +two with _ebraccata_ (Fouquette, 1960b) provides further evidence for +close relationship between the _Hyla leucophyllata_ and _Hyla +microcephala_ groups. + +In Mexico and northern Central America two small species, _Hyla picta_ +and _Hyla smithi_, comprise the _Hyla picta_ group. These frogs +resemble members of the _Hyla microcephala_ group by having a +yellowish tan dorsum with a dorsolateral white stripe and uniformly +yellow thighs. Furthermore the mating call is not unlike those of +the species in the _Hyla microcephala_ group. Despite these +similarities, the _Hyla picta_ group differs from the _Hyla +microcephala_ group by having a well-developed quadratojugal that +connects to the maxillary, tadpoles with teeth present and caudal fins +completely enclosing the caudal musculature, and a haploid number of +12 chromosomes. In all of these characteristics the frogs of the + +_Hyla picta_ group more closely resemble other Middle American _Hyla_ +than they do the _Hyla microcephala_ group. Therefore, it can best be +presumed that the superficial resemblances of coloration and the +mating call are the result of convergence. + +Since the _Hyla microcephala_ and _leucophyllata_ groups apparently +are related and since the greatest diversity of these frogs is in +South America (if _Hyla misera_ and its relatives are placed with the +_Hyla microcephala_ group), it seems appropriate to place the centers +of origins of these groups in South America. Therefore, the _Hyla +microcephala_ group and _Hyla ebraccata_ of the _Hyla leucophyllata_ +group either have immigrated into Central America, or they are +representatives of those groups that were isolated in Central America +during most of the Cenozoic when South America was separated from +Central America. + +The interspecific relationships of the species in the _Hyla +microcephala_ group are not clear. On the basis of coloration, _H. m. +microcephala_ and _H. robertmertensi_ are close, and _H. m. underwoodi_ +and _H. phlebodes_ are nearly identical. The mating calls of _H. +phlebodes_ and _sartori_ closely resemble one another, whereas the call +of _robertmertensi_ is intermediate between these and _microcephala_. + +In most respects _Hyla microcephala_ is distinct from the other +species, and with the exception of the amount of ossification of the +frontoparietals, the other species can be easily derived from a +_microcephala_-like ancestor. Possibly the slightly increased +ossification of the frontoparietals in _robertmertensi_, _phlebodes_, +and _sartori_ is secondary, or possibly after differentiation of the +species the amount of ossification was further reduced in +_microcephala_. If so, the species fall into a reasonable phylogenetic +scheme that has _microcephala_ as the extant species most like the +ancestral stock. + +We visualize the evolutionary history of the group to have followed +a course that began with the invasion of Central America by a +_microcephala_ ancestral stock that differentiated into two populations +in lower Central America--a _microcephala_-like frog on the Pacific +lowlands and a _phlebodes_-like frog on the Caribbean lowlands. +Differentiation could have been brought about by isolation by montaine +or marine barriers. The population on the Pacific lowlands either was +preadapted for subhumid conditions or became so adapted and dispersed +northward onto the Pacific lowlands of northern Central America. +Simultaneously the frogs on the Caribbean lowlands, which were adapted +to humid environments, dispersed northward in the humid forested +regions to southern Mexico and crossed the Isthmus of Tehuantepec onto +the Pacific slopes of Oaxaca and Guerrero northward to Jalisco. +Subsequent development of arid conditions, possibly in the Pliocene, +Pleistocene, or even as late as the Thermal Maximum in post-Wisconsin +time, resulted in a restriction of the ranges in northern Central +America, thereby isolating part of the _phlebodes_-stock on the +Pacific slopes of Mexico, where it adapted to drier conditions and +evolved into _sartori_. The rest of the _phlebodes_-stock was +restricted to the humid forests on the Caribbean lowlands of lower +Central America. The increased aridity on the Pacific lowlands +eliminated the _microcephala_-stock from southern Honduras and +northwestern Nicaragua and in so doing left an isolated population on +the lowlands of Chiapasand Guatemala, which differentiated into +_robertmertensi_. The original stock on the Pacific lowlands of Panama +and southeastern Costa Rica became _microcephala_. + +If the _microcephala_-stock was, as we believe, better adapted for +existence under subhumid conditions than was the _phlebodes_-stock, +the development of subhumid conditions in much of the lowland region +of northern Central America and southern Mexico would have permitted +the expansion of the range of _microcephala_ into the area now +inhabited by _H. m. underwoodi_, while _phlebodes_ was being eliminated +from this area by climatic conditions that were unsuited to its +survival there. Perhaps the similarity in coloration of _H. m. +underwoodi_ and _phlebodes_ is the result of convergence or possibly +hybridization occurred at the time the former was expanding its range +and the latter's range was being restricted. If hybridization did +occur, the differences in mating call subsequently were enhanced, +thereby providing a valid isolating mechanism in sympatric populations. + +_Hyla microcephala_ and _phlebodes_ range into northern South America. +Probably both species entered South America in relatively recent times +after they had differentiated from one another in Central America. + + + + +LITERATURE CITED + + +Boulenger, G. A. + 1898. Fourth report on additions to the batrachian collection in the + Natural-History Museum. Proc. Zool. Soc. London, 1898, + pp. 373-482, pls. 38-39. October 1. + 1899. Descriptions of new batrachians in the collection of the + British Museum (Natural History). Ann. Mag. Nat. Hist, ser. + 7, 3:273-277, pls. 11-12. + +Breder, C. M. Jr. + 1946. Amphibians and reptiles of the Rio Chucunaque Drainage, Darien, + Panama, with notes on their life histories and habits. Bull. + Amer. Mus. Nat. Hist, 86:375-436, pls. 42-60, August 26. + +Cole, L. J. and Barbour, T. + 1906. Vertebrata from Yucatan: Reptilia; Amphibia; Pisces. Bull. Mus. + Comp. Zool., 50:146-159. November. + +Cope, E. D. + 1886. Thirteenth contribution to the herpetology of tropical America. + Proc. Amer. Philos. Soc, 23:271-287. February 11. + 1894. Third addition to a knowledge of the Batrachia and Reptilia of + Costa Rica. Proc. Acad. Nat. Sci. Philadelphia, 1894, pp. + 194-206. + +Duellman, W. E. + 1956. The frogs of the hylid genus _Phrynohyas_ Fitzinger, 1843. + Misc. Publ. Mus. Zool., Univ. Michigan, 96:1-47, pls. 1-6. + February 21. + 1967. Additional studies of chromosomes of anuran amphibians. Syst. + Zool., 16:38-43, March 17. + +Duellman, W. E. and Cole, C. J. + 1965. Studies of chromosomes of some anuran amphibians (Hylidae and + Centrolenidae). Syst. Zool., 14:139-143. July 9. + +Duellman, W. E. and Trueb, L. + 1966. Neotropical hylid frogs, genus Smilisca. Univ. Kansas Publ., + Mus. Nat. Hist., 17:281-375, pls. 1-12. July 14. + +Dunn, E. R. + 1931. The amphibians of Barro Colorado Island. Occas. Papers Boston + Soc. Nat. Hist., 5:403-421. October 10. + 1933. Amphibians and reptiles from El Valle de Anton, Panama. + _Ibid._, 8:65-79. June 7. + 1934. Two new frogs from Darien. Amer. Mus. Novit., 747:1-2. + September 17. + +Fouquette, M. J. Jr. + 1960a. Call structure in frogs of the family Leptodactylidae. Texas + Jour. Sci., 12:201-215. October. + 1960b. Isolating mechanisms in three sympatric tree frogs in the Canal + Zone. Evolution, 14:484-497. December 16. + +Gaige, H. T., Hartweg, N. and Stuart, L. C. + 1937. Notes on a collection of amphibians and reptiles from + eastern Nicaragua. Occas. Papers Mus. Zool., Univ. Michigan, + 357:1-18. October 26. + +Gosner, K. L. + 1960. A simplified table for staging anuran embryos and larvae with + notes on identification. Herpetologica, 16:183-190. + September 23. + +Kellogg, R. + 1932. Mexican tailless amphibians in the United States National + Museum. Bull. U.S. Natl. Mus., 160:1-224. March 31. + +Rivero, J. A. + 1961. Salientia of Venezuela. Bull. Mus. Comp. Zool., 126:1-207. + November. + +Smith, H. M. + 1951. The identity of _Hyla underwoodi_ Auctorum of Mexico. + Herpetologica, 7:184-190. December 31. + +Stejneger, L. + 1906. A new tree toad from Costa Rica. Proc. U. S. Natl. Mus., + 30:817-818. June 4. + +Stuart, L. C. + 1935. A contribution to a knowledge of the herpetology of a portion + of the savanna region of central Peten, Guatemala. Misc. Publ. + Mus. Zool., Univ. Michigan, 29:1-56, pls. 1-4. October 4. + +Taylor, E. H. + 1952. The frogs and toads of Costa Rica. Univ. Kansas Sci. Bull., + 35-577-942. July 1. + + +_Transmitted July 11, 1967._ + + + +Transcriber's Notes + +This file was derived from scanned images. With the exception of the +list of typographical errors that were corrected below, the original +text is presented. + +In the copy of the original, the Plate text contains the notation +"X 2" after the caption to let the reader know that the image was +enlarged by a factor of two. + + +Emphasis Notation + + _Text_ = Italic + + +Text+ = Bold + + +Typographical Errors Corrected: + +Several minor typographical corrections were made (missing periods, +commas, incomplete italicization, etc.); but are not indicated here. +More substantial changes are listed below: + +Page 533 - UMZ => UMMZ +Page 534 - Diganosis => Diagnosis +Page 544 - fontanells => fontanelle +Page 545 - prrimary => primary +Page 547 - band of of frequencies => band of frequencies +Page 550 - ad => had +Page 551 - clumbs => clumps +Page 552 - acount => account +Page 557 - Minchigan => Michigan + + + + + + + +End of the Project Gutenberg EBook of Middle American Frogs of the Hyla +microcephala Group, by William E. 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