HARVARD UNIVERSITY LIBRARY OF THE Museum of Comparative Zoology THE HYLID FROGS OF MIDDLE AMERICA THE HYLID FROGS OF MIDDLE AMERICA Volume 2 WILLIAM E. DUELLMAN Curator Division of Herpetology Museum of Natural History The University of Kansas MONOGRAPH OF THE MUSEUM OF NATURAL HISTORY, THE UNIVERSITY OF KANSAS NUMBER 1 1970 MUS. COMP. ZOOL L!BRArjY JAN u '^n HARVARD UNIVERSITY 4'^ ?r- ^\jo\l- <^*iA_^ ^ ..> - j\ ^ ( • H. dendroscarta /-^ [ \I / \ r / / ^ J -. \ 4 J / / y /V -^ ' \ \ ,X / o ^L ^' - \ \ Y I . 20 20° •\ .,--'*■■* — ""^'^ — ^/ / -^^'..> V/ , ( ^_ -. "- - '.-»./ / f r 1 ' ■"( i~ \ '- — — i / 1 \ -- > / ^ 1 / « ^-'^^ "-■_. J 1 } 1 "* V 1 \ 1 1 / ■ \ ! } \ '' "^7 \ / 16° N^__^ ^,_/^' n ~ — < ~(L ^S^ r ^ ^^^ 16° o ^O < \^ \ 1 r ^^^"^— -^ -> r ~~^^-^/'' ° ■ / ^^— -^ o ^-=5-^ 1 16° 0 50 100 200 1 1 1 1 1 /■ / / / 16° KILOMETERS 1 98° 94° Fic. 230. Distribution of the species in the Hi/Zn tacniopii.i group. immaculate creamy white or creamy yellow venters and a dark postorbital mark; baiidinii ha.s a blunt .snout in both sexes, whereas cijanosticia has an acuminate, but not pro- truding snout in both sexes, and blue spots on the flanks and posterior surfaces of the thighs. DESCRn'TiON: This is a moderately large frog; males attain a maximum snout-\ent length of 65.9 mm., and females reach 70.0 mm. In a sample of IS adult males from throughout the range in eastern Mexico, the snout-\ent length is 4S.0 to 65.9 (mean, 5S.0) mm.; the ratio of tibia length to snout-\ent length is 0.450 to 0.500 (mean, 0.480); the ratio of head length to snout-vent length is 0.290 to 0..340 (mean, 0.310); the ratio of head width to snout-vent length is 0.260 to 0.300 (mean, 0.2S0), and the ratio of the diam- eter of the tympanum to that of the eye is 0.510 to 0.620 (mean, 0.560). In eight fe- males from the same area, the snout-vent length is 56.6 to 70.0 (mean, 64.2) mm. In most proportions, the females do not differ from the males, but females do have a slightly larger tympanum in relation to the diameter of the eye; the tvmpanum/eye ratio in fe- males is 0.540 to 0.660 ( mean, 0.620) . The head is as wide as the bod\', and the top of the head is flat. In dorsal profile, the snout is acuminate in males and truncate in females. In lateral profile, the snout gradually slopes downward anterior to the nostrils and protrudes beyond the lower jaw in males; it is rounded and barely protruding in fe- 1970 DUELLMAN: HYLID FROGS 447 males (fig. 231). The snout is moderately long in both se.xes, and the nostrils are situ- ated at a point about two-thirds the distance from the eyes to the tip of the snout. The nostrils are noticeabi\' protuberant. The can- thus is angular; the loreal region is noticeably conca\e, and the lips are thick and barely flared. A heavy dermal fold extends posteri- orly from the posterior edge of the eye, above the tympanum, and downward to a point above the insertion of the arm. In all speci- mens, this dermal fold covers the upper edge of the t\'nipanum; in some others it also ob- scures the posterior edge. Otherwise, the t\'mpanum is distinct and is separated from the eye by a distance equal to the diameter of the tympanum. The eyes are large and protuberant. Fig. 231. Se.xual dimorphism in the shape of the snout in Hiila taeniopus. A. Male, K.U. No. 53825. B. Female, K.U. No. 53833. x 8. The arms arc long and moderately slender. There is no a.xillary membrane. A row of tubercles forms a dermal fold along the ven- trolateral edge of the forearm; in most speci- mens this fold continues onto the fourth finger. A distinct transverse dermal fold is present on the wrist. The fingers are moder- ately short and robust and bear large discs. The disc on the third finger is about half again the size of the tympanum. The sub- articular tubercles are large and round; the supernumerary tubercles are small and sub- conical. The outer palmar tubercle is tripar- tite. An elongate tubercle is present on the base of the pollex. The pollex is greatly en- larged and in breeding males bears a horny nuptial excrescence. The fingers are about one-third webbed (fig. 225B). The webbing is vestigial between the first and second fin- gers and extends from the middle of the penultimate phalanx of the second to the middle of the antepenultimate phalanx of the third and on to the base of the penultimate phalanx of the fourth finger. The hind limbs are moderately short and robust; the heels of the adpressed Hmbs overlap by about one- fourth the length of the shank. The tibiotar- sal articulation extends to the middle of the eye. A transverse dermal fold is present on the heel, and a strong tarsal fold extends the full length of the tarsus. The inner metatar- sal tubercle is large, flat, elliptical, and broadly visible from above. The outer meta- tarsal tubercle, if present, is indistinct. The toes are long and slender, and bear discs that are noticeably smaller than those on the fingers. The subarticular tubercles are large and conical. The supernumerary tubercles are small and conical. The toes are about three-fourths webbed (fig. 226B). The web- bing extends from the distal end of the penul- timate phalanx of the first toe to the base of the penultimate phalanx of the second, from the base of the disc of the second to the base of the penultimate phalanx of the third, from the distal end of the penultimate phalanx of the third to the base of the penultimate pha- lanx of the fourth and onto the base of the disc of the fifth toe. The anal opening is directed ventrally at the level of the ventral surfaces of the thighs. The anal tube is long and tuberculate. The 448 MONOGRAPH MUSEUM OF iNATURAL HISTORY NO. 1 skin on the dorsum, anterior and posterior surfaces of the thighs, and ventral surfaces of the shanks and tarsi is smooth; that on the throat, belly, and ventral surfaces of the thighs is granular. The tongue is ovoid, emarginate or barely notched posteriorly, and free behind for about one-fourth of its length. The dentigerous processes of the prevomers form transverse ridges between the small round choanae. Four to eight teeth are pres- ent on each prevomerine process. The total number of prevomerine teeth is nine to 16 (mean, 12.7 in males, and 13.3 in females). The vocal slits are small and are situated posterolaterally near the angles of the jaws. The vocal sac is single, median, subgular, and barely distensible. The general coloration of Hyla taeniopiis is green or bro\\'n with darker green or brown blotches on the body and trans\erse bands on the limbs (pi. 6i, figs. 1 and 2). The brown dorsal coloration is more common in females than in males, but even the darkest brown females are capable of changing to pale greenish tan with olive-green markings. Typical coloration of an adult female consists of a reddish brown dorsum with irregular darker brown markings middorsally. The flanks are dark brown with lemon yellow spots narrowly bordered by black. The belly is brownish black with yellow flecks, and the throat is silvery white. The typical coloration of an adult male is greenish tan above with darker olive-green blotches and brown flecks. The flanks arc pale greenish \'ellow with dark brown spots. The posterior surfaces of the thighs are dark brown; the ventral surfaces of the limbs, anterior surfaces of the thighs, and webbing of the feet arc pale gray. The belly and throat are dusty white with gray spots. The iris is bronze or grayish bronze in adults of both sexes. Usually males have pale greenish yellow or yellow flanks with dark green, brown, or black mottling; females usually have dark green, dark brown, or sometimes black flanks with yellow spots. All individuals have dark dorsal markings on the body and dark trans\erse bands on the limbs and feet. Usually there are three or four transverse bands on the shank and thigh, and four and five bands on the feet. A north-south trend in \ariation in color- ation is apparent. Comparisons are made be- low between three series: 1) Vicinity of Tianguistengo, Hidalgo, 2) Rio Octapa, near Tezuitlan, Puebla, and 3) Central Veracruz. In general, there is a tendency toward a dark- er venter, especially in females, from north to south. All females and most males from Hidalgo have immaculate creamy white ven- ters; some males have scattered brovsn or black spots on the chest and flanks. Some males from the Rio Octapa are immaculate below, but most individuals have some spot- ting on the flanks, chin, and chest; others have brown throats and brown laterally on the belly. Some females are immaculate below, but most ha\'e darkened \enters and brown flanks. Some males from Veracruz ha\e pale venters with scattered dark spots, but most ha\'e a dusty brown \'enters with darker brown flanks, where there is a distinct yellow spotting or marbling. Females have a darker brown venter, often with distinctive darker flanks with yellow spots. Ontogenetic changes in coloration also are apparent. A recently metamorphosed indi- \idual (K.U. No. 65062) with a snout- vent length of 23.6 mm. had a bright green dorsum with small black flecks on the head and body in life. The flanks were pale green and the \enter was immaculate pale yellow. The limb bands were dark brown, and the ills was metallic green. With increased size, there is a gradual change in dorsal coloration from many small black flecks to fewer large spots, w hich in many specimens are fused to form irregular blotches. Increased melanophore development on the flanks, especially in fe- males, results in dark mottling or spotting on yellowish flanks; in large females, the flanks may be dark brown or black with \ello\v spots. Increased melanophore development also occurs on the belly. Juveniles having snout-vent lengths of about 30 mm. have a few large black spots on the throat and chest. Indi\iduals having snout-vent lengths of about 45 mm. have large round, dark spots on the venter or an overall general darkening of the xcntral surfaces. All indi\iduals ha\ing snout-\ent lengths less than 36 mm., in addi- tion to a few larger specimens, have a metallic green iris in life. Subadults (37 to 50 mm.) have a pale bronze iris, sometimes with a 1970 DUELLMAN: HYLID FROGS 449 silvery or greenish tint. In adults, the iris is bronze, often with a noticeably darker, some- times copper-colored, periphery. Although no noticeable geographic varia- tion in size or structural features is apparent, there is an ontogenetic change in the shape of the snout in males. The snout is truncate in juveniles, but in young males, having snout- \ent lengths of about 40 mm., a slight protru- sion of the snout is noticeable. The sloping, protruding snout, characteristic of the adult males, is developed by the time the frogs reach a length of 50 mm. Furthermore, in juveniles, the webbing on the hand is barely evident, and the feet are only about one-half webbed. The amount of webbing increases with age to the condition previously described for the adults. Tadpoles: Four tadpoles of this species are available for study. The smallest speci- men (developmental stage 25) has a body length of 13.6 mm. and a total length of 31.1 mm. The most advanced tadpole (develop- mental stage 30) has a body length of 18.5 mm. and a total length of 51.2 mm. A typical tadpole in developmental stage 25 has a body length of 16.5 mm. and a total length of 46.2 mm. The body is moderately depressed, slightly wider than deep. In dorsal profile the snout is bluntly rounded, and in lateral profile acutely rounded. The eyes are small and directed dorsolaterally; the diameter of the eye is equal to about one-sixth the depth of the body. The nostrils are small, directed anterolaterally, and situated about midway between the eyes and the tip of the snout. The spiracle is sinistral; the spiracular open- ing is directed posterodorsally at about mid- length of the body. The anal tube is dextral and long. The caudal musculature is heavy and extends nearly to the tip of the tail. At midlength of the tail, the depth of the muscu- lature is much greater than the depth of either the dorsal or ventral fins. The dorsal fin does not extend onto the body; distally, the fins are rounded (fig. 228B). The body is brownish black and the caudal musculature is slightly paler. Melano- phores form dense brownish black spots on the tail, and xanthophores form a distinct orange-yellow edge to the dorsal fin. Irido- phores form silvery green flecks on the body. The iris is pale bronze. In preservati\e the body is dark grayish brown, darker vcntrally. The caudal musculature is grayish tan; the caudal fin is translucent. Numerous dark brown spots are present on the caudal muscu- lature and fin. The mouth is small, located anteroven- trally, and directed ventrally. The lips have a shallow lateral fold. Two rows of small papillae completely border the mouth; medial to these is a row of larger papillae. The upper beak forms a broad arch with robust lateral processes. The lower beak is massive and broadly V-shaped. Both beaks bear fine ser- rations. There are two upper and three lower rows of teeth. The two upper rows are about equal in length, and the second upper row is broadly interrupted medially. The lower rows are complete, shorter than the upper rows, and the third lower row is the shortest (fig. 229B). Mating Call: Hijla taeniopus has not been heard to call in the field; recordings of the call are not available. One individual kept in captivity uttered one loud groan-like note. On the basis of this one observation and the presence of vocal slits, it may be assumed that this species does possess a voice. How- ever, the significance of voice in the mating behavior is quite questionable. Natural History: Hijla taeniopus in- habits cloud forests characterized by moder- ately low temperatures and high humidity. Individuals have been found on vegetation along cascading mountain sti-eams at night and on elephant ear plants, Hhes, and arbo- real bromehads by day. Breeding apparently takes place in the dry season, when the streams are clear and relatively quiet. Adults in breeding condition have been found in December, January, and February. Tadpoles were obtained from a gravel- bottomed pool in a rocky stream in a cloud forest. As pointed out by Duellman ( 1965b, p. 164) breeding males of Hyla taeniopus have greatly enlarged testes. In the breeding sea- son, the testes essentially fill the body cavity, much in the same way eggs do in a gravid female. Breeding apparently takes place in streams having a steep gradient. The rapidly flowing water would have a tendency to wash 450 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 away the sperm as they were being emitted over the eggs. Consequently, the develop- ment of large testes capable of producing great quantities of sperm possibly is an adap- tation to insure fertilization. Remarks: Duellman ( 1965b ) discussed the taxonomic status of the names Hyla tae- nioptii, Hyla hromeliana, Hyla (lalqtiesti, and Hyla cyclomaculata. The striking differences between adults of each sex and juveniles of Hyla taeniopus has resulted in the application of four specific names to this species. The syntype of Hyla taeniopus examined by me (B.M.N.H. No. 1947.2.2.3.32) is a juvenile having a snout-vent length of 30.3 mm.; the holotype of Hyla hromeliana (F.M.N.H. No. 100075) likewise is a juvenile having a snout- vent length of 26.9 mm. Taylor (1948a) named Hyla proboscidea [=Hyla dalquesti ( Taylor, 1949a ) J on the basis of five adult males from Jico, Veracruz, and Taylor (1949c) named Hyla cyclomaculata on the basis of a single female from Huatusco, Vera- cruz, Mexico. As demonstrated by Duellman (1965b) the acquisition of a series of these frogs has provided the necessary material to demonstrate that only a single species is in- volved. Etymology: The tri\ial name taeniopus, is derived from the Latin taenia, meaning band and the Latin pes, meaning foot. The name alludes to the transverse bands on the limbs. DiSTiuBUTiON: Hyla taeniopus occurs at elevations between 1200 and 2100 meters on the Atlantic slopes of the Sierra Madre Ori- ental from northeastern Hidalgo, southward through eastern Puebla to central Veracruz, Mexico (fig. 230). See Appendix 1 for the locality records of the 72 specimens examined. Hyla aitipotens Duellman Hyla aitipotens Duellman, 1968a, p. 572 [holotype, K.U. No. 101001 from 37 kilometers north (by road) of San Gabriel Mixtepec, Oaxaca, Me.xico, ele\ation 1860 meter.s; William E. Duellman collector]. Diagnosis: This is a large frog (adults of both sexes attain snout-vent lengths of 75 mm.) having a green or tan dorsum with darker green or brown spots on the body and transverse bands on the limbs and with or without a middorsal dark line. The licad is narrow, and the snout is acuminate in both sexes. The skin is smooth on the dorsum. The venter is immaculate yellow, and a bronze-colored canthal stripe is present. Hyla taeniopus differs from aitipotens by having a blunt snout in females. larger head, and shorter legs (see table 41). Furthermore, the \entcr in taeniopus is creamy white to brown, not yellow, and taeniopus lacks a canthal stripe. Hyla chanequc differs from aitipotens by having a blunt snout in both sexes, a tuber- culate dorsum, and in proportions (see table 41). JuN'eniles of aitipotens can be confused with Hyla pinorum, some individuals of which ha\ e a dark middorsal line. Otherwise, pinorum differs from aitipotens by ha\ing a smaller tympanum, less webbing on the hands, and a short, truncate snout. Description: The maximum known snout- vent length in both males and females of this large species is about 75 mm. In a sample of five adult males from the type locality, the snout- vent length is 68.8 to 75.1 (mean, 70.7) mm.; the ratio of tibia length to snout-vent length is 0.526 to 0.558 (mean, 0.537); the ratio of foot length to snout-vent length is 0.452 to 0.481 (mean, 0.472); the ratio of head length to snout-vent length is 0.281 to 0.300 (mean, 0.292); the ratio ''of head width to snout-vent length is 0.303 to 0.313 (mean, 0.308), and the ratio of the diameter of the tympanum to that of the eye is 0.414 to 0.5.52 (mean, 0.506). Two adult females from the type locality have snout-\ent lengths of 69.4 and 75.3 (mean, 72.4) mm. The females differ from the males by having slightly longer legs and larger tympani; the ratio of tibia length to snout-vent length in the two females is 0.558 and 0.562 (mean, 0.560), and the ratio of the diameter of the tympanum to that of the eye is 0.533 to 0.630 (mean, 0.588 ) . Most known specimens of this species are immature; 19 individuals have snout-vent lengths of 31.6 to 50.1 mm. There are no significant diflerences in proportions in these small specimens from the adults. The head is relatixely small; it is not as wide as the body. The top of the head is flat or barely convex. The snout in dorsal profile is acuminate; in lateral profile the snout is acutely rounded and protruding be- 1970 DUELLMAN: HYLID FROGS 451 vond the tip of the lower jaw. The snout is moderately long; the nostrils are shghth^ pro- tuberant, directed dorsolaterally, and situated about two-thirds of the distance from the eyes to the tip of the snout. The canthus is angu- lar, and the loreal region is flat; the lips are thick barely flared. A heavy dermal fold ex- tends posteriorly from the posterior corner of the eve over the dorsal edge of the tym- panum and curves \entralh- to a point abo\e the insertion of the arm. The fold obscures the upper edge of the tympanum, which otherwise is distinct. The tympanum is pos- teroventral to the eye and separated from the eye by a distance slightly greater than the diameter of the tympanum. The arms are moderately long and robust. An abbre\ iated axillary membrane is present. A thin dermal fold is present on the ventro- lateral edge of the forearm, and a distinct, trans\erse dermal fold is present on the wrist. The fingers are moderately short and broad and bear large discs; the width of the disc on the third finger is greater than the diame- ter of the tympanum. The subarticular tubercles are large, round, and conical; none is bifid. The supernumerary tubercles are large and granule-like; they are present only on the proximal segments of the digits. The prepollcx is enlarged, but breeding males ap- parently do not de\elop nuptial excrescences. The fingers are about one-half webbed (fig. 225C). The webbing connects the first and second fingers at the le\el of the distal end of the antepenultimate phalanx, extends from the middle of the penultimate phalanx of the second finger to the middle of the antepenul- timate phalanx of the third, and between the bases of the penultimate phalanges of the third and fourth fingers. The hind limbs are long and slender; the heels of the adpressed limbs o\erlap by about one-half of the length of the shank. The tibiotarsal articulation ex- tends to a point between the eye and the nos- tril. A thin, transverse dermal fold is present on the heel; the tarsal fold is strong and ex- tends the full length of the tarsus. The inner metatarsal tubercle is relatively small, elon- gate, and barely visible from above. The outer metatarsal tubercle is small and conical. The toes are moderately long and slender; the discs are slightly smaller than those on the fingers. The subarticular tubercles are large, round, and subconical. The supernumerary tubercles are large, conical, and arranged in a single row on the proximal segment of each digit. The toes are about four-fifths webbed (fig. 226C). The webbing extends from the base of the disc of the first to the base of the disc of the second and onto the base of the penultimate phalanx of the third toe, from the base of the disc on the third to the base of the penultimate phalanx of the fourth and onto the base of the disc of the fifth toe. The anal opening is directed posteroven- trally at the midlevel of the thigh; the anal sheath is long and tubular. The skin is smooth on the dorsal surfaces of the body and limbs and on the \entral surfaces of the shanks; it is granular on the throat, belly, and ventral surfaces of the arms and thighs. The tongue is ovoid, widest posteriorly, and neither notched nor noticeably free behind. The dentigerous processes of the prevomers are robust transverse ridges between the small, ovoid choanae. The number of pre- \-omerine teeth on each process varies from five to 10; the total number of prevomerine teeth is 10 or 12 (mean, two specimens, 11.0) in females, and 13 to 18 (mean, five speci- mens, 15.0) in males. The vocal slits and a \ocal sac are absent. The general coloration of Htjla altipotens is tan or pale green above with darker brown or green spots on the back and transverse bands on the limbs (pi. 60, fig. 1 and pi. 61, fig. 3). The typical coloration of an adult male is pale green abo\e with slightly darker green spots. The dorsal surfaces of the upper arms and thighs are tan with green transverse bars. The upper surfaces of the forearms and shanks are green with darker green trans- verse bars. The feet, fourth and fifth toes, and third and fourth fingers are tan with brown transverse bars; the other fingers and toes are tan with brown flecks. The ventral surfaces are creamy yellow, brightest on the throat and chest. The flanks and anterior sur- faces of the thighs are bright creamy yellow with dark brown reticulations and spots. The posterior surfaces of the thighs and ventral surfaces of the hand and webbing on the hands and feet are yellowish tan. There is a narrow, tan labial stripe. Narrow, cream- 452 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 colored stripes are present on the ventrolateral edge of the forearms, along the outer edge of the foot, and above the anus. A bronze-col- ored stripe e.xtends the length of the canthus, along the edge of the upper eyelid, and onto the supratympanic fold. The iris is pale bronze with black reticulations and a faint, median, horizontal copper-colored streak. The pupil is horizontally elliptical with a ventral notch. The palpebral membrane is clear above and pale bluish green with faint brown reticula- tions below. In preservative, the dorsum is pale brown with many darker brown spots on the back and dark brown transverse bands on the limbs. The flanks are white with dark brown spots; the anterior surfaces of the thighs are creamy white with brown reticula- tions and the posterior surfaces of the thighs are dark brown with creamy yellow flecks. The stripe on the snout, canthus, edge of up- per eyelid, and supratympanic fold is tan; the ventral surfaces of the feet are brown, and the rest of the venter is creamy white. All individuals have creamy yellow ven- ters and yellow flanks and anterior surfaces of thighs with brown or black spots and mottling. Most of the adults were pale green with darker green spots, but one individual was a much darker olive-green, and one was uniform brown above with a dark brown mid- dorsal stripe. Most subadults (snout-vent lengths, 31.6 to 50.1 mm.) were pale reddish tan above with darker reddish brown bars on the limbs and blotches on the back. The side of the head is dark brown and the stripe along the canthus, edge of upper eyelid, and the supratympanic fold is yellowish tan. Some individuals had a dark brown middorsal stripe. The posterior surfaces of the thighs were a dull yellowish tan; yellow flecks were present in the larger individuals. The num- ber of transverse bands on each thigh and shank varies from five to eight. The white stripe above the anus and the stripe from the snout along the side of the head are invariably present. In some of the largest individuals, the brown reticulations on the anterior sur- faces of the thighs extend onto the ventral surfaces; in these specimens, brown flecks are present on the ventral surfaces of the shanks. Tadpoles: A typical tadpole in develop- mental stage 25 from 1.3 kilometers north- northeast of Juchatengo, Oaxaca, Mexico, has a body length of 13.8 mm. and a total length of 41.1 mm. The body is slightly depressed and noticeably wider than deep. In dorsal profile, the snout is blunth' rounded; in lateral profile, it slopes gently from the eyes to a point above the nostrils and is further in- clined to an abbreviated, truncate snout. The eyes are small and directed dorsolaterally. The nostrils are slightly protuberant and are situ- ated about midway between the eyes and the tip of the snout. The spiracle is sinistral; the spiracular opening is directed posterodorsally at a point below the midline and about two- thirds the distance of the length of the body. The anal tube is moderately long and dextral. The caudal musculature is moderately robust. The tail is long; the caudal fins are low. At midlength of the tail the depth of the caudal musculature is greater than the depth of either the dorsal or ventral fins. The dorsal fin barely extends onto the body. Terminally, the fins are rounded (fig. 228C). The body is dark brown or black; minute golden flecks arc present on the sides and belly. The caudal musculature is brown lat- erally and ventrally and dark brown, nearly black dorsally. Faint brown flecks are pres- ent on the fins. In preservative the coloration is much like that in life, except that the gold flecks have disappeared and that the caudal musculature is creamy tan, dark brown dor- sally. In life, the iris is pale gold. The mouth is small and ventral. A mod- erately deep lateral fold is present in the lips. The mouth is completely bordered by two or three rows of small papillae; additional pa- pillae are present in the lateral fold. The upper beak is robust and in the form of a broad arch with long slender lateral processes. The lower beak is broadly V-shaped. Both beaks bear moderately long, blunt serrations. There are two and three lower rows of teeth. The upper rows are equal in length and ex- tend nearly to the edges of the lips; the sec- ond upper tooth row is broadly interrupted medially. The lower rows are complete, ap- proximately equal in length, and all slightly shorter than the upper rows (fig. 229C). Mating Call: The absence of vocal slits 1970 DUELLMAN: HYLID FROGS 453 and apparent absence of a vocal sac strongly suggest that this species lacks a voice. Natural History: This is a stream-breed- ing species that inhabits cloud forests and pine-oak forests. All known specimens were found in the dry season. At that time, adults and juveniles alike were found in trees and bushes near streams. Tadpoles were obtained from quiet pools in rocky streams. Hyla altipotens is like Hijla taeniopus in having greatly enlarged testes. If the large size of the testes is correlative with increased production of sperm, the large size of the testes may be an adaptation for successful breeding in torrential streams (fig. 232). Two tadpoles were raised to metamorpho- sis. The tadpoles were obtained in develop- mental stage 25 on February 19, 1966; they metamorphosed on March 26, 1966. The young had snout-vent lengths of 17.5 and 19.7 mm. The dorsum was dark green ( capable of changing to dark brown). A dark brown Fig. 232. Ventral view of the viscera of a male Hyla altipotens (K.U. No. 101008) showing the large granular tests. X 5. stripe extended from the nostril to a point above the insertion of the arm. The canthal stripe was pale greenish bronze. The an- terior and posterior surfaces of the thighs, ventral surfaces of the limbs, and hands were dark yellow. The chin and belly were pale yellow. The iris was bronze medially and a coppery color peripherally. Remarks: Duellman (1965b, p. 166) list- ed a specimen ( T.C.W.C. No. 16184 ) of sup- posed Htjla chaneque from Los Fustes, 3 kilometers east of San Sebastian, Oaxaca, Mexico. Re-examination of this specimen re- veals that it is Hijla altipotens. Etymology: The specific name altipotens is Latin, meaning mighty, used in allusion to the supposed potentiality of fertilization by the production of vast quantities of sperm in large testes. Distribution: Hyla aliipotens occurs on the Pacific slopes of the Sierra Madre del Sur in Oaxaca, Mexico, where it lives in cloud for- ests and pine-oak forests at elevations between 1100 and 1900 meters (fig. 230). See Appendix 1 for the locality records of the 31 specimens examined. The Hyla histincta Group Definition: The members of this group are medium-sized, stream-breeding species; males attain a maximum snout-vent length of 54 mm. and females, 56 mm. No marked sexual dimorphism in size is evident. Frogs in this group are rather drab in appearance. The dorsum is dull green, gray, yellow, or various shades of brown. The most distinctive aspect of coloration is the different color pat- terns on the flanks and posterior surfaces of the thighs. The flanks in all species are spot- ted or reticulated. The palpebral membrane is clear. The fingers are long and have little webbing (figs. 2.33 and 234), and the toes are at least two-thirds webbed (figs. 234 and 235 ) . A broad, flat, ossified prepollex is pres- ent but does not project as a spine. The skin of the dorsum is thick and glandular, but not tuberculate, in all but charadricola and chryses, in which it is thin. Dermal fringes and appendages are lacking on the limbs; an axillary membrane is present in charadricola, and chryses, and a thoracic fold is present in 454 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 pacliyderma, robertsortim, and siopela. The skull is moderately well ossified. The fronto- parietals arc widely separated medially throughout their lengths, and a large fronto- parietal fontanelle is present (fig. 2.36). The quadratojugal is reduced or absent; the max- illary does not articulate with the quadrato- jugal. The anterior arm of the squamosal does not extend more than one-half of the distance to the maxillary; the posterior arm of the s(juamosal is short, and the \entral arm is robust. The pterygoid is robust, and the medial ramus articulates with the prootic. The prevomers are unusually small and delicate. Teeth are present on the maxillary, premaxil- lary, and prevomer. Those on the maxillar\' and premaxillary are rather long, bifid, and moderately spatulate; some of the teeth on Fig. 2.33. liancLs of species in the Ht/hi bistincta group. A. H. bialincta. K.U. No. 6S077. B. il. pentheter. K.U. No. 100932. C. H. robcrtsorum, K.U. No. 71266. D. H. pachyderma, U.S.N. M. No. 115028. E. H. siopela, K.U. No. 100981. F. H. crassa, U.I.M.N.H. No. 2.5050. x 4. 1970 DUELLMAN: HYLID FROGS 455 Fig. 234. Hands and feet of species in the Htjla bistincta group. A and E. Hyla charadricola, K.U. No. 58414. B and F. Hyla chryses, K.V. No. 106306. C. Hyla bistincta, K.U. No. 68077. D. Hyla pentheter, K.U. No. 100932. X 4. 456 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fig. 235. Feet of species in the Hi/la histmcta group. A. H. rohertsorum, K.U. No. 71266. B. H. pachyderma, U.S.N. M. No. ir502S. C. H. siopcla, K.U. No. 100981. D. H. crassa, U.I.M.N.H. No. 25050. x 4. 1970 DUELLMAN: HYLID FROGS 457 the priMiiaxillaiy and anterior part of the pre- ina.\illar\' are hooked. The prcvonierine teeth are spatulate and bifid. The tadpoles ha\'e a long, terminally rounded tail and a small \en- tral mouth with lateral folds, two complete rows of fringing papillae and at least one additional irregular row medially, and two upper and three lower rows of teeth (figs. 237 and 238). Only one species, bistincfa, is known to have a voice; the other species usually lack vocal slits. The haploid number of chromosomes is 12 (known only in pcnthe- ther and rohertsorum) . Composition: Nine species (H. Insfincta, hogertae, crassa, charadricola, chnjses, pachij- derma, pentheter, rohertsorum, and siopela) comprise this group, which is endemic to the Mexican highlands. All of the species, as now recognized, are monotypic. Of the nine spe- cies, 386 preserved frogs, 15 skeletons, and eight lots of tadpoles were examined. Comments: Duellman (1964b) included five species in the histincia group. Adler (1965) named chnjses and pentheter and in- cluded them in the bistincta group. Duell- man ( 1968a ) named siopela in the group, and Straughan and Wright (1969) named hoger- tae. Members of the Hyla bistincta group in- habit mountain streams, and the e\'olutionary trend within the group is towards more aquat- ic habits in the adults. The tadpoles are moderately well-adapted for development in streams, but they show no advanced speciali- zations. Hyla bistincta, the least specialized frog in the group, has relatively short fingers with a moderate amount of webbing, a high, truncate snout, and vocal slits. Hyla charadricola and chryses apparently are closely related and represent a di\'ergence from the main evolutionary line within the group. These two species have relatively thinner skin on the dorsum, more slender bod- ies, an axillary membrane; furthermore, breeding males apparently lack nuptial ex- crescences. Both species lack vocal slits. Of the remaining .species in the group, pentlicter most closely resembles bistincta. but differs in having longer fingers and usual!)' no vocal slits. The other five species in the group (hogertae, crassa, pachyderma, rohertsorum, and siopela) are the most advanced. They ]ia\e siiort, blunt heads, thick glandular .skin, long fingers with little webbing, large webbed feet, nuptial excrescences in breeding males, and no \'Ocal slits or axillarv membranes (ta- ble 44). The frogs in the Hyla hi.stincta group pre- sent a classic picture of montane distribution. The most primitive species is the most wide- spread and is the only one that occurs sym- patrically with other species in the group. Hyla chryses occurs in the Sierra Madre del Sur in Guerrero, and charadricola occurs in the high mountains of Hidalgo. Hyla penthe- ter lives in the Sierra Madre del Sur in Oaxaca whereas rohertsorum, pachyderma, siopela, and crassa occur in that order from north to south in the Sierra Madre Oriental, and hoger- tae occurs in the Sierra Madre del Sur of Oaxaca. Each of the last four species is known from a single stream. Frogs in the Hijla bistincta group presum- ably are closely related to those of Plectro- hyla, inhabitants of montane streams in Nu- clear Central America. The two groups of species are alike in the absence of a quadra- tojugal, presence of thick, glandular skin, structure and form of the tadpoles, and in general appearance. They show parallel pro- gressive modifications for a stream existence in the lengthening of the fingers, reduction of webbing on the hand, and loss of vocal slits and a voice. The species in both groups have a broad, ossified prepollex; in Plectrohyla, the prepollex has one or more projecting spines. Plectrohyla is singularly distincti\'e in having robust premaxillaries with bifurcate alary processes. Hyla bistincta Cope Htjla bistincta Cope, 1877, p. 87 [holotype, U.S.N.M. No. 32261 from "most probably Veracruz," Me.xico; Francis Sumichra.st collector; type locality restricted to Acu'tzingo, \'eracruz, Mexico, by Smith and Taylor (1950, p. 346]. Broechi, 1882,' p. 43. Boulenger, 1882a, p. 401. Gunther, 1901 (1885- 1902), p. 265. Kellogg, 1932, p. 163. Smith and Taylor, 1948, p. 87. Duellman, 1964b, p. 475. Hijla bistincta laljcculata Shannon, 1951, p. 470 [holotspe. U.S.N.M, No. 123689 from San Lucas Caniotliin, Oaxaca, Me.xico; Walter S. Miller collector]. Hyla bistincta bistincta: Shannon, 1951, p. 472. Diagnosis: Hyla bistincta is a moderately large species with a truncate snout in dorsal 458 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fig. 2.36. Dorsal and lateral views of the skulls of two species in the Hijla bistincta group. A and B. H. charadricola, K.U. No. 55624. C and D. H. siopela, K.U. No. 117428. x 5. TABLE 44 Comparison of Sizes and Certain Proportions, with Means in Parentheses, of Males of the Species in tiie Ihjla bistincta Group. Species N H. bistincta 38 H. pentheter 7 //. clmradricola ._ 10 H. chrtjses 3 H. robertsorum 24 H. pacJnjderma 1 H. siopela 7 H. crassa 1 H. boaertae 1 Snout- vent Length Tibia Length/ S-V L Head Width/ S-V L Tympanum/ Eye 43.0-53.8 0.470-0.520 0.320-0.370 0..350-0.480 (46.3) (0.490) (0.340) ( 0.420 ) 43.3-52.1 0.502-0.525 0.350-0.381 0.520-0.586 (46.2) (0.514) (0.367) (0.555) 35.3-44.4 0.500-0.540 0.310-0..330 0.300-0.370 (40.4) (0.520) (0.320) (0..340) 36.3-37.3 0.494-0.498 0.313-0.327 0.595-0.634 (37.1) (0.496) (0.320) (0.574) 39.9-47.9 0.480-0.510 0.300-0.360 0.360-0.470 (43.1) (0.490) (0..320) (0.410) 39.9 0.530 0.320 42.1-46.2 0.472-0.500 0.291-0.317 0.363-0.468 (44.4) (0.487) (0.309) (0.438) 53.7 0.500 0.330 0.438 44.9 0.508 0.369 0.319 1970 DUELLMAN: HYLID FROGS 459 Fig. 237. Tadpoles of the species in the H. histincta group. A. H. pentheter, K.U. No. 104142. B. H. bistincta, U.M.M.Z. No. 115231. C. H. robertsonim, K.U. No. 60078. D. H. siopcia, K.U. No. 100118. A., X 4; other, X 2. profile. The fingers are short and about one- third webbed; the toes are about two-thirds webbed. A strong tarsal fold is present, but a thoracic fold and axillary membrane are ab- sent. The anal opening is at the level of the ventral surfaces of the thighs. Vocal slits and nuptial excrescences are present. The flanks and posterior surfaces of the thighs are creamy tan with brown reticulations or spots. Hijla pentheter resembles bistincta but differs in having longer fingers with less webbing and by lacking vocal slits and reticulations on the posterior surfaces of the thighs. Hyh siopeh differs in having a vertical rostral keel, less webbing, and no vocal slits. Other members of the Ihjia histincta group either have thin- ner, less glandular, skin and an axillary mem- brane and no nuptial excrescences, or they have round snouts in dorsal profile and long fingers \\ith little or no webbing. Aside from members of this group, bistincta cannot be confused with any other Middle American hylids, except Plectrohyla. Description: This is a moderate-sized species of the Hyla bistincta group; males at- tain a maximum snout-vent length of 53.8 mm. and females reach 67.6 mm. In a series of 19 males from Uruapan, Michoacan, Mex- ico, the snout-vent length is 43.0 to 48.7 (mean, 45.9) mm.; the ratio of tibia length to snout- vent length is 0.470 to 0.515 (mean, 0.492); the ratio of foot length to snout- vent length is 0.425 to 0.485 (mean, 0.452); the ratio of head length to snout-vent length is 0.2S9 to 0.:32.3 (mean, 0.306); the ratio of head width to snout-\'ent length is 0.317 to 0.364 (mean, 0.342), and the ratio of the diameter of the tympanum to that of the eye is 0.339 460 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 '«*^tf(n;J,AM>^>^^''^**^ ''i-"j5«t; Hitt'yaWtJ't^ii^' .>^ ?■ # Fic. 238, Mouths of tadpoles of the species in the H. bistincta group. .'\. //. pcnihetcr, K.U. No. 104112. B. H. bistincta, U.M.M.Z. No. 115231. C. H. rubertsorum, K.U. No. 60078. D. H. siopcla, K.U. No. 110118. A., X 20; others, X 10. to 0.479 (mean, 0.422). Two females from the same locality have snout-vent lengths of 43.8 and 51.4 mm.; in these specimens the ratio of the diameter of the tympanum to that of the eye is 0.440 and 0.420, respecti\ely. The head is as wide as, or slightly wider than, the body; the top of the head is slightly convex, and the eyes are large. In dorsal profile the snout is truncate; in lateral profile it is bluntly rounded. The snout is moderately short; the nostrils are barely protuberant and are at a point about two-thirds the distance from the eyes to the tip of the snout. The canthus is rounded, the loreal region is slight- ly concave, and the lips are thick and barely flared. A heavy dermal fold extends posterior- ly from the eye aboxe the tympanum, and downward to a point above the insertion of the arm. The fold obscures the upper edge of the tympanum, which otherwise is distinct and is separated from the eye by a distance slightly greater than the diameter of the tympanum. The arms are moderateh' long and robust; an abbre\'iated axillary membrane is present. A few small tubercles are present on the ven- tral surface of the forearm, but these do not form a distinctive row along the ventrolateral edge of the forearm. A heavy transverse der- mal fold is present on the wrist. The fingers are moderately short and stout and bear mod- erately large discs; the disc on the third fin- ger is about equal in size to the tympanum. The subarticular tubercles are large and round; none is bifid. The supernumerary tu- bercles are rather large and round; they are arranged in a single row on the proximal seg- ments of each digit. The outer palmar tu- bercle is low and rounded; in most specimens it is bifid or tripartite, and in some it is frag- mented into one large and one or two small tubercles. An elongate tubercle is present on the prepollex, which is greatly enlarged. Nup- tial excrescences, in the form of minute spin- ules are present on the prepollex, inner edge of the thumb, and inner edge of the second finger in breeding males. The webbing is ves- tigial between the first and second fingers and extends from the middle of the antepen- ultimate piialanx of the third to the distal end of the antepenultimate phalanx of the fourth finger (fig. 233A). The hind limbs are mod- 1970 DUELLMAN: HYLID FROGS 461 erately long and robust; the heels of the ad- pressed limbs overlap b>' about one-third of the length of the shank. The tibiotarsal ar- ticulation extends to the anterior corner of the eye. A trans\erse dermal fold is present on the heel, and the tarsal fold is moderately strong and extends the full length of the tar- sus. The inner metatarsal tubercle is large, elliptical, and has a raised median edge. The outer metatarsal tubercle is small, and conical. The toes are moderately long and slender and bear discs that are nearly as large as those on the fingers. The subarticular tubercles are large, round, and subconical. The supernu- merary tubercles are large, low, and arranged in a single row on the proximal segments of each digit. The toes are about three-fourths webbed" (fig. 234C). The webbing extends from the distal end of the penultimate pha- lanx of the first toe to the base of the penulti- mate phalanx of the second, from the base of the disc of the second to the base of the pen- ultimate phalanx of the third, from the base of the disc of the third to the distal end of the antepenultimate phalanx of the fourth and on to the base of the disc of the fifth toe. The anal opening is directed ventrally at the level of the ventral surfaces of the thighs in males and at the level of the middle of the thighs in females. The anal tube is long and curved downward. The skin on the ventral surfaces of the body and thighs is granular; elsewhere the skin is smooth. The tongue is broadly cordiform, shallowly notched behind, and free posteriorly for about one-fourth of its length. The prevomerine teeth are situ- ated on small, high, transverse elevations be- tween the small, ovoid choanae. Males have three to seven teeth on each prevomerine process and a total of six to 14 (mean, 9.8) teeth. Females have five to seven teeth on each process and a total of 10 to 13 (mean, 11.5) teeth. The vocal shts are situated along the inner edge of each ramus of the lower jaw. The vocal sac is single, median, sub- gular, and barely distensible. The general coloration of Hijla histincta is pale tan to dark brown dorsally with creamy yellow flanks with brown spots or reticulations (pi. 62, figs. 1 and 2). The posterior surfaces of the thighs are tan or brown with faint yellow spots. The flanks are cream with bold or fine dark reticulations that tend to enclose yellow spots. The ventral surfaces are a pale yellow. The iris is a pale copper color. There is considerable variation in color in the li\ing frogs. The dorsum varies from greenish tan to pale yellowish tan to reddish Ijrown, and in some individuals, dark choco- late brown. In the series of specimens from Uruapan, Michoacan, Mexico, the coloration of the flanks and the anterior surfaces varies from nearly uniform creamy yellow with only fine dark reticulations to bold reticulations enclosing yellow spots. In some specimens from Oaxaca and Veracruz, the markings on the flanks consist of irregular spots or dashes, instead of reticulations. In preservative, the dorsum is various shades of brown, and the ventral surfaces are creamy white. The flanks and anterior sur- faces of the thighs are creamy white with dark brown reticulations, and the posterior sur- faces of the thighs are tan or brown with creamy white spots. Tadpoles: Duellman (1961c, p. 47) pre- sented a description of the tadpoles of this species from Uruapan, Michoacan, Mexico. Tadpoles are available in developmental stages 25 through 36; the smallest tadpole has a total length of 33.0 mm., and the largest (developmental stage 36) has a total length of 61.0 mm. A typical tadpole in develop- mental stage .34 has a body length of 19.4 mm. and a total length of 57.6 mm. The body is moderately depressed, as wide as deep. In dorsal profile the snout is broad and rounded; in lateral profile the snout is rounded. The nostrils are small, directed anteriorly, and situated about midway between the eyes and the tip of the snout. The eyes are small and directed dorsolaterally. The spiracle is sinis- tral; its opening is on the midline at a point about two-thirds of the distance from the snout to the posterior edge of the thighs. The anal tube is short and dextral. The tail is about twice as long as the body with heavy musculature and relatively shallow fins. The caudal musculature does not extend to the tip of the tail. At midlength of the tail, the depth of the musculature is about equal to that of either the dorsal or ventral fin. The dorsal fin extends onto the body. Terminally, the cau- dal fins are rounded (fig. 2.37B). 462 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 In preservative the body is pale grayish brown dorsally and Literally and pale gray ventrally. The caudal musculature is brown and the fins are translucent with scattered melanophores. The color in life is not known. The mouth is ventral and moderately large; its width is equal to about two-thirds of the greatest width of the body. The lips are folded laterally; two rows of small papillae completely border the lips. A row of larger papillae is present between the upper lips and the first upper row of teeth, and a simi- lar row is present between the lower lips and the third lower row of teeth. Laterally, these rows of large papillae degenerate into small papillae in the lateral fold. The beaks are moderately robust and bear small peg-like serrations which are slightly larger on the lower beak. The upper beak is a broad arch with short, rounded lateral processes; the lower beak is broadly V-shaped. There are two upper and three lower rows of teeth. Tlie upper rows are nearly equal in length and slightly longer than the lower rows, which arc subequal in length. The second upper tooth row is narrowly interrupted medially in all specimens, and the first lower row is in- terrupted in about half of the specimens (fig. 238B). Mating Call: No recordings of the call of Hyla fmtincta exist. Shannon ( 1951. p. 473) remarked that the type specimen of Hyla bi- stincta lahectilata was singing when caught. At Uruapan, Michoacan, I heard a low growl- like call that possibly was produced by Hyla histincta, but I did not trace the call to a frog. Natural History: Hyla bistincta is an in- haliitant of pine-oak, pine-fir, and pine forests in the high mountains of Mexico. Individuals usually are found near streams. At Uruapan and at Dos Aguas, Michoacan, individuals were found by day clinging to roots and vines in heavily shaded areas immediately over cascading streams. At night, the frogs were sitting on rocks and low vegetation near the stream. Tadpoles were found in gravel-bottomed pools in torrential streams. A recently meta- morphosed indi\idual has a snout-vent length of 24. S mm. Remarks: Duellman (1964b, p. 477) dem- onstrated that Hyla bistincta labeculata was an unrecognizable subspecies. He reported that in general, specimens from western Mex- ico have reticulate mottling on the fianks as compared with the marbling on the flanks in specimens from eastern Mexico. The subspe- cies, labeculata was diagnosed by Shannon (1951, p. 470) as differing from the nominate subspecies by having "the gray reticulations of the sides entirely broken up into elongate black blotches; tarsal fold moderately ele- vated." Smith and Williams ( 1963, p. 23) reported a specimen of Hyla bistincta from San Vin- cente, Oaxaca. Duellman (1964b, p. 477) in- cluded this record in his account of Hyla bistincta. Examination of that specimen (U.I.M.N.H. x\o. 51346) revealed that it is a poorly preserved specimen of Hyla pinonim. Duellman (1964b, p. 478) also included a specimen (A.M.N.H. No. 13447) from Pluma Hidalgo, Oaxaca, in Hyla bistincta. Re-exam- ination of that specimen reveals that it is a small indi\idual of Hyla pentheter. Etymology: The specific name bistincta is derived from the Latin bis meaning twice and from the Latin tinctus, meaning paint or color; the name refers to the distincti\e darker coloration of the flanks as compared with the paler dorsum. Distribution: Hyla bistincta occurs at elevations from 1400 to 2S00 meters in the mountains of the Sierra Madre Occidental in southwestern Durango southward through the Cordillera Volcanica in Michoacan, \Iexico, and Morelos, in the Sierra de Coalcoman in Michoacan, and in the Sierra Madre del Sur in Guerrero; this species also occurs in the Sierra Madre Oriental from central Veracruz to central Oaxaca (fig. 239). See Appendix 1 for the locality records of the 114 specimens examined. Hyla penthether AdJer Ilijla pentheter .\dler, 1965, p. 5 [holotype, U.M.M.Z. No. 125381 from 37 kilometers north (by road) of San Gabriel Mi.xtepec, Oaxaca, Me.vico, ele- \ation 1860 meters; Kraig .^dler collector]. Diagnosis: This is a moderately large species with a truncate snout in dorsal pro- file, thick, glandular skin, long fingers with 1970 DUELLMAN: HYLID FROGS 463 • H bistincto O H pentheter 20' 0 100 200 KILOMETERS 20° 106° 102 Fig. 239. Distribution of Vl\.j\a bistincta and Hyla pentheter. only \-estigial webbing, a distinct tarsal fold, anal opening at level of ventral surfaces of thighs, and nuptial excrescences present in breeding males. Hyla pentheter lacks vocal slits, axillary membranes, and a thoracic fold. The dorsum is pale tan, yellow, or gray. The sides of the head, flanks, and inner and outer edges of the limbs are dark brown. Hyla bistincta resembles pentheter but differs by ha\ing vocal slits, shorter fingers that are about one-third webbed, and reticulations or spots on the flanks and posterior surfaces of the thighs. Hyhi siopela has a vertical rostral keel and no dark brown color on the sides of the body. Other species in the Hyla bistincta group either ha\'e thinner, less glandular skin, an axillary membrane, and no nuptial excres- cences, or they have round snouts in dorsal profile. The only other Middle American hy- lids with which pentheter can be confused are species of Plectrohyla. Description: This is a moderately large species; males attain a maximum snout-vent length of 52.1 mm., and females reach 56.4 mm. In a series of seven males from the Pa- cific slopes of the Sierra Madre del Sur in Oaxaca, Mexico, the snout-vent length is 43.3 to 52.1 (mean, 46.2) mm.; the ratio of tibia length to snout-vent length is 0.502 to 0.525 (mean, 0.514); the ratio of foot length to snout- vent length is 0.431 to 0.460 (mean, 0.447); the ratio of head length to snout-vent length is 0.350 to 0.381 (mean, 0.367), and the ratio of the diameter of the tympanum to that of the eye is 0.520 to 0.586 (mean, 0.5.55). Two females from the same area have snout- vent lengths of 56.0 and 56.4 mm. They show no differences in proportions from the males. 464 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 The head is as wide or slightly wider than the body, and the top of the head is flat. In dorsal profile the snout is truncate; in lateral profile it is acutely angular just anterior to the nostrils and barely rounded at the margin of the lip. The snout is short; the nostrils are slightly protuberant and situated about three- fourths of the distance from the eyes to the tip of the snout. The canthus is rounded; the loreal region is slightly concave, and the lips are thick and flaring. A heavy dermal fold extends posteriorly from the eye, above the tympanum, and downward to the point of insertion of the arm. The fold obscures the upper edge of the tympanum, which otherwise is distinct and separated from the eye by a distance equal to about two-thirds of the diameter of the tympanum. The arms are moderately long and robust; no axillary membrane is present. Tubercles are absent from the ventrolateral edge of the forearms, but a distinct transverse dermal fold is present on the wrist. The fingers are long and moderately slender and bear rather large discs; the disc on the third finger is slightly smaller than the diameter of the tympanum. The subarticular tubercles are large, round, flat on the third and fourth fingers, and coni- cal on the first and second. The supernu- merary tubercles are large, conical, and ar- ranged in a single row on the proximal seg- ments of each digit. None of the tubercles is bifid. A large, flat, bifid palmar tubercle is present, and an elongate tubercle is present on the prepollex. The prepollex is greatly en- larged, and in breeding males bears a nuptial excrescence composed of minute spinules; spinules are present on the inner surfaces of the thumb and second finger. A vestige of a web is present between the first and second and third fingers, whereas the web extends from the base of the antepenultimate phalanx of the third to the middle of the antepenulti- mate phalanx of the fourth finger (fig. 2.33B). The hind limbs are moderately long and ro- bust; the heels of the adpressed limbs o\erlap by about one-third of the length of the shank. The tibiotarsal articulation extends to the pos- terior corner of the eye. A faint transverse dermal fold is present on the heel, and the tarsal fold, which is indistinct in some speci- mens, extends the full length of the tarsus. The inner metatarsal tubercle is large, ovoid, elevated, and barely visible from above. The outer metatarsal tubercle is small and conical. The toes are long and slender and bear discs that are slightly smaller than those on the fingers. The subarticular tubercles are large and conical; the supernumerary tubercles are small, conical, and arranged in a single row on the proximal segments of each digit. The toes are about three-fourths webbed (fig. 2.34D). The webbing extends from the distal end of the penultimate phalanx of the first toe to the base of the penultimate phalanx of the second, from the base of the disc of the second to the distal end of the antepenulti- mate phalanx of the third, from the distal end of the penultimate phalanx of the third to the distal end of the antepenultimate phalanx of the fourth and on to the base of the disc of the fifth toe. The anal opening is directed \entrall\- at the midlevel of the thighs; a long anal sheath is present. The skin on the ventral surfaces of the body and posteroventral surfaces of the thighs is granular; elsewhere the skin is smooth. The tongue is broadly cordiform, shallowly notched posteriorly and barely free behind. The dentigerous processes of the prevomers are short transverse ridges between the small, round choanae. Males have three to six teeth on each process and ha\e a total of seven to 11 (mean, 8.7) teeth. Females have six to eight teeth on each process and a total of 13 to 15 (mean, 14.0) teeth. \'ocal slits are absent in most specimens. The general coloration of Hylu pentlieter is grayish brown or yellowish brown above with dark brown on the sides of the head, body, and limbs (pi. 62, figs. 3 and 4). The dorsum is uniformly colored and varies from a pale gra>ish brown to yellowish tan or yel- low with a slight greenish tinge. A broad, dark, chocolate brown band extends from the edge of the upper lip below the nostril to the eye, along the supratympanic fold and the side of the body. The flanks are dark brown with yellow spots. The anterior and posterior surfaces of the thighs, outer edges of the shanks and feet, and \entrolateral edges of the forearm are dark brown. Large yellow spots are present on the anterior and posterior sur- faces of the thighs. The venter is yellow. 1970 DUELLMAN: HYLID FROGS 465 darkest on the throat and chest. The throat is mottled with olive-brown in some speci- mens. The iris is reddish copper with black reticulation.s, and the palpebruni is clear. The nuptial excrescences are dark brown. Adler (1965, p. 8) stated: "Metachrosis is moderate. When the dorsum is light grayish tan (pi. IE) the brown band along the side of the body is bordered above by a thin whit- ish-tan line." This line was evident in li\ing specimens of all colors, but was most promi- nent in those individuals having a darker dorsum. In some specimens having grayish brown or yellowish tan dorsal color, small dark brown flecks are evident on the dorsum. In preservative, the dorsum is gray or dull brown with or without small brown flecks. The \enter is creamy yellow. The dark lateral markings are black or dark brown; and the spots on the flanks are dull yellow. T.\DPOLES: A series of tadpoles in develop- mental stages 25 to 27 are available from a small stream 37 kilometers north of San Ga- briel Mi.xtepec, Oaxaca, Mexico. A typical tadpole in developmental stage 25 has a body length of 9.2 mm. and a total length of 26.4 mm. The body is as wide as deep, only mod- erateh' depressed. In dorsal profile the snout is bluntly rounded; in lateral profile it is gent- ly rounded. The nostrils are small, directed anterodorsally, and situated slightly closer to the eyes than to the tip of the snout. The eyes are small and directed dorsolaterally. The spiracular opening is directed posterodorsally just below the midline at a point about two- thirds of the distance from the snout to the posterior edge of the body. The anal tube is short and dextral. The tail is about twice as long as the body and is shallow. The caudal musculature is moderately deep and does not extend to the tip of the tail. At midlength of the tail, the depth of the musculature is slight- ly less than the depth of either the dorsal or ventral fin. The dorsal fin barely extends onto the body and is deepest just posterior to the midlength of the tail. The ventral fin is of equal depth throughout its length; terminal- ly the fins are pointed (fig. 237A). The body is dark brown with greenish yellow flecks dorsally, golden flecks laterally and white flecks ventrally. The dorsal edge of the caudal musculature is dark brown and orange. The fins are transparent with faint brown flecks. The iris is bronze. In preserva- tive, the body and caudal musculature are pale creamy tan. The top of the body, a small area anterovential to the eye, and a large blotch on the side of the body posterior to the eye are dark brown. The dorsal edge of the caudal musculature is dark brown. Small brown flecks are scattered on the caudal mus- culature and on the dorsal fin. The mouth is ventral; its width is equal to about two-thirds the width of the body. A lateral fold is present in the lips. Two rows of small papillae border the lip anteriorly and posteriorly, and a single row is present later- ally. A row of larger papillae is present be- tween the fringing papillae and the first upper tooth row, and a row of large papillae is pres- ent between the fringing papillae and the third lower tooth row; laterally in the lateral fold the large papillae degenerate into scat- tered smaller papillae. The beaks are robust and bear large, blunt serrations. The upper beak forms a broad arch with laterally direct- ed, short, blunt, lateral processes. The lower beak is V-shaped. There are two upper and three lower rows of teeth. The upper rows are about equal in length and slightly longer than the lower rows. The second upper row is narrowly interrupted medially (fig. 2S3A). M.-\TiNG Gall: As indicated by the ab- sence of the vocal slits, this species apparent- ly lacks a voice. Natural History: Hyla pentheter inhabits humid montane pine-oak forest, where it lives in the vicinity of cascading mountain streams. Adler (1965, p. S) found a female "in the afternoon on the forest floor near the base of a clift, a dozen meters from the nearest stream." He found males on vines and twigs or on moss-covered boulders near or over the stream. At the type locality, I obtained indi- \iduals of both sexes from low trees near a stream at night, and one male was found on a boulder in the stream. One individual was found on a boulder in a stream 29 kilometers south-southeast of Juchatengo, Oaxaca. The tadpoles were found in a pool in the stream, where they laid quietly on the bottom, but when disturbed took refuge on the bottom in the leaf litter. Remarks: Duellman and Cole (1965, p. 466 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 141 ) reported the number of chromosomes in Hijla hhtincta. Re-examination of the speci- men from which the chromosome prepara- tions were made proves that it is an exam- ple of Hijla pentlieter. A faded juvenile (A.M.N.H." No. 13447) from Pluma Hidalgo, Oaxaca, was listed as Htjia bistincta by Duell- man (1964, p. 478). Etymology: The specific name penfheter is Greek, meaning mourner and is used in allusion to the black border of the body, a symbol of mourning. Distribution: Hyla pentlieter is known only from elevations between 1500 and 2000 meters on the Pacific slopes of the Sierra Madre del Sur in southern Oaxaca, Mexico (fig. 239 ).'■•' See Appendix 1 for the locality records of the 12 specimens examined. Hyla charadricola Duellman Htjla characricola Duellman, 1964b, p. 478 [holo- type, K.U. No. 58414 from the Rio Totolapa, 14.4 kilometers ( by road ) \\est of Huachinango, Puebla, Mexico, elevation 2280 meters; John Wellman collec- tor]. Diagnosis: This is a medium-sized species (maximum snout- vent length in males, 44.4 mm.) witli a truncate snout, relatively thin skin on the dorsum, and an axillary mem- brane. The dorsum is olive-green with black reticulations, and the flanks are grayish green with brown spots. Vocal slits, nuptial excres- cences, and a thoracic fold are lacking, and the anal opening is at the level of the middle of the thigh. The foregoing combination of characters distinguishes charadricola from other members of the Hyla bistincta group, of which chryses most closely resembles chara- dricola. The former differs by having a point- ed snout in dorsal profile, larger tympanum in relation to the eye (mean ratio, 0.574, as compared with 0.340 in charadricola) , and a golden yellow dorsum in life. Superficialh' Hyhi charadricola r(>scmbles miotympanum and arborescandens, both of which ha\'e round snouts and shorter fingers. ■' Dr. Robert G. Webb obtained one specimen of this species on July 22, 1968, at 11 kilometers south of Chicahuaxtla, Oaxaca, at an elevation of 1450 meters. Dr. Kraig Adier olitained five individuals from three localities in the moimtains west of Chil- pancingo, Guerrero, in December, 1969. Description: Males of this species attain a maximum snout-vent length of 44.4 mm., and females reach 50.9 mm. In a series of 10 males from Rio Totolapa, 14.4 kilometers west of Huachinango, Puebla, Mexico, the snout- \ent length is 35.3 to 44.4 (mean, 40.4) mm.; the ratio of tibia length to snout-\cnt length is 0.500 to 0.5.35 (mean, 0.517); the ratio of foot length to snout-vent length is 0.459 to 0..506 ( mean, 0.493 ) ; the ratio of head length to snout-vent length is 0.292 to 0.319 (mean, 0.308); the ratio of head width to snout- vent length is 0.311 to 0.334 (mean, 0.320), and the ratio of the diameter of the tympanum to that of the eye is 0.295 to 0.372 (mean, 0.340). Three females from the same locality have snout-\ent lengths of 43.4 to 50.9 ( mean, 48.1) mm. The tympanum is slightly larger in females than in males; the ratio of the diam- eter of the tympanum to that of the eve in females is 0.375 to 0.391 (mean, 0.384).' The head is as wide as the body; the top of the head is flat. The eyes are large and prominent. In dorsal profile, the snout is truncate; in lateral profile, it is bluntly round- ed. The snout is short; the nostrils are slightly protuberant and situated about three-fourths the distance from the eyes to the tip of the snout. The internarial region is slightly de- pressed. The canthus is rounded; the loreal region is barely concaxe, and the lips are thick and flaring. A moderately heavy der- mal fold e.xtends posteriorly from the eye, above the tympanum, and downward to a point above the insertion of the arm. The fold obscures the upper edge of the tympanum, uhich is barely distinct and separated from the eye by a distance ec|ual to half again the diameter of the tympanum. The arms are moderately long and slen- der; a distinct axillary membrane is present. A row of low tubercles is present on the ven- tral edge of the forearm, and a faint trans- \-erse dermal fold is present on the wrist. The fingers are long and slender and bear rela- tively small discs; the width of the disc on the third finger is less than the diameter of the tympanum. The subarticular tubercles are small and round; the supernumerary tuber- cles are small, subconical, and irregularly placed on the proximal segments of the sec- ond, third, and fourth fingers. The palmar 1970 DUELLMAN: HYLID FROGS 467 tubercle is flat and is usually bifid. The pre- pollex is greatly enlarged, flattened, and ovoid; nuptial excrescences are absent in breeding males. A vestige of webbing is pres- ent between the second and third and the third and fourth fingers (fig. 234A). The hind limbs are moderately long and slender; the heels of the adpressed limbs overlap by about one-third the length of the shank. The tibio- tarsal articulation extends to the anterior corner of the eye. A moderately heavy trans- verse dermal fold is present on the heel. The tarsal fold is weak and usually present only on the distal half of the tarsus. The inner metatarsal tubercle is moderately large, ovoid, flattened, and raised medially. The outer metatarsal tubercle is small and conical. The toes are long and slender and bear discs that are about equal to the size of those on the fingers. The subarticular tubercles are small and round, and the supernumerary tubercles are minute and subconical. The toes are three-fourths webbed (fig. 234E). The web- bing extends from the distal end of the pen- ultimate phalanx of the first toe to the base of the penultimate phalanx of the second, from the base of the disc of the second to the distal end of the antepenultimate phalanx of the third, from the middle of the penulti- mate phalanx of the third to the distal end of the antepenultimate phalanx of the fourth and on to the base of the disc of the fifth toe. The anal opening is directed posteroven- trally at the midle\'el of the thighs. A short, thin anal sheath is present. The skin on the belly and proximal posteroventral surfaces of the thighs is weakly granular; no thoracic fold is present. The skin on the other surfaces is smooth. The tongue is nearly round, shal- lowly notched behind, and free posteriorly for about one-fourth of its length. The dentig- erous processes of the pre\omer are narrow transverse ridges bet\veen the moderately large, round choanae. Males have two to five teeth on each process and a total of five to 10 (mean, 7.6) prevomerine teeth. Females have four or fi\e teeth on each process and a total of eight to 10 (mean, 9.0) prevomerine teeth. Vocal slits and a vocal sac are absent. The general coloration of Hyla charad- ricola is dark green with darker green reticu- lations on the back (pi. 6.3, fig. 1). The dorsal surfaces of the head, body and limbs are dark green; darker green reticulations usually are evident on the back. The flanks are dirty white with dark olive-gray mottling. A dark oli\e-gray stripe extends from the nostril to the eye and then to the insertion of the arm. The upper lips are pale green. The inguinal region, anterior and posterior surfaces of the thighs, and inner surfaces of the feet are dark yellowish orange. The ventral surfaces of the shanks, feet, and webbing are dusty yellow. The belly is white, and the iris is silvery gold. In some individuals, the dark reticulations on the dorsum are faint. Some adults, when collected, were pale green with faint or no dorsal reticulations; later these individuals became darker, and usually the reticulations were evident. In all specimens the anal stripe is absent and the flanks are heavily mottled. Juveniles have a dorsal color varying from rich brown with darker reticulations to pale green or gray with dark green reticulations. In preservative the dorsum is purplish brown with fine darker reticulations on the back. The flanks are pale tan with dark brown spots, and the posterior surfaces of the thighs are tan. The chin is creamy white with brown spots, and the belly is dusty white. The ven- tral surfaces of the thighs and shanks are pale yellow; the webbing is grayish brown. The ventral surfaces of the first two fingers are dusty white, and the ventral surfaces of the third and fourth fingers and of the feet are brown. There is no anal stripe, but small white flecks are present above and below the anal opening. Tadpoles: The tadpoles of Hyla charad- ricola are unknown. Presumably, they devel- op in mountain streams. Mating Call: The absence of vocal slits and a vocal sac precludes the presence of a call in this species. Natural History: Most specimens were obtained at the Rio Totolapa, a shallow rocky stream in a pine forest. There, Hyla charad- ricola was found beneath rocks at the edge of fast moving sections of the stream and be- neath rocks in shallow ripples in the stream. Most of the frogs were in water. At night, they were found sitting on rocks in the stream. At Lago de Tejocotal, Hyla choradricola was found beneath rocks at the shore of the lake 468 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 and by a stream in the pine forest. Individ- uals were found on low vegetation overhang- ing a small stream in pine-oak forest, 4 kilome- ters southwest of Tianguistengo, Hidalgo, Mexico. Hyla miotyuipanum is abundant at the Rio Totolapa. Indi\ iduals of this species were found beneath rocks at the edge of the stream by day and in bushes along the stream at night, but not in the ripples inhabited by Ili/la charadricola. Five recently metamorphosed young were found at the Rio Totolapa, on June 8, 1960; these specimens have snout-vent lengths of 22.4 to 24.0 (mean, 23.2) mm. Remarks: The presence of an axillary membrane and relatively thin skin on the dorsum, plus the absence of nuptial excres- cences in breeding males are characteristics shared by Hyla chryses. The latter differs in coloration and in having a pointed, instead of a truncate snout. Etymology: The specific name charad- ricola is derived from the Greek charadra, meaning mountain stream, and the Latin suffix, -cola, meaning an inhabitant; the name refers to the habitat of the frog. Distribution: Hyla charadricola inhabits streams in pine and pine-oak forests at eleva- tions of 2000 to 2.300 meters in northern Pu- ebla and in eastern Hidalgo, Mexico (fig. 240). See Appendix 1 for the locality records of the 59 specimens examined. Hyla chryses Adler lUjla chryses Adler, 1965, p. 1 [holotype, U.M.M.Z. No. 12.5.374 from between Puerto Chico and Asolea- dero ( aliout 4.5 kilometers airline west-northwest of Chitpancingo), Guerrero, Mexico, ele\'ation 2540-2600 meters; Kraig .\dler collector]. Diagnosis: This small (males attain a snout-vent length of 37.3 mm.) member of the Hyla bistincta group has relatively thin skin on the dorsum, a pointed snout in dorsal profile, and an axillary membrane. The dor- sum is golden yellow to dark greenish brown. Vocal slits, nuptial excrescences, and a tho- racic fold are lacking, and the anal opening is at the level of the middle of the thigh. The only other member of the Jlyla bistincta group having an axillary membrane, thin skin, and 100° 98° 2CP - - ' / ; rf'y / .^ - 20° 18° i 1 /^-v ' \ yd \ ^-^-^^ > 0 i 18° 1. 1 ^....^^ • /y. charadricola ^~^~\_^ 0 H. chryses 0 50 100""^-^ ^ '■' KILOMETERS 1 ^^^ 100° 98° Fig. 240. Distribution of Hijla cliaradricula and Hyla chryses. lacking nuptial excrescences is charadricola. That species has a green dorsum, truncate snout in dorsal profile, and a smaller tym- panum (mean t>inpanum/eye ratio 0.340, as compared with 0.574 in chryses). Some mem- bers of the Hyla pinorum and mixomaculata groups superficially resemble chryses. Mem- bers of both groups are smaller and either have blunt snouts or small or covered tym- pani. Description: This is the smallest species in the Hyla bistincta group. Males attain a maximum known snout-vent length of 37.6 mm., and the one female has a snout-\'ent length of 42.2 mm. In a series of three males from the type locality in the Sierra Madre del Sur in Guerrero, Mexico, the snout-vent length is 36.3 to 37.6 (mean, 37.1) mm.; the ratio of tibia length to snout-\ent length is 0.494 to 0.49S (mean, 0.496); the ratio of foot length to snout-vent length is 0.473 to 0.482 ( mean, 0.478 ) ; the ratio of head length to snout- vent length is 0.313 to 0.316 (mean, 0.315); the ratio of head width to snout-\-ent length is 0.313 to 0.327 (mean, 0.320), and 1970 DUELLMAN: HYLID FROGS 469 the ratio of the diameter of the t\nipanuni to that of the eye is 0.595 to 0.634 (mean, 0.610). The one female does not differ noticeably in proportion from the males, e.xcept that it has a slightly smaller tympanum; the t\mpanum/ eye ratio is 0.574. The head is as wide as the body; the top of the head is barely convex. The eyes are moderateh' large and prominent. In dorsal profile, the snout is broadly pointed \\ith a faint imitation of a rostral keel; in lateral profile the snout is bluntly rounded. The snout is moderately long; the nostrils are pro- tuberant and situated about three-fourths of the distance from the eyes to the tip of the snout. The internarial region is barely de- pressed. The canthus is round; the loreal re- gion is noticeably concave and the lips are thick and bareK' flared. A moderately heavy dermal fold extends from the eye, abo\'e the txmpanum, and downward to a point above the insertion of the arm. The fold obscures the upper edge of the tympanum, which otherwise is distinct and is separated from the eye by a distance equal to the diameter of the tympanum. The arms are moderately long and slender; an indistinct axillary membrane is present. A few low tubercles are present along the \'en- trolateral edge of the forearm, and a distinct transverse dermal fold is present on the wrist. The fingers are moderately long and slender and bear large discs; the width of the disc on the third finger is equal to the diameter of the tympanum. The subarticular tubercles are moderately large and round; none is bifid. The supernumerary tubercles are small, sub- conical, and irregularly arranged on the proxi- mal segments of the digits. The palmar tu- bercle is flat and bifid. The prepollex is mod- erately enlarged and ovoid; nuptial excres- cences apparently are lacking in breeding males. The webbing between the fingers is vestigial (fig. 234B). The hind limbs are moderately long and slender; the heels of the adpressed limbs overlap by about one-fourth of the length of the shank. The tibiotarsal articulation extends to the middle of the eyes. A weak transverse dermal fold is present on the heel, and a weak tarsal fold extends the full length of the tarsus. The inner meta- tarsal tubercle is elliptical, rounded, and bare- ly \isible from above. The outer m • /7. robertsorum o H pachyderma ■ H siopelo D H. crassa A H. bogertae 1 ^ N, V ( 6 \ \ ■ \ 18° y 1 \ A 18° 0 50 I00\ KILOMETERS ^^-^^^—-"^ 1 1 98° 96° Fig. 241. Distribution of Hijla robertsuium, pachyderma, siopela, crassa, and bogertae. bing between the fingers and moderately large nuptial spines on the prepollex in breed- ing males. The latter character is unique in the group and is present elsewhere in Middle American hylids only in Hyla echinata and in some species of Ftychohyla. Hyla echinata differs from pachyderma by having nearly fully webbed hands and dermal folds on the edges of the forearms and feet. Breeding males of the species of Ftychohyla have large \cntrolateral glands, vocal slits, and short, webbed fingers. Description: One male from Pan de Olla, Veracruz, Mexico, has a snout-vent length of .39.9 mm.; the ratio of tibia length to snout- \ cnt length is 0.526; the ratio of foot length to snout-vent length is 0.514; the ratio of head length to snout-vent length is 0.308, and the ratio of head width to snout-vent length is 0.321. The tympanum is not visible. Two females from the same locality have snout- 474 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 vent lengths of 52.7 to 55.7 (mean, 54.2) mm. The tympanum is visible in the females; the ratio of the diameter of the tympanum to that of the eye is 0..340 to 0.341 ( mean, 0.341 ) . The head is slightly narrower than the body. The top of the head is flat; the eyes are large and prominent. In dorsal profile the snout is rounded; in lateral profile it is round- ed above and truncate terminally. The snout is short; the nostrils are barely protuberant and situated about two-thirds the distance from the eyes to the tip of the snout. The canthus is rounded; the loreal region is barely concave, and the lips are thick, rounded, and not flared. A heavy dermal fold extends from the posterior corner of the eye to a point above the insertion of the arms. This fold completely obscures the upper part of the tympanum in all specimens; in the males the lower part of the tympanum is covered by thin skin so as not to be visible. The arms are moderately short and robust; an a.xillary membrane is missing. A few- small tubercles are present on the ventral sur- faces of the forearms and a distinct dermal fold is present on the wrist. The fingers are long and slender and bear moderately large discs. The subarticular tubercles are large and round; none is bifid. The supernumerary tubercles are large and conical; they are pres- ent in a single row on the proximal segment of each digit. The palmar tubercle is low, flat and bifid. The prepollex is greatly enlarged; in a breeding male it bears a large clump of moderately large spines. The spines are present on the inner surfaces of the thumb and second finger. Only a vestige of webbing is present between the fingers (fig. 2.33D). The hind limbs are robust; the heels of the adpressed limbs overlap by about one-fourth of the length of the shanks. The tibiotarsal articulation extends to the anterior corner of the eye. A transverse dermal fold is present on the heel, and a thick low tarsal fold is present on the distal two-thirds of the tarsus. The inner metatarsal tubercle is moderately large, elliptical, and raised medially. The outer metatarsal tubercle is small and conical. The toes are long and slender and bear discs that are only slightly smaller than those on the fingers. The subarticular tubercles are moderately large and round, and the super- numerary tubercles are distinct and subconi- cal. The toes arc about three-fourths webbed (fig. 235B). The webbing extends from the distal end of the penultimate phalanx of the first toe to the base of the penultimate phalanx of the second, from the base of the disc of the second to the base of the antepenultimate phalanx of the third, from the base of the penultimate phalanx of the third to the distal phalanx of the third, from the base of the fourth and on to the middle of the penulti- mate phalanx of the fifth. The anal opening is directed posteroven- trally at the midlevel of the thighs. A short anal sheath is present, and a distinct trans- verse dermal fold is present above the anal sheath. The skin on the dorsum and \entral surfaces of the limbs, except the thighs, is smooth; the skin on the chin, belly and \entral surfaces of the thighs is granular. A distinct thoracic fold is present. The tongue is ovoid, shallowly notched anteriorly and posteriorly and barely free behind. The dentigerous pro- cesses of the prevomers are small, ovoid, and situated in a transverse plane between the moderately large, nearly round choanae. One male has three teeth on each process for a total of six pre\omerine teeth, whereas two females have four teeth on each process and a total of eight prevomerine teeth. Vocal slits and a vocal sac are absent. No knowledge of the color of this species in life exists. In preservative the general col- oration is dull grayish brown with indistinct, scattered, darker flecks on the dorsal surfaces ( pi. 4, fig. 1 ) . The flanks are grayish brown with cream reticulations, and the posterior surfaces of the thighs are tan. The chin is cream, mottled with brown. The belly is creamy yellow and is mottled with brown anteriorly in the females. A creamy white anal stripe is present, and in the females the stripe extends laterally in the form of a row of creamy white dashes and spots onto the pos- terodorsal surfaces of the thighs. T.\DPOLES: No tadpoles of this species are known. Presumabh' they de\elop in mountain streams. Mating C.\ll: The absence of vocal slits and a vocal sac precludes the presence of a call in this species. Natur.\l History: Taylor and Smith 1970 DUELLMAN; HYLID FROGS 475 (1945, p. 588) stated that the known speci- mens of Hyla pachijderma were found on bushes and weeds beside a small, bounding stream near Pan de Olla. Veracruz, Mexico. I have searched unsuccessfully for this species in the area around Pan de Olla and Tezuitlan. Nothing more is knou'n about the natural history of this species. Remarks: On the basis of the four speci- mens a\ailable for study, HyJa pachyderma seems to be closely related to Hyla crassa and Hyla rohertsonim. Perhaps, these three spe- cies, as known now, are merely representa- tives of one ta.xon, but, if so, the differences between the known populations are distinc- tive. Hyla pachyderma is unique in the Hyla histincta group by having moderately en- larged nuptial spines. Etymology: The specific name pachy- derma is deri\'ed from the Greek, pachys, meaning thick, and the Greek derma, mean- ing skin; the name refers to the thick, glandu- lar skin on the dorsum. Distribution: Hyla pachyderma is known only from a stream at an elevation of about 1600 meters on the Atlantic slopes of the Si- erra Madre Oriental in central Veracruz, Mex- ico (fig. 241). See Appendix 1 for the locality records of the four specimens examined. Hyla siopela Duellman Hijla siopela, Duellman, 1968a, p. 570 [holotype, K.U. No. 100981 from the west slope of Cofre de Perote, Veracruz, Mexico, elevation 2500-2550 meters; William E. Duellman collector]. Diagnosis: This medium-sized (males at- tain snout-vent lengths of 46.2 mm.) member of the Hyla histincta group lacks an axillary membrane, and vocal slits. It has webbing between the two outer fingers, and the toes are about two-thirds webbed. Small nuptial spines and a weak thoracic fold are present. The snout is truncate and has a weak, vertical rostral keel, a character not present in other members of the group. In northern Middle America the only other hylids with a rostral keel are some species of the genera Ptycho- hyla and Plectrohyla. In those species vocal shts are present; breeding males of Ptycho- hyla have spinous nuptial excrescences and ventrolateral glands, and males of Plectro- hyla have projecting prepollical spines. Description: Hyla siopela is a medium- sized species, in which the males attain a maximum snout-vent length of 46.2 mm., and females reach 52.5 mm. In a series of seven males from the Cofre de Perote, Veracruz, Mexico, the snout-vent length is 42.1 to 46.2 (mean, 44.4) mm.; the ratio of the tibia length to snout-vent length is 0.472 to 0.500 (mean, 0.487); the ratio of foot length to snout-vent length is 0.456 to 0.495 (mean, 0.474 ) ; the ratio of head length to snout- vent length is 0.286 to 0..304 (mean, 0.296); the ratio of head width to snout-vent length is 0.291 to 0.317 (mean, 0.309), and the" ratio of the diameter of the tympanum to that of the eye is 0.363 to 0.468 (mean, 0.4.38). Five females from the same locality have snout- vent lengths of 45.1 to .52.5 (mean, 49.6) mm. Females have noticeably larger tympani than do the males; the ratio of the diameter of the tvmpanum to that of the eyes in females is d..500 to 0.545 (mean, 0.516). The head is about as wide as the body; the top of the head is barely convex, and the eyes are large and prominent. In dorsal pro- file the snout is truncate with a faint, vertical rostral keel; in lateral profile the snout is truncate. The snout is short; the nostrils are protuberant and situated at about four-fifths of the distance from eyes to the tip of the snout. The canthus is angular; the loreal region is concave, and the lips are thick and not flared. A heavy dermal fold extends pos- teriorly from the eye, above the tympanum, and thence downward to the point of inser- tion of the arm. The fold obscures the upper one-third of the tympanum, which otherwise is distinct and separated from the eye by a distance slightly greater than the diameter of the tympanum. The arms are moderately long and robust; an axillary membrane is lacking. A row of small tubercles is present on the ventro- lateral edge of the forearm, and a distinct dermal fold is present on the wrist. The fingers are long and slender and bear mod- erately large discs; that on the third finger is as large as the tympanum. The subarticular tubercles are moderately small and round; none is bifid. The supernumerary tubercles are small and in some specimens barely dis- tinguishable; they are arranged in a single 476 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 row on the proximal segment of each digit. The palmar tubercle is low, flat, and barely visible. The prepollex is greatly enlarged and flattened ventrally. In breeding males it bears nuptial excrescence composed of minute horny spinules. The nuptial excrescence is also present on the inner surface of the thumb. Little webbing is present between the fingers ( fig. 233E ) . The webbing is vestigial between the first and second fingers; the web extends from the base of the penultimate phalanx of the second to the base of the antepenultimate phalanx of the third and on to the base of the penultimate phalanx of the fourth finger. The hind limbs are relatively short and ro- bust; the heels of the adpressed Hmbs over- lap by about one-third of the length of the shank. The tibiotarsal articulation extends to the posterior edge of the eye. A transverse dermal fold is present on the heel, and a thin tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is large, elongate, flat, and visible from above. The outer metatarsal tubercle is absent. The toes are moderately long and slender and bear discs that are slightly smaller than those on the fingers. The subarticular tubercles are moderately small and round; the supernu- merary tubercles are small and arranged in a single row on the proximal segments of each digit. The toes are about two-thirds webbed (fig. 235C). The webbing extends from the middle of the penultimate phalanx of the first toe to the base of the penultimate of the second, from the middle of the penultimate phalanx of the second to the middle of the antepenultimate phalanx of the third, from the middle of the penultimate phalanx of the third to the middle of the antepenultimate phalanx of the fourth and on to the middle of the penultimate phalanx of the fifth toe. The anal opening is directed posteriorly at the midlevel of the thighs. A short anal sheath is present. The skin on the chin, belly, and posteroventral surfaces of the thighs is granular; elsewhere it is smooth. A weak thoracic fold is present. The tongue is broad- ly cordiform, notched posteriorly, and barely free behind. The dentigerous processes of the prevomers are posteromedially inclined elevations between small ovoid choanae. Males have three to five teeth on each process and a total of six to nine (mean, 7.9) pre\o- merinc teeth. Females have four or five teeth on each process and a total of eight or nine (mean, 8.4) prevomerine teeth. A \ocal sac and vocal slits are absent. The general coloration of Hyla siopela is green or tan with darker reticulations (pi. 63, f^g. 5). A typical adult male has a pale green dorsum with black spots and reticulations. The flanks are mottled dark brown and creamy white. The outer edges of the feet are silvery white with brown spots; the an- terior and posterior surfaces of the thighs are dull brown, and the webbing and the first three toes are dull yellowish tan. The belly is creamy gray, and the throat is silvery white mottled with gray. The iris is dull bronze with black reticulations. Some individuals have an olive-green or dark green dorsum with darker green or black flecks or reticulations: other individuals are pinkisli tan or brown with dark brown flecks or reticulations. All specimens have some white markings above the anus and on the postcrodorsal surfaces of the thighs; in some individuals the white flecks are expand- ed and interconnected to form an irregular white line. In preser\ative the dorsum is dull grayish brown with small, irregularly shaped black spots on the head, back, and limbs. The flanks are gray mottled with creamy tan; the an- terior and posterior surfaces of the thighs are tan. The belly is dull creamy tan, and the throat is marked with gra>- blotches. The anal region and posterior surfaces of the thighs are marked with small white spots. In most preserved specimens the dorsum is heav- ily marked with dark spots or flecks, but in some specimens relatively few dark flecks are present. Juveniles have notably different coloration in life. The dorsum is uniformly pale green (pi. 63, fig. 4). The anterior and posterior surfaces of the thighs, fingers, first three toes and the webbing are deep yellow. The anal stripe is creamy white and the flanks are pale gray with black flecks. The upper lip, supra- tympanic fold, and canthal stripe are a bronze color. The belly is pale \ellow with a silver cast on the throat. Juveniles haxing snout- vent lengths from 24.5 to 36.6 mm. are so col- 1970 DUELLMAN: HYLID FROGS 477 ored in life and are uniform dark bluish gray dorsally in preservati\e. Tadpoles: Four specimens in advanced stages of de\elopment are available. Three specimens in developmental stage 41 have body lengths of 23 to 26 (mean, 24) mm. One specimen in de\elopmental stage 37 has a bod\' length of 26.5 mm. and a total length of 66.0 mm. The body is moderately depressed; the width is noticeably more than the depth. In dorsal profile the snout is bluntly rounded; in lateral profile the snout is inclined antero- ventrally from the nostril to a bluntly rounded tip. The nostrils are small, directed anteriorly, and situated slightly closer to the eyes than to the tip of the snout. The eyes are small, slighth^ ele\ated and directed dorsolaterally. The sinistral spiracle is short; the spiracular opening is directed posterodorsally at a point on the midline slightly less than half the dis- tance from the snout to the posterior edge of the body. The anal tube is long and dextral. The tail is long, low, and bluntly rounded terminally. The caudal musculature is heavy and does not extend to the end of the caudal fin. At midlength of the tail, the depth of the musculature is equal to the depth of the ven- tral fin and is deeper than the depth of the dorsal fin. The dorsal fin does not extend onto the body ( fig. 2.37D ) . In preser\'ative the body is dark grayish brown with bluish gray flecks ventrally. The tail is creamy tan with dark brown flecks. Only the periphery of the caudal fins is transparent. The mouth is ventral and relatively small; its width is equal to about one-half of the greatest width of the body. The lateral folds in the lips are barely discernible. The mouth is completely bordered by two rows of small papillae; medial to these is an irregular row of larger papillae. The beaks are moderately slender and bear long, pointed serrations. The upper beak is in the form of a broad arch with short, blunt lateral processes. The lower beak also forms a broad arch. There are two upper and three lower rows of teeth. The upper rows are about equal in length, and the second upper row is narrowly interrupted medially. The lower rows are shorter than the upper ones and progressively shorter from the first to the third row (fig. 238D). Mating Call: The absence of vocal slits and a vocal sac precludes the presence of a mating call in this species. Natural History: This species inhabits relati\cly dry pine forests. All individuals were found along a stream, where both adults and juveniles were found in crevices and on rocks behind small cascading waterfalls by day or sitting on rocks or branches in the spray of cascades by night. Tadpoles were found in pools in the stream, where they hid under moss-covered banks. Remarks: Messrs. Macreay J. Landy and John D. Lynch obtained the first specimens of this species on July 30-31, 1964. Their specimens were tentatively identified as Htjla paclnjderma. I visited the stream on Cofre de Perote in February, 1966; at that time, no frogs were found. Mr. Howard L. Freeman visited the locality on June 18, 1966, and ob- tained several frogs and four tadpoles. I re- turned to the stream on July .30, 1966, and obtained several frogs, but no tadpoles. The examination of the fresh material and com- parison of it with the specimens obtained by Land}' and Lynch and with the type series of Hyla paclnjderma led to the conclusion that the frogs inhabiting Cofre de Perote repre- sented a distinct and previously unnamed species (Duellman, 1968a, p. 570). Etymology: The specific name is derived from the Greek siopelos, meaning silent, and is used in allusion to the absence of a voice in the species. Distribution: Hyla siopela is known only from a small stream on the west slope of Cofre de Perote, in the Sierra Madre Oriental in cen- tral Veracruz, Mexico, at an elevation of 2500 to 2550 meters (fig. 241). See Appendix 1 for the locality records of the 54 specimens examined. Hyla crassa (Brocchi) Cauphias crassus Brocchi, 1877b, p. 130 [liolotype, M.N. H.N. No. 6331 from "Mexico"; Adolpe Boucard collector]. Cauphias crassum Brocchi, 1882a, p. 64. Kellogg, 1932, p. 118. Hyla crassa: Boulenger, 1882a, p. 396. Guntlier, 1901 (1885-1902), p. 281. Smith and Tavlor, 1948, p. 86. Duellman, 1964b, p. 486. Hypsiboas crassus: Cope, 1887, p. 14. Hyla rohustofemora Taylor, 1940c, p. 239 fholo- type, U.I.M.N.H. No. 25050 (formerly E.H.T.-H.M.S. 478 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 No. 16.314) from Cerro San Felipe, 1.5 kilometers northeast of Oa.xaca de Juarez, Oa,\aca, Mexico; Ed- ward H. Taylor collector]. Smith and Tavlor, 1948. p. 86. Plcctrohyla criissa: Hartweg, 1941, p. 1. Diagnosis: This large (males attain a length of 53.7 mm.) member of the Hyh histincta group has a strong tarsal fold, small nuptial spines, a round snout, and vestigial webbing on the hand. Vocal slits, axillary membranes, and a thoracic fold are absent. The feet are webbed to the base of the discs. By this character alone crassa can be distin- guished from all other members of the Htjia histincta group. Description: One adult male from Cerro San Felipe, Oaxaca, Mexico, has a snout-vent length of 5.3.7 mm.; the ratio of tibia length to snout- vent length is 0.501; the ratio of foot length to snout-vent length is 0.473; the ratio of head length to snout-vent length is 0.29S; the ratio of head \\'idth to snout-vent length is 0..32S, and the ratio of the diameter of the tympanum to that of the eye is 0.278. A fe- male from an unknown locality has a snout- vent length of 48.2 mm. In the female, the tympanum is completely concealed; other- wise, it resembles the male in proportions. The head is slightly narrower than the body and barely convex on top. In dorsal profile, the snout is broadly rounded and in lateral profile, bluntly rounded. The snout is short; the nostrils are barely protuberant and are situated about two-thirds the distance from the eyes to the tip of the snout. The canthus is rounded; the lorcal region is barely concave, and the lips are thick and not flared. A heavy dermal fold extends posterovcntrally from the posterior corner of the eye to a point above the insertion of the arm; this fold obscures the entire tympanum in one female and the upper half of the t\'mpanum in the one male. Otherwise, the tympanum is barely discernible and is separated from the eye by a distance half again the length of the diameter of the tympanum. The arms arc short and thick; no axillary membrane is present. Tubercles are absent along the ventrolateral edge of the forearm, but a distinct dermal fold is present on the wrist. The fingers arc moderately long and slender and bear moderately large discs; the disc on the third finger is somewhat larger than the diameter of the tympanum. The subarticular tubercles are moderately small and round; none is bifid. The supernumerary tubercles are large and subconical. The pal- mar tubercle is large, flat, and partially bifid. The prepollex is greatly enlarged and round- ed; in the one male it bears a nuptial excres- cence composed of minute spinules. he nup- tial excrescence also is present on the penulti- mate phalanges of the first and second fingers. The fingers essentially lack webbing (fig. 233F ) . There is no web between the first and second fingers and only rudimentary web be- tween the others. The hind limbs arc short and robust; the heels of the adpressed limbs overlap by about one-fourth the length of the shank. The tibiotarsal articulation extends to the posterior corner of the eye. A trans- verse dermal fold is present on the heel, and a thick tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is rather small, o\oid, and rounded. The outer metatarsal tubercle is small, flat, and indis- tinct. The toes are moderately long and slen- der and bear discs that are somewhat smaller than those on the fingers. The subarticular tubercles are moderately large and round; the supernumerary tubercles are large, low, and arranged in a single row on the proximal segments of each digit. The toes are fully webbed (fig. 2.35D). A dermal fringe is pres- ent on the inner edge of the first toe and the outer edge of the fifth toe. The anal opening is directed postero\en- trally at the midlevel of the thighs; an elon- gate anal sheath is present, and small tuber- cles are present below the anal opening. The skin on the dorsal surface of the body is smooth, but it is somewhat granular on the dorsal surfaces of the limbs. The skin on the chin and belly is moderately granular, and tliat on the ventral surfaces of the thighs is lu'a\ily granular. There is no thoracic fold. The tongue is nearly round, shallowly notched posteriorly and free behind for about one- fourth of its length. The dentigerous pro- cesses of the pre\omcrs are elliptical in shape and situated on a transverse plane between the small ovoid choanae. One male has five teeth on each process, and one female has seven and eight teeth on each process. Vocal sHts and the \ocal sac are absent. 1970 DUELLMAN: HYLID FROGS 479 The general coloration of this frog in pre- servative is dull brown (pi. 4, fig. 2). The venter is dull creamy tan with brown suflFu- sion on the throat and ventral surfaces of the hind limbs. A few creamy yellow spots are present on the flanks. Taylor (1940c, p. 392), in his description of Hijla robustofemora, stated: "Above, a uni- form dull o]i\e-green, somewhat lighter on the sides of the head and body; chin, gray with >ellow flecks; abdomen, creamy yellow with some pigmentation posteriorly, especial- ly under posterior part of femur; palms and soles, dark la\cnder-gray; posterior side of femur gra\- with wash of yellow; a cream spot under forearm; a few cream spots on side, on anterior face of femur, and at knee and heel; a dim spot of cream on anal flap." The only other known specimen, a female (M.N.H.N.No. 6331, the holotype of HijJa crassa) has more cream mottling on the flanks and posterior surfaces of the thighs and more distinct mottling on the throat than does the male; these differences were used as the basis of description of Hijla rohustofemora. T.-^DPOLES: No tadpoles of this species have been found. Mating C.\ll: The absence of vocal slits and a vocal sac preclude the presence of a call in this species. Natural History: The only knowledge of the natural history of this species is in- corporated in a brief statement by Taylor (1940c, p. 389): "In the summer of 19.38, I obtained a specimen of an undescribed Hijla at night, hopping along the edge of a small spring-fed rivulet at an elevation of about 2.300 meters on the Cerro San Felipe. The frog, frightened by my approach, jumped into the rivulet, swam to the opposite side and clamored up the bank, without attempting to hide under the water." Remarks: The systematic status of Cau- phias crassus Brocchi remained in doubt from the time of its original description until Duell- man (1964b, p. 488) re-examined the type specimen and compared it with the holotype of Hijla rohustofemora (U.I.M.N.H. No. 250.50). Brocchi (1877b, p. 1.30) and Kellogg (19.32, p. 118) erroneously stated that the terminal phalanges in the holotype of Cau- phias crassus were T-shaped and that the terminal phalanx was not preceded by an intercalary cartilage. Duellman ( 1964b, p. 489) stated: "The type of Catiphias crassus possesses intercalary cartilages between the penultimate and terminal phalanges; the latter are not T-shaped, but as in the type of Hijla rohustofemora resemble those typical of Hyla." Etymology: The specific name is derived from the Latin crassus, meaning thick or fat and alludes to the robust appearance. Distribution: Hyla crassa is known only from a small stream at an elevation of 2300 meters in the mountains of central Oaxaca, Mexico (fig. 241). 1^ See Appendix 1 for the locality records of the two specimens examined. Hyla bogertae Straughan and Wright Hyla bogertae Straughan and Wright, 1969, p. 1 [holotype, L.A.C.M. No. 44400 from a tributary of the Rio Atoyac, below Vix-ero El Tapanal, 1.6 kilometers south of La Cofradia, Distrito Sola de Vega, Oa.\aca, Mexico, elevation 2652 meters; Ian R. Straughan and John W. Wright collectors]. Diagnosis: Hyla hogertae is a moderately large (snout-vent length in one adult male, 45.1 mm.) member of the Hyla histincta group with a rounded snout, discontinuous tarsal fold, and non-spinous nuptial pads in breeding males. Vocal slits, axillary mem- branes, thoracic fold, and webbing on the hand are absent. Hyla bogertae is similar to rohertsorum, which has vestigial webbing be- tween the fingers and a weak thoracic fold; furthermore, there are differences in colora- tion: rohertsorum has dark brown flanks with small cream spots and a gray belly, whereas hogertae has olive-green flanks with large yellow spots and a white belly. Hyla siopela differs from hogertae by having a more trun- cate snout with a weak rostral keel and less webbing (2/3 in siopela; 4/5 in hogertae). Hyla pachyderma differs from hogertae by having a strong thoracic fold and large nup- " In April, 1969, Mr. Chuck McClung collected li\c specimens of this species from rock crevices at a locality 14.4 kilometers northeast of Ciudad Oaxaca, Oaxaca. I ha\'e no further details concerning the localit>' but suspect that it is in the highland mass associated with Cerro San Felipe. Three specimens are preserved in the United States National Museum, and one is in the Museum of Natural History at the University of Kansas. 480 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 tial spines in breeding males. Hijla crassa differs by having the feet fully webbed and a strong tarsal fold. Hyla bistincta and penthe- ter differ from bogertae by having a strong tarsal fold and an elongate anal sheath (short in bogertae ) . Description: The one adult male has a snout-vent length of 45.1 mm.; three adult females have snout-vent lengths of 43.3 to 50.1 (mean, 47.6) mm. In the one male the ratio of tibia length to snout-vent length is 0.505; the ratio of head length to snout-vent length is 0.262; the ratio of head width to snout-vent length is 0.365; and the ratio of the diameter of the tympanum to that of the eye is 0.404. The proportions of the three females are about the same as those of the male, ex- cept that the tympanum is proportionately smaller to the eye in the females; the ratio of the diameter of the tympanum to that of the eye is 0.333 to 0.400 (mean, 0.371). The head is slightly narrower than the body; the top of the head is barely conve.x. In dorsal profile the snout is bluntly rounded; in lateral profile, the snout is truncate and rounded above. The snout is short; the nos- trils are slightly protuberant and situated about three-fourths of the distance from the eyes to the tip of the snout; the internarial region is slightly depressed. The canthus is rounded; the loreal region is concave, and the lips are thick and rounded. A heavy dermal fold extends posteriorly from the eye, above the tympanum, and angles downward to a point above the insertion of the arm. The fold covers the upper edge of the tympanum, which otherwise is distinct and separated from the eye by a distance equal to half again the diameter of the tympanum. The arms are moderately long and robust; an axillary membrane is absent. A few small tubercles are present on the ventrolateral edge of the forearm, and a weak dermal fold is present or absent on the wrist. The fingers are long and slender and bear small discs; the width of the disc on the third finger is equal to the diamet(;r of the tympanum. The sub- articular tubercles are large and round; none is bifid. The supernumerary tubercles are moderately large and low; they are arranged in a single row on the proximal segment of each digit. The outer palmar tubercle is low. rounded, and bifid. The prepoUex is enlarged and rounded. A horny nuptial excrescence is present in the one breeding male. Webbing is absent between the fingers. The hind limbs are robust; the heels of the adpressed limbs barely o\erlap. A hea\y transverse dermal fold is present on the heel. The tarsal fold is interrupted and essentially consists of a series of low tubercles. The inner metatarsal tubercle is small and ovoid; the outer meta- tarsal tubercle is small and subconical. The toes are moderately long and slender and bear discs that are slightly smaller than those on the fingers. The subarticular tubercles are large and round. The supernumerary tuber- cles are small and present in a single row. The toes are about four-fifths webbed; the webbing extends to the base of the discs on all toes, except the fourth, where it extends to the base of the penultimate phalanx. The anal opening is directed posteroven- trally near the midlevel of the thighs; it is covered by a short anal sheath. The skin on the dorsum and ventral surfaces of the arms, slianks, and feet is smooth; that on the throat, belly, and ventral surfaces of the thighs is granular. A thoracic fold is lacking. The tongue is broadly cordiform, barely notched behind, and free posteriorly for about one- fourth of its length. The pre\omerine teeth are situated on posteromedially inclined ele- vations between the small, ovoid choanae. In one female there are nine prevomerine teeth, and in one male there are six teeth. Vocal slits and a \ocal sac are absent. The general coloration of Hyla bogertae is dark brown with indistinct tan spots dor- salh'. The brown gi\es way to gray lateralK' with gray and white spots on the flanks and lips. The dorsal surfaces of the limbs are mottled brown and gray. The posterior sur- faces of the thighs are dark brown. A white anal stripe is present. The throat is dark brown with cream spots. The rest of the ven- tral surfaces are uniform creamy white, ex- cept the hands and feet, which are dark brown. Straughan and Wright (1969, p. 3) de- scribed the coloration in life of the female Iiolot>pe: "Dorsal surface of body and limbs oii\e green (gun metal gray in alcohol) with extensive silver to pale bronze reticulation, 1970 DUELLMAN: HYLID FROGS 481 largely maintained in alcohol. Light bar above level of cloaca at beginning of \entral granularity. Ventral surface of body white with yellow wash along flanks. Limbs mainly darker with yellow in groin area, around heels, upper arm, and elbow. Throat dark- er olive with large yellow spots and mi- nute creamy pustules." Straughan and Wright (1969, p. 6) noted the coloration of the male allotype: "Color darker than in holotype with essentially the same pattern, but slightly less de\elopment; throat color darker and more extensive than in holotype. In all other characters allotype agrees with holotype." Tadpoles: Three tadpoles in develop- mental stage 25 ha\e total lengths of 30 to .32 mm., and four in developmental stage 30 have total lengths of 52 to 57 mm. A typical tadpole in developmental stage 30 has a body length of 19.1 mm. and a total length of 55.7 mm. The body is depressed and slightly wider than deep. In dorsal and lateral profiles the snout is bluntly rounded. The nostrils are small, directed anterolaterally, and situated about midway between the eyes and the tip of the snout. The eyes are moderately small and directed dorsolaterally. The spiracle is sinistral; its opening is directed posteriorly at a point on the midline at about midlength of the body. The cloacal tube is moderately long and dextral. The tail is long, low, and terminally rounded. The caudal musculature is robust and moderately deep; at midlength of the tail the depth of the caudal musculature is only shghtly less than the depth of the dorsal fin, which does not extend onto the body. The body is dark grayish brown dorsally and slightly paler laterally with scattered black flecks and golden lichenous markings. The caudal musculature is tan, and the fins are translucent tan. Dark brown spots are scattered on the caudal musculature and fins. The mouth is ventral and moderately large; its width is equal to about two-thirds of the width of the body. Lateral folds are present in the lips, which are bordered by two rows of small papillae. Median to the small labial papillae there is one row of larger papillae on the upper lip and two rows on the lower lip. A few large papillae are present in the lateral folds. The beaks are moderately robust and bear small serrations. The upper beak is in the form of a broad arch with long, rather slender, terminally rounded lateral processes. The lower beak is broadly \'-shaped. There are two upper and three lower rows of teeth. The upper rows are about equal in length and slightly longer than the lower rows. The second upper row is narrowly interrupted medially. Tadpoles in developmental stage 25 have dark spots on the caudal musculature and dorsal fin but lack spots on the ventral fin. These tadpoles also ha\'e a proportionately shorter tail than do tadpoles in stage .30. Matixg Call: The apparent absence of vocal slits and a vocal sac preclude the pres- ence of a mating call in this subspecies. Natliral History: The only information on the habits and habitat of this species was gi\'en by Straughan and Wright (1969, p. 8): "All individuals were collected from a system of small and medium sized streams flowing down steep slopes in pine-fir forest. Adults and juveniles were encountered sitting on rocks or piles of detritus deposited by flood. The main stream consisted of small pools (two to three m wide) with sandy bottoms partially covered with leaf litter, and small water falls and rapids (one-half to one and one-half m wide). When disturbed, the frogs jumped into the water and remained sub- merged for a short period before re-emerging. Water temperature at time of capture of frogs wasl4° C. Tadpoles were found in the larger pools resting on the bottom in quieter water." Straughan and Wright (1969, p. 8) meta- morphosed one tadpole in the laboratory. At metamorphosis the young frog had a snout- \ent length of 20.2 mm. Remarks: This species was described af- ter the manuscript for the present publication was completed. Subsequently I examined the known specimens and included data on the species in the text; however, because the illus- trations had been mounted and arranged, I was not able to insert illustrations of this spe- cies. The reader is referred to the type de- scription (Straughan and Wright, 1969) for photographs of adults and young and draw- ings of tadpoles and the mouth of a tadpole. I concur with Straughan and Wright (1969, p. 8) that Hyla bogertae is related to 482 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 the high montane complex of species con- taining crassa, pacliydermu, rohertsorum, and siopela. Until osteological data are available for all of those species, no further comments on relationships can be justified. Etymology: The specific name is a patro- nym for Martha M. Bogert. Distribution: Hyla hogertae is known only from one small stream system at an ele- vation of 2652 meters in the Sierra Madre del Sur in Oa.xaca, Mexico. See Appendix 1 for the locality records of the 15 specimens examined. The Hyla eximia Group Definition: The members of this group are moderate-sized species; males attain a maximum snout-vent length of 44 mm., and females 47 mm. The dorsum in most species is green, with or without brown spots or stripes, but in some the dorsum is tan or gray with darker spots. A dark face mask is pres- ent (except in cadaverina), and the posterior surfaces of the thighs are uniformly colored (pale yellow spots in cuphorhiacea) . The pal- pebral membrane is clear. The fingers have vestigial webbing, and the hands are no more than two-thirds webbed. Dermal fringes and appendages are absent on the limbs. A tarsal fold is present, but an axillary membrane is lacking. Males have single, median, subgular vocal sacs and usually have small, horny, nup- tial excrescences on the thumbs. The cranial elements are weakly ossified, and a large fron- toparietal fontanelle is present (fig. 242). The nasals are moderately small and not in bony contact with the sphenethmoid, which is not ossified anteriorly between the nasals. The quadratojugal is present and in contact with the maxillary. The squamosal is not in bony contact with the crista parotica, and the an- terior arm of the squamosal extends only about one-third of the distance to the maxil- lary. The columella is expanded distally. The prevomers are poorly ossified and bear teeth. The palatine is weak, and the medial ramus of the pterygoid does not articulate with the prootic. The tadpoles ha\e deep fins and small anteroventral mouths with two upper and three lower rows of teeth. The mating calls consist of a series of short notes or a series of rattling notes. The haploid number of chromosomes is ] 2. Fig. 242. Dorsal view of the skull of Hijla eximia, K.U. No. 59903. x 6. Composition: Seven species comprise the group, which is widespread in North America. One species, Hyla squirella of southeastern group, which is widespread in North America, all of the others occur in Mexico, although the greatest part of the range of Hyla regUla is in the United States. Of the six species oc- curring in Middle America, .3843 preser\ed frogs, 22 skeletons, 15 lots of tadpoles, and one clutch of eggs have been examined from Mexico and Guatemala. Additional material, principally skeletons and tadpoles, has been examined from the United States. Comments: The arrangement of species used here differs notably from that presented by earlier workers. Taylor (1939b) first rec- ognized an eximia group, in which he placed lafrentzi, regilla, euphorhiacea, and eximia; he named two other species {cardenasi and nri'^htorum ) in the same group. Taylor con- cluded that Hyla hocoiirti was a synonym of euphorhiacea and that Hyla gracilipes was a s\nonym of eximia. Taylor (1941) named Hyla arhoricola, an c.vi??na-like frog from the Sierra Madre del Sur in Guerrero. Stuart (1954b) named Hyla walkeri from Guate- mala; he suggested that ualkeri was most like arJ>oricola. Maslin (1957) mimed Hyla micro- cxiinia from Jalisco, but Duellman (1961c) showed that Maslin's species was based only on a- common pattern of eximia. Thus, for about two decades the eximia group remained only a simple assortment of nondescript tree frogs that inhabited the highlands of western North America .southward to Guatemala. Then, seemingly as though by explosive evo- 1970 DUELLMAN: HYLID FROGS 483 lution the complexities of the group multi- plied. Blair (1960) and Bogert (1960) point- ed out the apparent mosaic of call-types in eximia on the Mexican plateau. Gorman (1960) showed that the populations of "Hylci arenicolor" west of the Colorado Desert were not really arenicolor but represented a distinct species, which he named californioe and placed in the eximia group. Blair (1960) added Hijk squirella to the eximia group. The structure of the adult and tadpole, the life history, and the mating call seem to ally this vicariant species with the eximia group. However, Blair's suggestion that Hyla statifferi be placed in the eximia group is as preposterous as Kellogg's (1932) inclusion of Hijla smitlui as a synonym of exi- mia. Blair's idea of the relationships of statif- feri was based entirely on similarities in the mating call of statifferi and eximia and with- out regard to morphology and distribution. Jameson, Mackey, and Richmond (1966) presented the most diverse arrangement yet of the eximia group. On the basis of a multi- variate discriminant analysis of 10 measure- ments of each of 454 specimens they recog- nized ten subspecies of Htjla regilla, including lafrentzi and tcrightorum. Thus, where Tay- lor in 1939 recognized six taxa, 18 exist today. Most of the races of Hijla regilla do not occur in Mexico and consequently will not be dealt with here. In order that the recognizable populations can be discussed here, the taxo- nomic status of the \arious nominate species and subspecies is outlined below. Each is elaborated upon more fully in the accounts of the appropriate species. Hyla eximia possibly is a composite spe- cies comprised of two or more populations not, or but little, differentiated morphologi- cally. This possibility notwithstanding, sev- eral named species {gracilipes, cardenasi, tcrightorum, arboricola, and microeximia) are considered to be synonyms; all were distin- guished from eximia by minor morphological characters. The northern populations former- ly assigned to tcrightorum are not con- specific with Htjla regilla. Hyla euphorbiacea includes Htjla hocotirti and is specifically distinct from eximia. Hyla icalkeri is a distinct vicariant most closely re- lated to etiphorhiacea. Hyla plicata is an earher name for Hyla lafreittzi, a species distinct from regilla and occurring in partial sympatry with eximia. The Mexican populations of Htjla regilla can be assigned to two subspecies; those to the south of the 'Viscaino Desert are H. r. curta, and those to the north of the desert are con- sidered to be representatives of H. r. hypo- chondriaca {deserticola is a synonym). The frogs named Hyla calif orniae by Gor- man (1960) were originally named Hyla ne/Mv/osa by Hallowell (1854). Cope (1866a) pointed out that Hallowell's name was pre- occupied and proposed the replacement name Htjla cadaverina, which is the correct name for the species that for so many years mas- queraded under the name Hyla arenicolor and for less than a decade enjoyed specific recog- nition under a junior objective synonym (Duellman, 1968c). The definition and recognition of species in the eximia group is difficult due to subtle differences that are inconspicuous in light of gross similarities and to the absence of easily definable characters in the preserved frogs. In this respect, the species in the eximia group are like those in the Hyla microcephala group. Osteological differences among the species are lacking or insignificant. The rugose dorsal skin immediately distinguishes cadaverina and regilla from the other species in the group. Some slight \'ariation in the structure of the hands and feet and in the amount of webbing is evident (figs. 243 and 244). Diff^erences in coloration are useful specific characters but do not tend to elucidate interspecific relation- ships. Hijla euphorbiacea is distinctive by having yellow spots on the posterior surfaces of the thighs, and plicata has a white stripe on the shank; these are the only distinctive color difl[erences in the smooth-skinned spe- cies. Slight differences in size and proportions are evident (table 45). Htjla plicata is the largest species and has the longest legs. Hijla regilla and cadaverina have proportionately larger heads and smaller tympani than the other species. These differences, except for the large size and long legs of plicata, are negated in eximia by the extreme variation in that species (see account of Hijla eximia for details of variation). The differences in the tadpoles are very slight, except for those of 484 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fig. 24.3. Hands of members of the Hyla cximia group. A. Hi/la regilla, K.U. No. 109875. B. Hyla cadaverina, K.U. No. 109866, C. Hyhi eximia, U.M.M.Z. No. 119163. D. Hyla ciiphorbiacea, K.U. No. 100924. E. Hyla walked, K.U. No. 57835. F. Hyla plicata, K.U. No. 57384. x 5. cadaverina and regilla (figs. 245 and 246); unfortunately, the tadpole of plicata is un- known. The mating calls offer some excellent clues to the relationships of the species (table 46; pis. 12-14). The calls of euphorhiacea and walkeri consist of groups of quickly repeated short notes, whereas the calls of the other species are made up of equally dispersed notes. The notes produced by plicata are longer and have a lower dominant frecjuenc}' than those of the other species. The mating calls of members of sympatric pairs of spe- cies (eximia-plicata and cadaverina-regilla) differ in several parameters and doubtlessly act as important reproducti\c isolating mech- anisms. The relationships of the eximia group seem to be with the Hyla cinerea group and with Pseuclacris in North America. Hyla regilla Baird and Gixard Hyla regilla Baird and Girard, 1852, p. 174. Diagnosis: This moderately small species has small discs, little or no webbing between the fingers, and the toes about two-thirds webbed. The dorsal ground color is green, gray, tan, brown, or reddish bro\\n, and the 1970 DUELLMAN: HYLID FROGS 485 Fig. 244. Feet of members of the Hijla eximia group. A. Hijla regilla, K.U. No. 109875. B. Hijla cada- verina, K.U. No. 109866. C. Hyla eximia, U.M.M.Z. No. 119163. D. HijIa euphorbiacea. K.U. No. 100924. E. Hyla walkeri. K.U. No. 57835. F. Hyla plicata, K.U. No. 57384. x 5. 486 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fig. 245. Tadpoles of members of the Hyla cxiinia group. A. Hyla regilla. K.U. No. 118150. B. HsS'^'' Fig. 246. Mouths of tadpoles of members of the Hyla eximia group. A. Hijla rcgilla, K.U. No. 118150. B. Htjla cadaverina, K.U. No. 118149. C. Hyla eximia, K.U. No. 104133. D. Hyla euphorbiacea, K.U. No. 59988. E. Hijh walkeri, K.U. No. 60003. x 30. regilla. Duellman (1968c) showed that Hyla regiUa lafrentzi {^Hyla plicata) was spe- cifically distinct from regilla. I here conclude that Hyla regilla icrightorum is indistinguish- able from eximia but consider eximia to be specifically distinct from regilla. Furthermore, I consider Hyla regilla deserticola and hypo- chondriaca to be the same. According to this arrangement, seven subspecies are recognized; fi\e of these occur only to the north of Me.\- ico and ha\e not been studied by me. The Me.xican populations are assignable to Hyla regilla curia and H. r. hypochondriaca. DiSTRiBUTiox: Hyla regilla ranges from sea level to elevations of about .3400 meters from southern British Columbia, Canada, southward through the mountains and along the coastal regions of western United States to the southern tip of the peninsula of Baja California, Mexico, and eastward to western Montana and Idaho and eastern Nevada in the United States (fig. 247). 488 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 W < C QJ (X, 3 C 2 G OJ O -G O •= O , c >, H •5 "Be/) ■5 o ■5 bo , 2 §•5 IS CO 10 o 05 o O 05 ,^10 _^a5 t^ o o5 o ^ .^ i^ ^ O C-l o t^ o o en 05 O ^. CO O CO 00 t- o i^ in cq t^ ^ (M 1^ ^'^ CD OioOctjOcdC^CO'^COOco ;£, CO ^ CO 1. ro -i. r^ I. ro .1 ro (M O CO — ^ CO - ' '"O UJ ^-^ U^ ^— ^ "' ^— ' CO — - • .. OC000500000 CO^--CO~_''^(M'^cD<^g? ro^CO'CO-iCO'OJ. ^^ --: C* ^ C_J fC O t^ O f^ (N "-^oq — 'CM 0000 o ^ CO o ^ r-; ^ S '■^ i^ <=■ CO <— ' 05 ® " '(M - - O O ■ CO 00 C-- CD M* O ■* 05 , ,t-^ ^-^CO , ,0 , ,05 ^^05 ,— ^ "^CO'^— I^Cd''^— l"^0O"^O5 OIOO-^OCOOCDO-^OIO co;^csj;^CT>^io;3o^c6;^ (M ■<*< O Tt< t^ -* 0_^CO_,,t^,^C' cq in O cq ol Tt< cq 10 , — , CO . ^O ^-^00 ^-^ t^ ^-^ M55''^£^oo5r;oor1o t^ rH Q, Cq ^/^ C-l (£) '^l rH OcO^Ogcor^loO; c^cq-^-^c-'io'^ic^ ^-'CO ^-t^ ■ — ^CD "-T^ --^ CO o CO ,-^05 CM O CO CO 5",4. ^ r^ -< V5 -H -00 t:- .CO ■ CO —CO ^'=> '~-'=> '-^ S "^ S 2, CO CO 10 ^ ^ 2^ — 't^ Tf --'CD — LO i-H O OCq o O CO 05 ^^ CM ^-^ r-l -^ . . OCOOCMOSOCO -^c>]'^in'^4Scoq OlOO.— lOC^OO "-'O ^^o ^^o o ' ^ — ' ^-^ ' — > O ''i f,, CD CO ^ Tf ' CO CO 05 CO o ( — I 10 CO CM 00 CO CO o 5: Si S ^ .2 o a. 3 to 'a 1970 DUELLMAN: HYLID FROGS 489 32 28" 24° • H. regit la cur fa o H. regilla hypochondriaca 200 — I KILOMETERS 114° Fic. 247. Distribution of the subspecies of Hijla regilla in Mexico. 490 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Hyla regilla curta Cope Hyla curta Cope, 1887b, p. 313 [syntvpes, U.S.N. M. No. 5293 (19 specimens) from Soria, 15 miles north of Cabo San Lucas, Baja California Sur, Me.\ico; John Xanthus collector]. Hyla regilla laticeps Cope, 1889, p. 356 [syntypes, U.S.N.M. No. 5308 (7 specimens) from Cabo San Lucas, Baja California Sur, Mexico; John Xantus col- lector]. HtjUola regilla (part): Mocquard, 1889b, p. 339. Hyla regilla (part): Smith and Taylor, 1948, p. 82. Hyla regilla curia: Jameson, Mackey, and Rich- mond, 1966, p. 585. Diagnosis: This subspecies of Hyh regilki generally is more robust than hijpochondriaca and has smooth skin on the dorsum, slightly more webbing on the feet, and slightly short- er hind limbs. Most specimens of curta tend to have more diffuse dorsal markings than do those of hijpochondriaca. Description': Males of this moderately small species attain a snout-vent length of 37.8 mm., and females reach 44.1 mm. Nine males from San Ignacio, Baja California Sur, Me.xico, have snout-vent lengths of 26.6 to 30.7 (mean, 28.2) mm.; the ratio of tibia length to snout-vent length is 0.460 to 0.500 (mean, 0.477); the ratio of foot length to snout-vent length is 0.442 to 0.4S9 (mean, 0.462); the ratio of head length to snout- vent length is 0..322 to 0.350 (mean, 0..334); the ratio of head width to snout-vent length is 0..326 to 0.361 (mean, 0.340), and the ratio of the diameter of the tympanum to that of the eye is 0.344 to 0.412 (mean, 0.386). Seven females from the same locality have snout- vent lengths of 26.8 to 38.0 (mean, 31.3) mm. and a proportionately larger tympanum; the ratio of the diameter of the tympanum to that of the eye is 0.353 to 0.545 (mean, 0.440). Specimens from the southern part of the peninsula are somewhat larger and have a proportionately larger tympanum. Two males from Todos Santos, Baja California Sur, have snout- vent lengths of 35.8 and 37.8 mm.; in these specimens, the ratio of the diameter of the tympanum to that of the eye is 0.488 and 0.500. Two females from the same locality have snout-\'ent lengths of 41.6 and 44.6 mm.; in these specimens, the ratio of the diameter of the tvmpanum to that of the eve is 0.579 and 0.605. The head is as wide as the body, and the top of the head is barely convex. In dorsal and lateral profiles, the snout is rounded. The snout is moderately long; the nostrils arc no- ticeably protuberant at a point about two- thirds the distance from the eyes to the tip of the snout. The canthus is rounded; the loreal region is barely concave, and the lips are moderately thick and barely flared. A thin dermal fold extends posteriorly from the eye, above the tympanum, and downward to a point above the insertion of the arm. The fold obscures the upper edge of the tympanum, which otherwise is distinct and separated from the eye by a distance less than the diam- eter of the tympanum. The arms are moderately short and slen- der; an axillary membrane is absent. No dis- tinct row of tubercles is present on the \entro- lateral edge of the forearm, but a distinct transverse dermal fold is present on the wrist. The fingers are moderately short and bear rather small discs; the width of the disc on the third finger is somewhat less than the diameter of the eye. The subarticular tuber- cles are large and conical; none is bifid. The supernumerary tubercles are large, elevated, and conical. A distinct, bifid palmar tubercle is present. The prepollex is moderately en- larged and in breeding males bears a small nuptial excrescence. Webbing on the hands is absent. The hind limbs are moderately short and robust. The heels of the adpressed limbs o\erlap by about one-fifth of the length of the shank. The tibiotarsal articulation ex- tends to the posterior corner of the eye. A thin transverse dermal fold is present on the heel, and a distinct tarsal fold is present. The inner metatarsal tubercle is ele\'ated, elliptical, and barely visible from above. The toes are moderately long and slender and bear discs that are noticeably smaller than those on the fingers. The subarticular tubercles are mod- erately large and conical, and the supernu- merary tubercles are small, but distinct. The toes are about two-thirds webbed. The web- bing extends from the base of the penultimate phalanx of the first toe to the distal end of the antepenultimate phalanx of the second, from the middle of the penultimate phalanx of the second to the base of the antepenultimate phalanx of the third, from the distal end of 1970 DUELLMAN: HYLID FROGS 491 thf antoponultimatc phalanx of the third to the base of the antepenultimate phalanx of the fourth and on to the base of the penulti- mate phalanx of the fifth toe. The anal opening is directed posteriorly at the upper level of the thighs; a short, broad anal flap is present. The skin on the dorsum is smooth; that on the throat, belly, and proxi- mal postcro\entral surfaces of the thighs is granular. The tongue is broadly cordiform, shallowly notched posteriorly, and free behind for about one-third of its length. The dentig- erous processes of the prevomers are narrow- ly separated, transverse elevations between the posterior margins of the small, ovoid cho- anae. Males have four to six teeth on each process and a total of eight to eleven ( mean, 9.8) prevomerine teeth. Females likewise have four to six teeth on each process and a total of eight to twelve (mean, 10.3) prevo- merine teeth. The vocal sHts extend from the midlateral base of the tongue to the angles of the jaws. The vocal sac is single, median, and subgular. I ha\e no knowledge of the coloration of Hyla regiUa ctirta in life. In preservative the dorsum is dull grayish tan with darker gray- ish brown markings (pi. 1, fig. 3). All speci- mens have a distinct dark mark beginning on the snout and extending to the nostril and eye and posteriorly to the anterior part of the flank. In most individuals, this mark is bor- dered above by a narrow white line. The lips are pale grayish white. The markings consist of a pair of irregular dark dorsolateral marks, which are confluent anteriorly in some speci- mens. Most individuals possess a well-defined triangular mark on the top of the head. The dorsal surfaces of the limbs are faintly barred with dark brown, and the posterior surfaces of the thighs are dull tan. The flanks are pale gray with dark brown flecks or spots. A faint white anal stripe is present. The venter is creamy tan, and the vocal sac is heavily flecked with gray. Tadpoles: The tadpoles of this subspecies are unknown; presumably they are like those of Hyla regiUa hypochondriaca. Mating Call: The call of this subspecies consists of a series of short diphasic notes, "aah-aah, aah-aah, aah-aah." The analysis of the call of one individual from Todos Santos, Baja California Sur, reveals that the note repe- tition rate is 30 notes per minute, and that the duration of the note is about 0.10 of a second; the fundamental frequency is at 121 cycles per second and the dominant frequency at 2420 cycles per second ( pi. 12, fig. 1 ) . Natural History: I have had no first- hand field experience with this subspecies, nor has there been any published record con- cerning its ecology. The frogs apparently congregate around any depression containing moisture and breed there when sufficient wa- ter accumulates. Dr. Laurence M. Hardy ob- tained a calling male from a water-filled ditch at Todos Santos, on July 9, 1963, and he found several indi\ iduals in clumps of grass near an outlet from an artificial tank containing water. Remarks: Although the type specimens of Hyla curta and Hyla regiUa laticeps are rather faded, there is no doubt but what they represent the same species of tree frogs. All workers in the present century have regarded laticeps and curta as synonyms of Hyla regilla. Jameson, Mackey, and Richmond ( 1966, p. 585) resurrected Hyla curta Cope, 1867, as a subspecies of Hyla regiUa. On the basis of a so- phisticated mathematical analysis of a variety of measurements of possible doubtful signifi- cance, those authors partitioned the wide- spread Hyla regiUa into numerous subspecies. The populations of regiUa in the southern part of Baja California seem to be moderately distinct from those populations to the north. Consequently. I am recognizing the southern population as Hijla regiUa curta. Etymology: The subspecific name is de- rived from the Latin ciirtus, meaning short, and possibly refers to the length of the legs in this subspecies. Distribution: Hyla regilla curta occurs at elevations from sea level to approximately 1000 meters in the peninsula of Baja Califor- nia south of the Desierto de Vizcaino, Mexico (fig. 247). See Appendix 1 for the locality records of the 98 specimens examined. Hyla regilla hypochondriaca Hallowell Ihjla scapularis \ar. hypochondriaca Hallowell, 1854, p. 97 [synt>-pes, U.S.N.M. 3235 (8 specimens) from Tejon Pass, Los Angeles County, California, U.S.A.; A. L. Heermann collector]. Hyla regilla (part) : Smith and Taylor, 1948, p. 82. 492 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Htjla regilla descrticola Jameson, Mackey, and Richmond, 1966, p. 582 [holotype, S.D.N.H.M. 54176 from San Borja, Baja California del Norte, Mexico; David L. Jameson collector]. Hyla regilla htjpochondiiaca: Jameson, Mackey, and Richmond, 1966, p. 588. Dl\gnosis: This subspecies of Hijla regilla is more slender than ciirta and has slightly longer hind limbs and slightly less webbing. The skin on the dorsum is weakly pustulate in some specimens, whereas it is smooth in cur- ta. The dorsal markings in hypochondriaci are more distinct than in curt a. Description: Males of this subspecies at- tain a snout- vent length of 37.1 mm., and females reach 38.2 mm. In a series of 25 males from Ramona, San Diego County, Cali- fornia, the snout-vent length is 27.8 to 36.9 (mean, 32.9) mm.; the ratio of tibia length to snout- vent length is 0.451 to 0.502 (mean, 0.477); the ratio of foot length to snout- vent length is 0.407 to 0.498 (niean, 0.452); the ratio of head length to snout-vent length is 0.312 to 0.346 (mean, 0..327); the ratio of head width to snout-vent length is 0.344 to 0.378 (mean, 0.359), and the ratio of the diameter of the tvmpanum to that of the eye is 0.424 to 0.624 (mean, 0.540). Morphologically this subspecies is like Hyla regilla curia, except that the skin on the dorsum is weakly pustulate in some speci- mens. Most indi\'iduals possess a row of tu- bercles along the ventrolateral edge of the forearm. The webbing is vestigial or absent on the hands (fig. 24,3A), and the feet are somewhat more than one-half webbed (fig. 244A). Males have three to five teeth on each prevomerine process and a total of si.x to ten (mean, 7.2) prevomerine teeth. Specimens from Ramona, San Diego Coun- ty, California are highly variable in dorsal coloration (pi. 65, figs. 1-4). The dorsum is green with darker green markings, tan with brown markings, grayish tan with grayish brown markings, or reddish brown with dark brown markings. The flanks are creamy white, pale grayish tan with brown flecks. The groin, anterior and posterior surfaces of the thighs, the inner edges of the tarsi, the bases of the first and second toes, and the ventral surfaces of the hind limbs are dull yellow. There is a narrow white canthal and supratympanic white stripe and an indistinct white anal stripe. A broad white labial stripe is present, a dark brown stripe extends from the tip of the snout, through the nostril and eye, and posteriorly to a point above the insertion of the arm; from that point it continues on to the flank as a series of brown spots in some specimens. The belly and ventral surfaces of the arms are creamy white. The vocal sac is dull yellow with greenish gray flecks. The iris is dull bronze with a median horizontal brown streak. In preservative, the dorsum varies from pale bluish gray and yellowish tan to dull grayish brown. Most individuals have a pat- tern consisting of a pair of longitudinal dark brown stripes that are continuous or inter- rupted one or more times. All specimens have some form of a triangular dark mark between the eyes with the apex of the triangle directed posteriorly. The dorsal surfaces of the limbs are strongly banded with dark brown. T.\DPOLES: A typical tadpole in develop- mental stage 35 has a body length of 12 mm. and a total length of 30 mm. The body is ro- bust and ^s deep as wide. In dorsal profile, the snout is bluntly rounded, and in lateral profile it is round. The eyes are widely sepa- rated and directed laterally. The nostiils are directed anterolaterally at a point somewhat closer to the eyes than to the tip of the snout. The opening of the sinistral spiracle is di- rected posterodorsally at a point just below the midline about two-thirds of the distance from the tip of the snout to the posterior end of the body. The anal tube is dextral. The caudal musculature is slender and tapers grad- ually to the tip of the rounded tail. The fins are deep, at midlength of the tail, both the dorsal and ventral fins are about half again as deep as the caudal musculature. The dor- sal fin extends onto the body (fig. 245A). The coloration in life was described by Gaudin (1965, p. 122). He noted that in early developmental stages the body is rather e\en- Iv co\ered with melanophores and that gold- en chromatophores and a few guanophores are scattered over the dorsal and lateral parts of the body. He stated that the dorsal and lateral parts of the tail musculature ha\e me- lanophores scattered throughout with a sprinkling of golden chromatophores and guanophores. Gaudin noted that in stage 30 1970 DUELLMAN: HYLID FROGS 493 the distribution of melanophores is relatively stable. He stated: "Anterior to the spiracle, melanophores occur in a rather heavy concen- tration dorsally and extend down to the \'en- tral surface of the body, while posterior to the spiracle, melanophores extend only one-half to two-thirds of the distance down the sides of the body. The intestines are still completely obscured by an opaque layer of melanophores lining the coelom. Golden chromatophores and guanophores are scattered over the dor- sal and lateral parts of the body and tail musculature and contribute \arying amounts of sheen to the body, depending on the degree of contraction of the chromatophores." In preservative, the tadpoles are dull brown above and transparent below. The caudal musculature is creamy tan with dense brown flecks, especially anteriorly. The fins are trans- parent and are flecked with brown above and distalK- on the ventral fin. The mouth is moderately small and antero- ventral in position. The lips have a shallow lateral fold. The median half of the upper lip is bare; the lower lip is bordered by a single row of blunt papillae, but two rows of pa- pillae are present laterally. The beaks are moderately robust and bear blunt serrations. The upper beak is in the form of a broad arch with robust, short lateral processes; the lower beak is broadly V-shaped. There are two up- per and three lower rows of teeth. The upper rows are equal in length, and the second up- per row is broadly interrupted medially. The first and second lower rows are nearly as long as the upper rows, whereas the third lower row is short ( fig. 246A ) . Mating Call: The call of Hyla rcgiUa hypochondhaca consists of a series of short, usually diaphasic notes (see Snyder and Jameson, 1965, p. 1.31 ) for a discussion of the variation in mating call in Hyla regiUa). Natur.\l History: No observations on the natural history of this subspecies have been made in Me.xico; the reader is referred to a general account of the species bv Stebbins (1951, 1. 322). Remarks: Although Jameson, Mackey, and Richmond (1966, p. .5.54) arrived at the con- clusion that hypochondriaca was an available name for a population of Hyla regiUa in southern California, thev did so without the benefit of examination of the type in question and miraculously were correct in their con- clusions. Jameson, Mackey, and Richmond (1966) analyzed the variation in Hyla regiUa and recognized ten computer-generated sub- species, some of which are distinguished on extremely superficial characters. I have been unable to justify the recognition of two sub- species of Hyla regilla in northern Baja Cali- fornia, Mexico. Consequently, I conclude that Hyla regilla cleserticola Jameson, Mackey, and Richmond, 1966, is a synonym of Hyla regilla hypochondriaca Hallowell, 1S54. Etymology: The subspecific name is de- rived from the Greek, hypochondriakos, meaning literally of the abdomen. I am un- sure of its reference to the frog concerned. DisTRiBUTiox: Hyla regiUa hypochondria- ca occurs at elevations from sea level to about 1400 meters from the northern end of the interior valley of California southward through southern California and extreme southern Nevada to the northern half of the peninsula of Baja California, Mexico; the subspecies also occurs on the islands off the Pacific coast of California and Baja Califor- nia (fig. 247). See Appendix 1 for the locality records of the 109 specimens examined. Hyla cadaverina Cope Hyla arenicolor (part): Kellogg, 1932, p. 156. Smith and Taylor, 1948, p. 89. Hijla nebulosa Hallowell, 1854, p. 96 [syntypes, A.xN.S.P. Nos. 1987 and 1988 from Tejon Pass, Los Angeles Countj-, California, U.S.A.; A. L. Heemiann collector (not Hyla nebulosa Spi.x, 1824, from Brazil)]. Hyla cadaverina Cope, 1866a, p. 84 [replacement name for Hyla nebulosa Hallowell, 1854, preoccupied by Hyla nebulosa Spi.x, 1824]. Duelbnan, 1968c, p. 200. Hyla californiae Bogert, 1958, p. 11 [iionien nu- dum]. Hyla californiae Gorman, 1960, p. 214 [holotype, M.V.Z. No. 31773 from Canon de Llanos, 9 miles south-southwest of "Alaska" (La Rumorosa), Baja California del Norte, Mexico; Robert R. Miller and J. Davis collectors]. Diagnosis: This moderately small species has tubercular skin on the dorsum and lacks webbing on the hand; the feet are about three- fourths webbed. The dorsum is gray or brown with numerous small, irregular spots. This species diflPers from all other members of the 494 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 eximia group by having a distinctly tubcrcu- late dorsum, small dorsal spots, and no dark brown band on the side of the head posterior- ly onto the body. Hyla caclaverina differs from regilla by being more pustulate and by having more webbing on the feet (web to base of penultimate phalanx of the fourth toe in ca- daverina and only to base of antepenultimate phalanx in regiUa). Hyla cadaverina resem- bles arenicolor, which is somewhat larger and differs by having less webbing on the feet (about one-half webbed), larger discs, more numerous and distinct supernumerary tuber- cles, and a larger tympanum; the diameter of the tympanum in caclaverina is about half of the diameter of the eye, whereas in arenicolor it is about two-third of the diameter of the eye. Other Middle American hylids that might be confused with caclaverina all have web- bing between the fingers. Description: Males of this moderately small species attain a snout-vent length of 36.0 mm., and females reach 45.0 mm. In a series of 16 males from Boulder Park, San Diego County, California, the snout-vent length is 29.0 to 35.9 (mean, 33.0) mm.; the ratio of tibia length to snout-vent length is 0.474 to 0.523 (mean, 0.503); the ratio of foot length to snout-vent length is 0.432 to 0.477 (mean, 0.447); the ratio of head length to snout-vent length is 0.334 to 0.368 ( mean, 0.354); the ratio of head width to snout-vent length is 0.383 to 0.417 (mean, 0.397), and the ratio of the diameter of the tympanum to that of the eye is 0.432 to 0.529 (mean, 0.478). Nine females from the same locality have snout-vent lengths of 39.4 to 43.9 ( mean, 40.9) mm. and do not differ significantly in proportions. The head is slightly broader than the body, and the top of the head is barely convex. In dorsal profile the snout is acutely rounded; in lateral profile it is round. The canthus is round and barely evident; the loreal region is slightly concave and the lips are moderately thick and not flared. A thin dermal fold ex- tends posteriorly from the eye, above the tym- panum, and downward to a point just pos- terior to the angle of the jaw. The fold ob- scures the upper edge of the tympanum, which otherwise is distinct and separated from the eye by a distance equal to about one-half of the diameter of the tympanum. The arms are moderately short and slen- der; no axillary membrane is present. A row of low tubercles is present on the ventrolateral edge of the forearm, and a weak transverse dermal fold is present on the wrist. The fin- gers are moderately long and slender and bear relatively small discs; the width of the disc on the third finger is equal to about two- thirds of the diameter of the tympanum. The discs are truncate. The subarticular tubercles are moderately large and conical; in some individuals one or more tubercles are bifid. The supernumerary tubercles are small and conical. A large, flat palmar tubercle is pres- ent. The prcpollex is moderately enlarged and in breeding males bears a weak nuptial ex- crescence. Webbing is absent between the fingers (fig. 243B). The hind limbs are mod- erately short and slender; the heels of the adpressed limbs barely overlap. The tibio- tarsal articulation extends to the eye. A weak tarsal fold is evident distally on the tarsus; a few small tubercles are present on the outer edge of the tarsus. The inner metatarsal tu- bercle is moderately small, ovoid and flat. No outer metatarsal tubercle is evident. The toes are moderately long and slender and bear small discs. The subarticular tubercles are small and conical, and the supernumerary tubercles are minute. The toes are about three-fourths webbed (fig. 244B). The web- bing extends from the base of the penultimate phalanx of the first toe to the middle of the antepenultimate phalanx of the second, from the middle of the penultimate phalanx of the second to the middle of the antepenultimate phalanx of the third, from the middle of the penultimate phalanx of the third to the base of the penultimate phalanx of the fourth and on to the base of the penultimate phalanx of the fifth toe. The anal opening is directed posteriorly near the upper level of the thighs; a short anal sheath is present. The skin on the dorsum has numerous scattered tubercles; that on the throat, chest, and proximal posteroventral sur- faces of the thighs is granular. Elsewhere, the skin is smooth. The tongue is narrowh- cordi- form, shallowly notched posteriorly, and bare- ly free behind. The dentigerous processes of 1970 DUELLMAN: HYLID FROGS 495 the prevomers are small o\oid ele\ations be- tween the round choanae. Males ha\e two to four teeth on each process and a total of four to seven ( mean, 6. 1 ) prevomerine teeth. The vocal slits extend from the midlateral base of the tongue towards the angles of the jaws. The vocal sac is single, median, and subgular. The general coloration of Hyla cadaverina is dull grayish browai or olive-brown with darker spots dorsally (pi. 64, fig. 1). In liv- ing individuals from San Diego Count)', Cali- fornia, the dorsum is dull oIi\e-brown with dull olive-green spots on the back and bars on the hmbs. The flanks are pale olive-tan with dull olive-green flecks. The groin, an- terior surfaces of the thighs, ventral surfaces of the shanks, and the inner surfaces of the tarsi are pale dull yellow. The posterior sur- faces of the thighs are a darker dull yellow. The anterior part of the throat is dark gray with white flecks; the posterior part of the throat and the belly are pale white. A nar- row silvery cream labial stripe is present. The iris is pale bronze with black reticulations and a median horizontal brown streak. In preservative the dorsum varies from dull gray to dull brown with darker markings, which consist of small irregular spots scattered over the dorsum and transverse bars on the dorsal surfaces of the limbs. The flanks are pale tan or pale gray with small dark flecks. The venter is dull white, and the posterior surfaces of the thighs are creamy tan. A faint white anal stripe is present and numerous white tipped tubercles are evident in the anal region. Tadpoles: A typical tadpole in develop- mental stage 25 has a body length of S.2 mm. and a total length of 18.2 mm. The body is as wide as deep; the snout in dorsal profile is bluntly rounded and in lateral profile, it is round. The eyes are moderately small, broad- ly separated, and directed laterally. The nos- trils are directed anterolaterally at a point about midway between the eyes and the tip of the snout. The opening of the sinistral spiracle is directed posterodorsally at a point slightly below the midline at about two-thirds of the distance from the tip of the snout to the posterior end of the body. The anal tube is short and de.xtral. The caudal musculature is slender and does not extend to the tip of the bluntly rounded tail. The caudal fins are deep; at midlength of the tail, the depth of the dorsal fin is half again the depth of the caudal musculature. The dorsal fin does not extend on to the body ( fig. 245B ) . In life the dorsal and lateral surfaces of the body are dark brown, whereas the venter has a yellowish or cream tinge. Dark irregu- lar blotches are present on the caudal muscu- lature. A few brown flecks are present on the dorsal fin. In preservative, the tadpoles are pale brown; the caudal musculature is creamy tan u'ith dark brown blotches tending to form transverse bands on the dorsal surface of the musculature. The mouth is moderately small and di- rected anteroventrally. Weak lateral folds are present in the lips. The median two-thirds of the upper lip is bare; elsewhere, the lips are bordered by a single row of large, elongate papillae. The beaks are moderately slender and bear long, blunt serrations. The upper beak is in the form of a high, acutely rounded arch with long, slender lateral processes; the lower beak is broadly U-shaped. There are two upper and three lower rows of teeth. The upper rows are equal in length, and the sec- ond upper row is broadly interrupted me- dially. The first and second lower rows are equal in length and somewhat shorter than the upper rows, whereas the third lower row is much shorter than the others (fig. 246B). Gaudin ( 1964 ) described the tadpole of this species under the name of Hyla califor- niac. In 1965, he compared the lar\al devel- opment of this species with that in Hyla re- gilla. M.\TiNG Call: The call of Hyla cadav- erina consists of a long series of short notes: "aah-aah-aah." The analysis of recordings of four individuals from Sentenac Caiion, San Diego County, California, reveals that the note repetition rate is 44 to 50 (mean, 46.7) notes per minute and that the notes have a duration of 0.12 to 0.15 (mean, 0.1.35) of a second. The pulse rate is 125 to 135 (mean, 131.2) pulses per second. The fundamental frequency varies from 130 to 137 (mean, 131.7) cycles per second and the dominant frequency varies from 2055 to 2080 (mean, 2073) cycles per second (pi. 12, fig. 2). N.A.TURAL History: According to Gorman 496 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 (1960, p. 220) Htjla cadaverina occurs in canyons where they are rarely found in trees, but usually are found on rocks adjacent to pools of water in the bottom of the canyon. The frogs breed following rain storms from Mid-March to mid-June. Remarks: Duellman (1968c, p. 200) dis- cussed the allocation of the specific name ca- daverina, which is a replacement name for Hyla nebtdosa Hallowell, 1854, preoccupied by Htjla nebidosa Spix, 1824. For many years Hijla cadaveww masqueraded as a western population of Hyla arenicolor. Bogert ( 1950, p. 11) showed that the California populations of "Hyla arenicolor" had a distinctively differ- ent caU from that of Hyla arenicolor east of the Mojave and Colorado deserts. Gorman (1960, p. 214) named the western population Hyla califortiiae, a name that has been used subsequent to 1960, until Duellman (1968c, p. 200) showed that cadaverina was an earlier name for the western population of tree frogs that characteristically inhabits canyons. Etymology: The specific name is derived from the Latin cadaver, meaning corpse, and the diminitivc suffix -ina and means literally, little corpse, possibly in allusion to the pallid appearance of the species. Distribution: Hyh cadaverina occurs at elevations usually less than 500 meters in the mountains of the southern part of California in the United States and in the northern part of the peninsula of Baja California, Mexico (fig. 248). Sec Appendix 1 for the locality records of the 44 specimens examined. Hyla plicata Brocchi Hyla pUccita Brocchi, lS77b, p. 126 [holot\-pe, M.N. H.N. No. 6317 from "Mexico"; Marie-Firiiiin Bocourt collector]; 1882, p. 35. Boulenger, lS82a, p. 396. Gunther, 1901 (1885-1902), p. 261. Kellosg, 1932. p. 173. Smith and Taylor, 1948, p. 88. Duell- man, 1968c, p. 201. Hyla lafrentzi Mertcns and VVolterstorff, 1929, p. 235 [holotype, M.M. No. 49/27 from Desierto de los Leone.s, Distrito Federal, Mexico; K. Lafrentz collector (holotype destroyed; S.N.M. No. 30997 from the same locality designated as neotype by Jameson, Mackey, and Richmond, 1966, p. 596]. Smith and Taylor, 1948, p. 84. Hyla gracilipcs: Kellogg, 1932, p. 168. Hyla regilla lafrentzi: Jameson, Mackey, and Rich- mond, 1966, p. 596. 116° 114° 1 1 V •• r -.. 32° *K 1 ^^ 32° 30° 0 100 ^^ N"^ 30° KILOMETERS \ 1 K • 116° 114° Fig. 248. Distribution of Hyla cadaverina in Mexico. Diagnosis: This medium-sized green frog has a brown face and brown postorbital band extending to midflank, bordered above by a narrow white line; the fingers lack webbing; and the toes are about two-thirds webbed, and the discs are small. The posterior sur- faces of the thighs are uniform brown, and the dorsal surfaces usually are uniform green. The smooth skin on the dorsum and absence of many small irregular spots or a large inter- orbital triangular mark distinguish plicata from cadaverina and regilla. Hyla euphorhia- cea differs by having many yellow spots on the posterior surfaces of the thighs. Hyla eximia and walkeri are slightly smaller (males to 36 mm. as compared with 44 mm. in pli- cata) and have slightly less webbing on the feet; the web usually extends only to the distal end of the antepenultimate phalanx of the fifth toe in eximia and ualkeri, whereas it ex- tends to the base of the penultimate phalanx in plicata. The dorsum in plicata is green with or without a pair of dorsolateral brown longitudinal marks posteriorly; the dorsum in tcalkeri is similarly marked in most specimens, whereas the dorsal pattern in eximia usually consists of dark stripes and/or spots. Description: Males of this medium-sized species attain a snout-vent length of 44.0 mm., and females reach 47.4 nuu. In a series of 15 males from El Chico Parque Nacional, Hi- 1970 DUELLMAN: HYLID FROGS 497 dalgo, Mexico, the snout-vent length is 36.7 to 41.6 (mean, 39.7) mm.; the ratio of tibia length to snout-vent length is 0.482 to 0.570 (mean, 0.501); the ratio of foot length to snout-vent length is 0.449 to 0.525 (mean, 0.479); the ratio of head length to snout-\cnt length is 0.291 to 0.316 (mean, 0.306); the ratio of head width to snout-vent length is 0.337 to 0.379 (mean, 0.360), and the ratio of the diameter of the tympanum to that of the eye is 0.500 to 0.650 (mean, 0.570). Five fe- males from the same locality have snout-vent lengths of 37.9 to 47.4 (mean, 43.8) mm. and a proportionately larger tympanum; the ratio of the diameter of the tympanum to that of the eye is 0.590 to 0.684 (mean, 0.613). In a series of 25 males from San Gregorio, Michoa- can, Mexico, the snout-\ent length is 32.7 to 39.0 (mean, 36.7) mm., and the ratio of the diameter of the tympanum to that of the eye is 0.444 to 0.583 (mean, 0.504). Although the frogs from San Gregorio arc smaller and have a proportionately smaller tympanum than those from El Chico, the other proportions are nearly the same. The head is slightly narrower than the body, and the top of the head is barely con- vex. The eyes are large and prominent. The snout in dorsal profile is rounded; and in lat- eral profile it is bluntly rounded. The snout is short; the nostrils are protuberant at a point about three-fourths of the distance from the eyes to the tip of the snout. The canthus is rounded but evident, and the loreal region is noticeably concave; the lips are moderately thick and barely flared. A moderately heavy dermal fold extends posteriorly from the eye, above the tympanum, and downward to a point behind the angles of the jaws. The fold obscures the upper edge of the tympanum, which otherwise is distinct and separated from the eye by a distance slightly less than the diameter of the tympanum. The arms are moderately short and somewhat robust; an axillary membrane is absent. A row of low tu- bercles is present on the ventrolateral edge of the forearm, and a distinct transverse der- mal fold is present on the wrist. The fingers are moderately long and slender and bear small discs; the width of the disc on the third finger is equal to about three-fifths of the diameter of the eye. The subarticular tuber- cles are large and round; none is bifid. The supernumerary tubercles are large and coni- cal. A large, elevated partially bifid palmar tubercle is present. The prepollex is moder- ately enlarged and bears a horny nuptial ex- crescence in breeding males. Webbing is ab- sent between the fingers ( fig. 243F ) . The hind limbs are moderately long and slender; the heels of the adpressed limbs overlap by about one-fourth of the length of the shank. The tibiotarsal articulation extends to the anterior corner of the eye. A thin transverse dermal fold is present on the heel, and a strong flap- Hke tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is mod- erately large, flattened, and elliptical. A coni- cal outer metatarsal tubercle is present. The toes are long and slender and bear small discs. The subarticular tubercles are moderately large and round, and the supernumerary tu- bercles are large, subcorneal, and numerous on the proximal segments of each digit. The toes are about two-thirds webbed (fig. 244F). The webbing extends from the base of the penulti- mate phalanx of the first toe to the distal end of the antepenultimate phalanx of the second, from the middle of the penultimate phalanx of the second to the base of the antepenulti- mate phalanx of the third, from the base of the penultimate phalanx of the third to the base of the antepenultimate phalanx of the fourth and on to the base of the penultimate phalanx of the fifth toe. The anal opening is directed posteriorly at the upper level of the thighs; it is covered by a short, broad anal sheath. The skin on the dorsum is smooth; that on the throat, belly, and posteroventral surfaces of the thighs is heavily granular. The tongue is cordiform, shallowly notched behind, and barely free posteriorly. The dentigerous processes of the pre\'omers are small, transverse or slightly posteromedially inclined, narrowly separated elevations between the anterior margins of the small, ovoid choanae. Males have four to six teeth on each process and a total of 8 to 12 (mean, 10.3) prevomerine teeth. Females have five to seven teeth on each process and a total of 10 to 14 (mean, 11.8) prevomerine teeth. The vocal slits extend from the mid- lateral base of the tongue to the angles of the 498 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 jaws. The \ocal sac is single, median, and subgular. The general coloration of Hyla plicata is dark green with a brown lateral stripe ( pi. 66, fig. 2). In most individuals the dorsum is dark green and is marked only by a dorso- lateral brown stripe or series of dashes con- necting in the sacral region and extending to, or nearly to, the vent. A dark brown stripe ex- tends from the tip of the snout, through the nostril, eye, and tympanum, and thence onto the flank; in most individuals, the brown stripe is continuous to the groin. The brown stripe is bordered above by a narrow white line. The outer edges of the feet, shanks, and forearms are dark brown, bordered above by narrow white lines. A faint white anal stripe and a narrow white labial stripe are present. The posterior surfaces of the thighs are uniform dull tan, and the venter is creamy white. The vocal sac in breeding males is gray with white flecks. The iris is dull bronze. In preservative the dorsum is dark bluish gray, and the venter is creamy tan. The pos- terior surfaces of the thighs are dull tan to dark brown. The dark markings on the side of the head and body are present in all speci- mens. The only noticeable variation in color pattern is in the dorsal markings, which are absent in some specimens. A few individuals have a dark brown stripe extending anteriorly to the scapular region, and in some specimens small round, brown spots are present pos- teriorly on the dorsum in addition to the longi- tudinal brown markings. Tadpoles: The only tadpoles available for this species are recently hatched ones that are unusable for description of the larval charac- teristics. Mating Call: The call of Hyla plicata consists of a long, low note, "waah." The anal- ysis of caUs of three individuals revealed the presence of four to 16 notes in a call-group and a note repetition rate of 24 to 60 (mean, 40) notes per minute. The duration of the notes varies from 0.52 to 0.72 (mean, 0.63) of a second, and the pulse rate varies from 78 to 98 (mean, 90) pulses per second. The fundamental frequency varies from S3 to 109 (mean, 96) cycles per second, and the domi- nant frequency varies from 1328 to 1632 (mean, 1495) cycles per second (pi. 14, fig. 3). Natural History: Hyla plicata inhabits humid pine and fir forests. At El Chico Parque Nacional, Hidalgo, Mexico, in the month of June in 1960, 1962, and 1966, calling males were found on rocks in the surface of the wa- ter in a quiet pool in a stream in a meadow, on junipers and bunch grass at the edge of a meandering stream in a meadow and from the ground at the edge of a shallow pond. A clasping pair was observed in the water of a shallow pond in the meadow on June 16, 1966; the following morning a clutch of eggs was found attached to sticks in the water. Taylor (1939b, p. 436) stated: "The specimens col- lected near Vigas, Veracruz, were found about a small rainpool beside the highway during the morning. The males were calling. Those taken at Zcmpoala were calling most of the day. A single pair was found clasping. A few immature tadpoles, presumably of this species were found in small pools in the bog near the lake edge." Duellman (1961c, p. 50) reported adults and recently metamorphosed young from be- neath logs and rocks in a damp canyon on the west slope of Cerro San Andres, Michoacan, Mexico, in March. The limited obser\ations on breeding sites suggest that this species probably utilizes small temporary pools in montane meadows as well as quiet pools in the streams. It is highly unlikely that the tad- poles are adapted for life in torrential streams. Remarks: Duellman (1968c, p. 201) res- urrected Brocchi's name Hyla plicata for those frogs that had been known as Hyla lafrentzi Mertens and Wolterstorff. Jameson, Mackey, and Richmond ( 1966, p. 596 ) placed lafrentzi {=zplicata) as a subspecies of Hyla regilla. Duellman ( 196Sc, p. 203) noted the extreme ditterences in mating calls, as well as different morphological characters between regilla and plicata and concluded that plicata was a spe- cies distinct from regilla and from eximia, which occurs sympatrically with lafrentzi in the lower part of the range of the latter. Jameson, Mackey, and Richmond ( 1966, p. 555) suggested that Hyla cardenasi was a synonym of Hyla lafrentzi (^plicata). Ex- amination of the type specimen of cardenasi reveals that it is identical with eximia; there- fore, the suggestion of these authors should be disregarded. 1970 DUELLMAN: HYLID FROGS 499 Etymology; The specific name is Latin, meaning folded; I am uncertain as to the significance of the name with reference to this species of frog. Distribution: Hyla plicata occurs princi- pally at high elexations (2400 and 3600 me- ters) in pine and fir forest in the mountains of the Sierra Madre Oriental and the Cordil- lera \'olcanica along the southern edge of the Mexican Plateau (fig. 249). The species oc- curs at somewhat lower elevations on eastern slopes of the Sierra Madre Oriental in central Veracruz, where specimens have been ob- tained between 1400 and 1500 meters in the vicinity of Vigas. See Appendix 1 for the locality records of the 403 specimens examined. Hyla eximia Baird Hyla eximia Baird, 1854, p. 61 [syntypes, U.S.N.M. No. 3248 (2 .specimens) from "Valley of Me.xieo," (Distrito Federal), Me.xieo; William Rich collector]. Brocchi, 1882, p. 32. Boulenger, 1882a, p. 378. Gunther, 1901 ( 1885-1902), p. 261. Kellogg, 1932, p. 164. Smith and Taylor, 1948, p. 83. Hyla gracilipes Cope, 1865b, p. 194 [syntypes, U.S.N.M. No. 15318-15321 from Mirador, Veracruz, Me.\ico; Charles Sartorius collector], Brocchi, 1882, p. 36. Boulenger, 1882a, p. .378. Gunther, 1901 ( 1885-1902), p. 262. Kellogg, 1932, p. 168. HyJa eximia eximia: Cope, 1887, p. 14. Hyla cardenasi Taylor, 1939b, p. 430 [holotype, U.S.N.M. No. 84403 from Puebla, Puebla, Me.xico; H. Ruano collector]. Smith and Taylor, 1948, p. 83. Hyla wrightorum Taylor, 1939b, p. 436 [holotype. U.M.M.Z. 79141 from 11 miles south of Springerville, .\pache County, .\rizona, U.S.A.; Irving J. Cantrall collector]. Smith and Taylor, 1948, p. 84. Hyla arborieola Taylor, 1941, p. 118 [holotype, F.M.N. H. No. 100131 (formerly E.H.T.-H.M.S. No. 24556) from 6 miles east of Omilteme, Guerrero, Mexico; Edward H. Taylor collector]. Smith and Taylor, 1948, p. 83. Hyla eximia wrightorum: Schmidt, 1953, p. 71. Hyla microeximia Maslin, 1957, p. 81 [holotype, U.S.N.M. No. 139246 from 3 miles northwest of Joco- tepec, Jalisco, Me.xico; T. Paul Maslin collector]. Hyla regilla wrightorum: Jameson, Mackey, and Richmond, 1966, p. 594. Diagnosis: This moderately small species with smooth dorsal skin, small discs, no web- bing between the fingers, and the toes about two-thirds webbed has a brown face mask, uniformly tan posterior surfaces of the thighs, and a green dorsum that is variously marked with a linear arrangement of brown spots or stripes in most specimens. Hyla eximia differs from cadaverina in color (green instead of brown or gray) and in lacking the tubercular skin of cadaverina. Hyla regilla can be dis- tinguished from eximia by the presence in the former of a dark interorbital triangular-mark and dark spots or flecks on the flanks; in eximia the dorsal and lateral color is sepa- rated by a narrow white line, below which the flanks are uniform creamy tan. Hyla eu- phorbiacea differs from eximia by having yel- low spots on the posterior surfaces of the thighs. Hyla plicata and walkeri are extreme- 104° 100° 20' ~n r "< i ^<:;=0'"^'-^-\ I \ 104° 100° 96° Fig. 249. Distribution of Hyla plicata. 500 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 ly difficult to distinguish from eximia; both usually lack transverse bars on the dorsal surfaces of the thighs and either lack dorsal markings or have only a pair of short brown lines posteriorly. In most specimens of eximia transverse bars are present on the thighs and the dorsum is marked by spots and/ or dark lines. Hyla plicata is larger (males to 44 mm.) than eximia (35 mm.). Description: Males of this moderately small species attain a maximum snout-vent length of 35.0 mm., and females reach 36.2 mm. In a series of 25 males from 3.2 kilo- meters west of Arandes, Jalisco, Mexico, the snout-vent length is 24.6 to 30.9 (mean, 27.8) mm.; the ratio of tibia length to snout-vent length is 0.432 to 0.495 (mean, 0.457); the ratio of foot length to snout-vent length is 0.427 to 0.478 (mean, 0.446); the ratio of head length to snout-vent length is 0.278 to 0.326 (mean, 0.307); the ratio of head width to snout-vent length is 0.304 to 0.371 (mean, 0.337), and the ratio of the diameter of the tympanum to that of the eye is 0.500 to 0.680 (mean, 0.572). Three females from the same locality have snout-vent lengths of 27.2 to 29.4 (mean, 28.5) mm. They exhibit no sig- nificant differences in proportions from the males. A mosaic of minor variation in sizes and proportions exist throughout the range of this species; this variation is illustrated in part by five samples (table 47). The head is narrower than the body, and the top of the head is barely convex. In dor- sal profile the snout is acutely rounded; in lateral profile it is round. The snout is moder- ately long; the nostrils are barely protuberant at a point about three-fourths of the distance from the eyes to the tip of the snout. The canthus is rounded, and the loreal region is barely concave; the lips are moderately thin and barely flared. A thin dermal fold extends posteriorly from the eye, above the tympanum, and downward to a point posterodorsal to the angles of the jaws. The fold obscures the up- per edge of the tympanum, which otherwise is distinct and separated from the eye by a distance equal to about one-half of the diam- eter of the tympanum. The arms are moderately short and slen- der; an axillary membrane is absent. A row of low, inconspicuous tubercles is present on the ventrolateral edge of the forearm, and a distinct transverse dermal fold is present on the wrist. The fingers are moderately long and slender and bear small discs; the width of the disc on the third finger is equal to about three-fifths of the diameter of the tym- panum. The subarticular tubercles are mod- erately large and round; none is bifid. The supernumerary tubercles are conical, conspic- uous, and numerous on the proximal segments of each digit. An elevated palmar tubercle is present. The prepollex is moderately large and in breeding males lacks a horny nuptial excrescence. Webbing is absent on the hands (fig. 24.3C). The hind limbs are short and moderately robust; the heels of the adpressed limbs barely overlap. The tibiotarsal articula- tion extends to the tympanum or to the pos- terior corner of the eye. A thin transverse dermal fold is present on the heel, and a strong tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is elevated and ovoid. A small conical outer metatarsal tubercle usually is evident. The toes are long and slender and bear discs that are about the same size as those on the fingers. The subarticular tubercles are moderately large and conical, and the supernumerary tu- bercles are small and usually evident only on the proximal segments of each digit. The toes are a little more than one-half webbed (fig. 244C). The webbing extends from the base of the penultimate phalanx of the first toe to the base of the penultimate phalanx of the second, from the base of the penultimate pha- lanx of the second to the base of the ante- penultimate phalanx of the third, from the distal end of the antepenultimate phalanx of the third to the base of the antepenultimate phalanx of the fourth and on to the distal end of the antepenultimate phalanx of the fifth toe. The anal opening is directed posteriorly near the upper edge of the thighs; a short anal sheath is present. The skin on the throat, belly, and proximal posteroventral surfaces of the thighs is granular; elsewhere the skin is smooth. The tongue is cordiform, shallowly notched posteriorly and free behind for about one-fourth of its length. The dentigerous pro- cesses of the prevomers are small, wid(>ly sepa- rated medially, posteromedialK' inclined pro- cesses between the small ovoid choanae. 1970 DUELLMAN: HYLID FROGS 501 -2 'a 4-' C 05 O a o PL, C ctf QJ _N cJ5 c c o > 3 C 0) e •5 c J J3 o r DC - C i> r-i o t- oq =Po "^ oT'^ (N «. CO S (M oix)Oc] '^ -*- CO Ol CO CO CO (NO'— 'OOO'-^oroo CO^^CO-— ^CO- -co — -(M- - CO t^ CO ro CO O CD CO t^ CO ^ CO Tji ro CO ^ CI t^ I.-.; ^ ■— I O O O O O O CO ~r CO rX CO ■ CO I — I - - --J ■ ^ . ^ O O O' .^, .t-, .coico 'oa d d d d d CO -^ 00 Ol CO 05 , ,0 „-^t^ ^-^05 , — ,03 CD or— dt^d'^diod'o Tf ^ CD '*. t^ ^- CD "^ ■^oinOcqOcoo ■^ d lO ■^ in lo 00 CM ^ ^ OM _^ 03 _^ C» " lo ;:Z^ -^ O '^ C^ '^ -. ^ ^ CT ^. t- "^ lo ;i CO OcoOcoOlOOCDOCD (M ^ t^ ^ (>] "* 00 ^. 6 ^. CDOCOOCOOCOOCOO ■^ -^ T^ Tj* Tj^ d o o d d CO 05 05 o <; o til n (/3 ^ o 0( U ti Rbn C o 60 c hJ c« o ^ri 00 o> ID CO O t^ 00 Ol o ^ di d d d d d eg o r- CO 05 00 in ^^ o lo CO 1— I d d d d d d 00 CD in 00 CO CO 'S' ^ o •— I in CO d d d d d d O O O CD CM -rfi 00 CO t- -^ d d d d d d CO CO -^ r- in '—I CO -^ ^ CO (M m d d d d d d o t- o o o o ^ o o o o o d d d d d d CM O CO t- CM CO -'^ t- Tjf 00 t- Tt< CS 0.& ^1 o c -CS o u o 502 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Males ha\c three to five teeth on each process and a total of six to ten (mean, 8.1) prevo- merine teeth; females have three to si.\ teeth on each process and a total of 6 to 11 (mean, 8.5) prevomerine teeth. The vocal slits ex- tend from the midlateral base of the tongue to the angles of the jaws. The vocal sac is single, median, and subgular. The general coloration of Hijla eximia is bright green above, usually with dark brown spots or dashes, and a dark brown lateral .stripe (pi. 65, fig. 5; pi. 66, figs. 1 and .3). All individuals have a green dorsal ground color. This \'aries from a bright pale green to dark green or green with a tint of tan. A dark brown stripe begins on the snout and passes through the nostril, eye, and tympanum to extend onto the flank, as far as the groin in some specimens. This brown stripe is bor- dered above by a narrow white line. The pos- terior surfaces of the thighs are dull brown. The dorsal surfaces of the upper arms, thighs, shanks, and feet are marked by transverse dark brown bands or spots. The dorsal mark- ings are highly variable. Some indi\iduals lack dark markings on the dorsum, but in most there is some form of a dorsolateral series of dashes or a dorsolateral stripe. In addition to these marks, or in place of them, the dorsum in some individuals is marked by numerous small brown spots. The venter is creamy white, and the vocal sac in breeding males is dusty yellow with white flecks. The iris is dull bronze. Three aspects of the color pattern were analyzed in six samples from throughout the range of the .species (table 48). The lateral light stripe usually extends to the middle of the flanks or to the groin, but in specimens from the northern part of the range ( Chihua- hua) the stripe extends only to the axilla in 40 per cent in a sample of 42 specimens. On the other hand, in a series from Tamaulipas the lateral light stripe extends to the groin in 86 per cent of a sample of 4.3 specimens. In samples from the northern part of the range (Chihuahua, Durango, and Tamaulipas) the dorsal dark stripes are absent in more than 60 per cent of the specimens, whereas the stripes are solid in 42 per cent or fragmented in .39 per cent of the frogs in a sample of 72 specimens from Michoacan. Small spots are present on the dorsum in some specimens. Most individuals from the southern part of the range (Michoacan and Puebia) lack dor- sal spots, and few indi\iduals from there have spots all over the dorsum. However, in sam- ples from Durango, and Tamaulipas, small spots are present over most of the dorsum in more than three-fourths of the frogs exam- ined. Uniformly green frogs are present throughout the range; Holman (1965, p. .34) noted the uniformly green frogs in a sample from Durango and postulated a polymorphic gene in this species. In preservative, the dorsum is bluish gray. The anterior and posterior surfaces of the thighs, the groin, and the \entral surfaces of the limbs are creamy tan. The belly is creamy white, and the vocal sac in most breeding males is dark gray. The dorsal markings and the lateral stripe are dark brown. A distinct white anal stripe and labial stripe usually are evident. T.^DPOLES: Series of tadpoles are available from \ arious parts of the range of the species. Although some variation in pigmentation, par- ticularly on the tail, is evident, the tadpoles in the various samples are very nearly alike. Series of tadpoles ha\'e been examined from Arizona, Durango, Na\'arit, and Jalisco. A typical tadpole in developmental stage 37 from Buenos Aires, Durango, Mexico has a body length of 14.4 mm. and a total length of 32.1 mm. The body is deep; in dorsal pro- file the snout is bluntly rounded, and in lat- eral profile it is inclined anteroventrally from a point abo\c the nostrils. The eyes are rela- ti\ely small, widely separated, and directed laterally. The nostrils arc directed anterolat- erally at a point somewhat closer to the eyes than to the tip of the snout. The opening of the sinistral spiracle is directed posterodor- sally at a point on the midline about three- fifths of the distance from the tip of the snout to the posterior end of the body. The anal tube is long and dcxtral. The caudal muscu- lature is slender and tapers gradually to the tip of the acutely rounded tail. The fins are deep; at midlength of the tail the depth of the dorsal fin is slightly gi'eater than that of the ventral fin and is equal to about twice the depth of the caudal musculature. The dorsal fin extends onto the body (fig. 245C). 1970 DUELLMAN: HYLID FROGS 503 The dorsal and lateral svirfaces of the body are brown with minute sihei")' gold flecks. The venter is dark with an overlying tinge of pale gold. The caudal musculature is pale tan with dark brown flecks, especially con- centrated on the dorsal aspect of the pos- terior two-thirds of the caudal musculature. The fins are transparent with dark flecks and reticulations on all of the dorsal fin and on the posterior two-thirds of the ventral fin. In preservative, the gold tinge on the venter and the silvery gold flecks on the dorsum are lost. The mouth is small and directed antero- ventrally. Lateral folds in the lip are absent, and the median half of the upper lip is bare. The lips are bordered by two rows of small papillae; the beaks are rather massive and bear short serrations. The upper beak forms a broad arch with long, slender lateral pro- cesses. The lower beak is broadly V-shaped. There are two upper and three lower rows of teeth. The two upper rows are equal in length, and the second upper row is narrowly interrupted medially. The first and second lower rows are slightly shorter than the upper rows, and the first lower row is narrowly in- terrupted medially in some specimens. The third lower row is extremely short ( fig. 246C ) . Tadpoles from 40 kilometers northeast of Lagos de Moreno, Jalisco, Mexico, were col- ored like those from Buenos Aires, Durango, but were more pallid in appearance; they lack dark pigmentation on the caudal musculature and fins. Zweifel ( 1961 ) presented a detailed description of the development of the tadpoles from the northern part of the range; he dis- cussed these under the name of Hyla icrigh- tonim. Mating C.\ll: The mating call of Hijla eximia consists of a series of short, relatively low-pitched notes; in the calls of some indi- viduals these notes are distinct and separated, whereas in others they are so closely spaced that the call resembles a trill. No typical call can be described, because the variation in each of the parameters of the call seems to vary independently from the others (table 49). Some indi\iduals emit a slow call, and others have a fast call, whereas specimens from some areas emit an intermediate tvpe of call (pi. 13). Blair (1960) first pointed out the variation in the call of Htjia eximia; he noted the exis- TABLE 49 Geographic Variation in the Mating Calls of Hijla eximia. (Means are given in Parentheses.) Locality N Repetition Rate (min.) Duration Notes (sec.) Pulse Rate (per sec.) Frequencies (cps) Fundamental Dominant Apache County Arizona.. Buenos Aires, Durango. Tepic, Nayarit Lagos de Moreno, Jalisco Queretaro, Queretaro Huachinango, Puebla Patzcuaro, Michoacan Toluca, Mexico Sanctorium, Tlaxcala Ixtapan de la Sal, Mexico 3 42-84 0.16-0.18 110-120 109-122 1635-2078 (68) (0.167) (115) (118) (1889) 3 42-60 0.39-0.42 80-85 74-87 1740-1826 (51) (0.400) (82) (82) (1781) 1 58 0.20 60 87 1.560 7 90-1.37 0.15-0.19 110-130 90-131 20.34-2498 (119) (0.170) (120) (123) (2271) 1 156 0.15 100 83 2158 3 88-96 0.24-0.25 70-80 89-111 2112-2221 (92) (0.247) (75) (98) (2184) 1 92 0.23 95 100 2300 2 90 0.21-0.24 78-80 77-91 2233-2462 (0.225) (79) (84) (2297) 3 45-60 0.28-0.30 48-52 87-91 2349-2457 (54) (0.290) (50) (89) (2297) 0 40-57 0.15-0.24 48-70 81-113 2300-2704 (47) (0,200) (55) (103) (2501) 504 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 tence of slow-calling populations and of other fast-calling populations. He suggested that possibly two or more species were involved. Bogert (1960, 1. 296) stated: "Even with the limited information thus far obtained for a few representative populations of Hyla eximia ... it is possible to show that intermediate stages ranging from an un-trilled to trilled calls may occur within populations currently assigned, and correctly so in all probability, to a single species. The variation within these populations ... do not appear to be clinal in nature. On the contrary, the variations more closely resemble a mosaic pattern of dis- tribution, insofar as can be judged by repre- sentative calls from seven populations." The recordings that I have analyzed provide data in support of Bogert's suggestion. However, I have been able to obtain data from only .34 recordings from 10 different samples. Much more work needs to be done on this aspect of the biology of Hyla eximia. Natural History: Hyla eximia inhabits subhumid highlands, where it occurs in mes- quite-grassland, scrub forests, and pine-oak forest. The species is an opportunistic breed- er and utilizes shallow rainpools in undis- turbed as well as artificial situations. Calling males have been found as early as June 11 and as late as August 21 on the Mexican Pla- teau. Breeding usually takes place in the shallow grassy ponds. The males call from shallow water or while floating on the surface of the water usually grasping a blade of grass or a stick with the hands. The eggs are laid in loose clumps attached to grasses in shallow water. In December, 1959, adults were found secreted in bromeliads growing on pine trees near Tianguistengo, Hidalgo, Mexico. Hol- man (1965, p. 34) reported finding adults and many juveniles beneath rocks in pine forests at La Ciudad, Durango, Mexico, in March. The tadpoles develop in shallow grassy ponds, where they seek refuge amidst the aquatic vegetation. Tadpoles have been found as early as June 27 near Ixtapan de la Sal, Mexico, and as late as August 25 at Buenos Aires, Durango. Remarks: The synonymy of Hijla eximia has had a varied and confused historv. Kel- logg (1932, p. 164) included Hyla euphorbia- cea Giinther and Hyla smithii Boulenger in the synonymy of eximia, but he considered Hyla gracilipes Cope to be a separate species and having as a synonym Hyla lafrentzi Mer- tens and Wolterstorff. Giinther (1901, p. 261) first placed Hyla gracilipes in the synonymy of Hyla eximia. Taylor (1939b, p. 423) disa- greed with Kellogg's recognition of Hyla gra- cilipes and concluded, as did Giinther, that gracilipes is the same as eximia. I have ex- amined the syntypes of gracilipes (U.S.N.M. Nos. 15318-15321) and agreed with Taylor that these specimens are representatives of Hyla eximia. Taylor (1939b, p. 426) showed that Hyla euphorbiacea Giinther was a valid species, distinct from Hyla eximia. Hyla smithii long has been recognized as a distinct species not closely related to Hyla eximia. Maslin ( 1957, p. 81 ) named Hyla micro- eximia from 5 kilometers northwest of Jocote- pec, Jalisco, Mexico. Duellman (1961c, p. 49) discussed the variation in Hyla eximia. A comparison of the holotype of microeximia (U.S.N.M. No. 139246) with the syntypes of Hyla eximia (U.S.N.M. No. 3248) reveals that the holotype of microeximia is larger than either of the two syntypes of eximia! In the foregoing synonymy of Hijla exiinia I have included three other species for the first time; these are Hyla cardenasi Taylor, 19.39b; Hyla wrightorum Taylor, 1939b; and Hyla arboricola Taylor, 1941. Following is a justification of these assignments. The holo- type of Hyla cardenasi (U.S.N.M. No. 84403) from Puebla, Puebla, Mexico is a gra\'id fe- male having a snout-vent length of 39 mm. and essentially no dorsal dark markings what- soever. The comparison of this specimen with a series of Hyla eximia and plicata reveals that the detailed structure of the hands and feet, especially the amount of webbing on the feet is like that of eximia and not of plicata. Hyla eximia is a rather common frog in the vicinitN' of Puebla, and I conclude that name Hyla cardenasi was based on an unpatterned indi\idual of Hyla eximia; dorsal spots are absent in more than 50 per cent of the speci- mens from the vicinity of Puebla, and dorsal dark stripes are absent in more than 25 per cent of the specimens from that area. 1970 DUELLMAN: HYLID FROGS 505 Specimens of Htjla exiinia from the north- ern part of the range (Arizona and New Mex- ico in the United States, and Chihuahua in Mexico) are somewhat larger, more robust, and have proportionately longer legs than do those frogs from the southern part of the range. Ho\\'ever, these differences in size and proportions, notwithstanding, the variation in color patterns indicates a very close relation- ship between the northern and southern popu- lations. Analysis of mating calls of individuals formerly assigned to Hyla urightorum from Apache County, Arizona, with those from throughout the Mexican Plateau, reveal no outstanding differences. The fundamental frequency is slightly higher in those individ- uals from Apache County, Arizona, than in the other samples, but the range of variation in the former is included in the latter. On the basis of the absence of any distinctive morphological characters and on the basis of general similarity of mating call, I conclude that Hyla urigl^torum is the same as Hyla eximia. Taylor (1941, p. 118) diagnosed Hyla ar- horicola as different from eximia by having a broader head, more webbing on the feet, limbs lacking dark marks, and the absence of a well-defined dark mark on the side of the head. Few adult specimens have been ob- tained from the highlands of Guerrero, but of these, several have dark markings that are typical of Hyla eximia. Furthermore, the pro- portions of head width and the amount of webbing on the feet fall within the range of variation of Hyla eximia on the Mexican Pla- teau. Unfortunately, recordings of the calls of frogs of the populations in the highlands of Guerrero are not available. Thus, my con- clusion that Hyla arhoricola is a synonym of eximia is based solely on morphological evi- dence, without the benefit of a knowledge of the mating call or the tadpoles of the frogs formerly assigned to arhoricola. Etymology: The specific name is Latin meaning uncommon! Distribution': Hyla eximia occurs in a variety of upland environments but princi- pally associated with pine forests, in highland areas in central Arizona and New Mexico, in the Huachuca Mountains of southern Arizona, in the Sierra Madre Occidental in northwest- ern Mexico, and throughout the southern part of the Mexican Plateau, the Sierra Madre Ori- ental, and Cordillera Volcanica in central Mexico (fig. 250). The species occurs at de- lations between 900 and 2900 meters. See Appendix 1 for the locality records of the 2209 specimens examined. Hyla euphorbiacea Giinther Hyla euphorbiacea Giinther, 1859, p. 109 [syn- types, B.M.N.H. No. 1947.2.24.19 from "Cordilleras," Mexico; E. Parzudaki collector; B.M.N.H. Nos. 1947.2.24.15 and 16 from "Me-xico," B.M.N.H. No. 1947.2.24.18 from "Cordilleras," Mexico, and B.M.N.H. No. 1947.2.24.17 from Cordoba (Veracruz?), Mexico; Auguste Salle collector]. Taylor, 1939b, p. 426. Smith and Taylor, 1948, p. 82. Hijliola bocourti Mocquard, 1889b, p. 341 [syn- t>pes, M.N.H.N. Nos. 1266 (2 specimens), 6370 (6 specimens), 6371 (6 specimens) from Alta Verapaz, Guatemala; Marie-Firmin Bocourt collector]. Hyla bocourti: Gunther, 1901 (1885-1902), p. 263. Stuart, 1963, p. 35. Diagnosis: This moderately small green frog with a brown face mask and brown spots or stripes dorsally has smooth skin, small discs, no webbing between the fingers, and the toes about two-thirds webbed. The presence of small yellow spots on the dark brown pos- terior surfaces of the thighs immediately dis- tinguishes this species from other members of the eximia group. Other Middle American hylids with yellow spots on the posterior sur- faces of the thighs include Hyla pictipes and xanthosticta in Costa Rica; both of those frogs have relatively large discs and have webbing between the fingers. Description: Males of this moderately small species attain a maximum snout-vent length of 29.6 mm., and females reach 40.6 mm. In a series of 25 males from the Valley of Oaxaca, Oaxaca, Mexico, at an elevation of about 1500 meters, the snout-vent length is 31.6 to 37..3 (mean, 34.7) mm.; the ratio of tibia length to snout-vent length is 0.434 to 0.480 (mean, 0.457); the ratio of foot length to snout-vent length is 0.410 to 0.469 (mean, 0.440 ) ; the ratio of head length to snout-vent length is 0.270 to 0..304 (mean, 0.287); the ratio of head width to snout-vent length is 0.304 to 0.335 (mean, 0.320), and the ratio of the diameter of the tympanum to that of the eye is 0.548 to 0.733 (mean, 0.629). Five 506 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 110° 106° 102° 98° 1 ^ _ i • "n < stf "V • • y''\ V, /■ V. \ .' y Jl scr 26° " ( 26° S \ \ -' '- '- ^ ■■--- 22° 22° 18° 0 100 300 \" r- <^ r^^*i^ 18° ■■ KILOMETERS I 110° 106° 102° 98° Fig. 250. Distribution of Utjla cximia. 1970 DUELLMAN: HYLID FROGS 507 females from the same locality ha\'c snout- vent lengths of 34.0 to 39.6 (mean, 36.4) mm. and show no significant differences in pro- portions. In a series of 25 males from Llano de las Flores, Oaxaca, Me.xico, at an elevation of 3100 meters, the snout-vent length is 33.3 to 39.6 (mean, 36.6) mm. These slightly larger frogs do not differ from those from the Valley of Oa.xaca in proportions, except that the tym- panum is proportionately smaller; the ratio of the diameter of the tympanum to that of the eye is 0.472 to 0.658 (mean, 0.570). The head is slightly narrower than the body, and the top of the head is barely con- vex. In dorsal profile the snout is acutely rounded, and in lateral profile it is round. The snout is moderately long; the nostrils are barely protuberant at a point about two-thirds of the distance from the eyes to the tip of the snout. The canthus is rounded, and the loreal region is barely concave; the lips are thin and barely flared. A moderately thin dermal fold extends posteriorly from the eye, abo\e the tympanum, and downward to a point behind the angle of the jaw. The fold obscures the upper edge of the tympanum, which other- wise is distinct, and separated from the eye by a distance equal to about one-half the diameter of the tympanum. The arms are moderately long and slender; an axillary membrane is absent. An indistinct row of tubercles is present on the ventrolateral edge of the forearm, and a thin dermal fold is present on the wrist. The fingers are moder- ately long and slender and bear small discs; the width of the disc on the third finger is equal to about three-fifths of the diameter of the tympanum. The subarticular tubercles are large and conical; none is bifid. The super- numerary tubercles are conical and distinct on the proximal segments of the digits. An elevated, usually bifid, palmar tubercle is pres- ent. The prepollex is barely enlarged and in breeding males bears a thin horny nuptial ex- crescence. The webbing between the fingers is vestigial (fig. 243D). The hind limbs are moderately short and robust; the heels of the adpressed limbs overlap by about one-fifth of the length of the shank. The tibiotarsal articulation extends to the tympanum or to the posterior comer of the eye. A thin trans- verse dermal fold is present on the heel and a distinct tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is ovoid and elevated. A small conical outer metatarsal tubercle is present. The toes are long and slender and bear discs that are smaller than those on the fingers. The sub- articular tubercles are large and round, and the supernumerary tubercles are low, indis- tinct, and present only on the proximal seg- ments of the digits. The toes are slightly more than one-half webbed (fig. 244D). The webbing extends from the base of the penulti- mate phalanx of the first toe to the distal end of the antepenultimate phalanx of the second, from the base of the penultimate phalanx of the second to the base of the antepenultimate phalanx of the third, from the distal end of the antepenultimate phalanx of the third to the base of the antepenultimate phalanx of the fourth and on to the base of the penulti- mate phalanx of the fifth toe. The anal opening is directed posteriorly at the upper level of the thighs; it is covered by a short anal sheath. The skin on the throat, belly, and posteroventral surfaces of the thighs is strongly granular; elsewhere, the skin is smooth. The tongue is cordiform, mod- erately notched posteriorly, and free behind for about one-third of its length. The dentig- erous processes of the prevomers are small, posteromedially inclined elevations between the small, ovoid choanae. Males have two to fi\'e teeth on each process and a total of five to nine (mean, 7.8) prevomerine teeth; females have three to five teeth on each pro- cess and a total of six to ten (mean, 8.1) prevomerine teeth. The vocal slits extend from the midlateral base of the tongue to the angles of the jaws. The vocal sac is single, median, and subgular. The general coloration of Htjla euphorbia- cea is green or pale tan above with or without dark brown dorsal markings (pi. 66, fig. 6). The dorsum varies from pale green and olive- green to pale tan. Usually the dorsum is marked by elongate dark brown streaks or small round brown spots; in approximately 40 per cent of the specimens examined the dorsal markings are absent or reduced to a few small spots posteriorly. A dark brown stripe extends from the snout through the nostril, eye, and tympanum onto the flank, and 508 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 usually to the groin. This stripe is bordered above by a narrow white line. The anterior and posterior surfaces of the thighs, the outer edges of the shanks, and the inner edges of the tarsi are orange-brown to black with bright yellow spots. The belly is creamy white. The vocal sac is yellow or brown with creamy yellow spots anteriorly. The iris is pale coppery bronze. In preservative, the dorsum is pale bluish gray or grayish tan. Dorsal markings are brown. Individuals lacking brown marks on the back usually lack dark transverse marks on the hmbs. A distinct white anal stripe is invariably present, and a distinct white stripe on the outer edge of the shank usually is evi- dent. Pale spots are present on the posterior surfaces of the thighs in all specimens; how- ever, in some individuals the spots are absent in the groin and on the anterior surfaces of the thighs. Tadpoles: A typical tadpole in develop- mental stage .31 has a body length of 11.5 and a total length of 28.0 mm. The body is slightly deeper than wide; in dorsal profile the snout is bluntly rounded, and in lateral profile it is round. The eyes are small, widely separated, and directed laterally. The nostrils are directed anterolaterally at a point about midway between the eyes and the tip of the snout. The spiracle is directed posterodorsally at a point below the midline and about three- fifths the distance from the tip of the snout to the posterior end of the body. The anal tube is long and dextral. The caudal muscu- lature is slender and tapers to the tip of the acutely rounded tail. The caudal fins are deep; at midlength of the tail the depth of the dorsal fin is half again as great as the depth of the caudal musculature. The dorsal fin extends onto the body (fig. 245D). Tadpoles are pale tan above and pale golden below. The throat is dark gray with silvery flecks. The caudal musculature is tan with faint grayish brown reticulations on the musculature and fins. In preservative, the body and caudal musculature is creamy tan; faint gray flecks and reticulations are evident on the musculature and fins. The mouth is small and directed antero- ventrally. Lateral folds are absent in the lips. The median one-third of the upper lip is bare, whereas the rest of the lips are bordered by two or three rows of small papillae. The beaks are moderately robust and bear small serrations. The upper beak is in the form of a broad arch with slender lateral processes, and the lower beak is broadly V-shaped. There are two upper and three lower rows of teeth. The upper rows are equal in length and the second upper row is broadly inter- rupted medially. The first and second lower rows are nearly as long as the upper rows, and the third lower row is noticeablv shorter (fig. 246D). Mating Call: The call of Hijla eitphor- biacea consists of a short series of quickly repeated, low-pitched notes. An analysis of the calls of 12 individuals recorded at temper- atures of 17° to 18°C. from the Valley of Oaxaca show that there are five to ten ( mean, 7.4) notes per call group. The note repetition rate is 600 to 900 (mean, 773) notes per min- ute, and the duration of each note is 0.03 to 0.06 (mean, 0.05) of a second. The call rate is 18 to 39 (mean, 25.5) call groups per min- ute, and the pulse rate is 100 to 120 (mean, 112) pulses per second. The fundamental frequency varies from 104 to 130 (mean, 114.3) cycles per second and the dominant frequency varies from 2080 to 2736 (mean, 2345.8) cycles per second (pi. 14, fig. 1). Calls of Hyla euphorbiacea have been re- corded at temperatures between 12.0°C. and 21.5°C. Analysis of these records indicate that there is little variation in the number of notes per call group, but the note repetition rate, call rate, pulse rate, and the fundamental and dominant frequencies increase at higher tem- peratures, whereas the duration of the notes decreases at higher temperatures (table 50). Natural History: Hijla euphorbiacea is especially ubiquitous in the Valley of Oaxaca, where the frogs call by the thousands from flooded grassy fields after heavy rains in July and August. The frogs also occur at high elevations in pine-oak and pine forest, where they breed in shallow temporary ponds. Males call while sitting in shallow water or while floating on the water and holding onto blades of grass or small sticks. The eggs are laid in loose clumps in grassy parts of the pond. Tad- poles have been found in shallow grassy ponds, a shallow muddy pool in oak forest on 1970 DUELLMAN: HYLID FROGS TABLE 50 Comparison of Certain Parameters of the Mating Calls of Hyla eupJwrbiacea at Different Temperatures (Means are given in Parentheses.) Parameter 12.5°C 17-18°C JV 5 12 12-18 18-39 Call Rate ( min. ) -. ( 15.0 ) ( 25.5 ) 300-466 600-900 Note Repetition Rate (min.) (368) (773) 65-90 100-120 Pulse Rate (sec.) (72) (112) 70-87 104-130 Fundamental Frequency (cps) (81.8) (114.3) 1653-2175 2080-2736 Dominant Frequency (cps) (1793) (2346) 0.08-0.11 0.03-0.06 Durationof Notes (sec.) (0.098) (0.050) 509 21.5°C 6 28-36 (31.3) 686-935 (820) 100-120 (115) 104-139 (118.7) 2300-2782 (2518) 0.04-0.05 (0.047) Cerro Machin, and in roadside ditches be- tween June 23 and August 31. In the dry season frogs of this species seek shelter in bromeliads; adults have been taken from bromeliads at Cumbres de Acultzingo, Veracruz, in January and at Llano de las Flores, Oaxaca, in March. Remarks: The status of HyJa bocourti (Mocquard) is doubtful. The only specimens that have been referred to this species are from the vicinity of Coban on the Atlantic slopes of the Guatemalan highlands. Three subadults (F.M.N.H. Nos. 20684-20686) have snout-vent lengths of 26.9, 29.8, and 30.6 mm., respectively; an adult female (U.M.M.Z. No. 90870) has a snout- vent length of .39.8 mm. The latter is partially dried, brittle, and for- malin burned. This specimen was obtained from a bromeliad at Finca Samac, Alta Vera- paz, Guatemala, by Laurence C. Stuart on April 26, 1938. In his field notes, Stuart stated: "Above light brown with slightly darker brown longitudinal streaks — trace of similar colored broad band between eyes — a distinct dark brown streak from nostril to eye and along sides where it widens — sharply demarked above but indistinct below — belly brown mottled with gray — posteriorly and somewhat laterally bright yellow spotted with brown — legs brown above with several darker bars — below light yellow with brown mot- thng." The three subadults from Coban all have dark thighs with pale spots. I am con- vinced that on the basis of morphology and coloration, HyJa bocourti cannot be distin- guished from Hyla eiiphorbiacea. The tad- poles and mating calls of the Guatemalan pop- ulation herein referred to as eiiphorbiacea are unknown; consequently, the possibility does exist that there are biological differences be- tween the two populations. However, at the present time on the basis of the existing knowledge, it seems best to me to consider the Guatemalan and the Oaxacan specimens as examples of one species. Of course, this poses a zoogeographic problem. Hyla eiiphor- biacea is known from elevations in excess of 1500 meters in Oaxaca and at elevations of about 1000 meters on the northern slopes of the highlands in Guatemala. Intervening be- tween the ranges of these two populations are the lowlands of the Isthmus of Tehuan- tepec and the highlands of Chiapas and Gua- temala, which are inhabited by Hyla icalkeri, a species obviously closely related to, but distinct from, Hyla eximia. Etymology: The specific name is derived from the Latin euphorbea, referring to plants of the family Euphorbiacea and the Latin suffix -aceus, meaning belonging to. Distribution: Hyla euphorbiacea occurs in the Sierra Madre Oriental southward from central Veracruz into Oaxaca, in the Valley of Oaxaca, and in the mountains to the south of 510 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 • H euphorbiacea o H. wolkeri 0 50 ^ ^ KILOMETERS 98° 94° Fic. 251. Distribution of Hxjla euphorbiacea and Hyla tcalkeri. the Valley of Oaxaca; in addition, the species is known from the Atlantic slopes of the high- lands in Alta Verapaz, Guatemala (fig. 251). In Mexico, the species is known from eleva- tions between 1600 and 3150 meters, and in Guatemala it is known from elevations of about 1000 meters. See Appendix 1 for the locality records of the 810 specimens examined. Hyla walkeri Stuart Hyla walkeri Stuart, 1954b, p. 165 [holotype, U.M.M.Z. No. 106817 from Aserradero San Lorenzo ( 12 kilometers airline, slightly east of north of Jalapa), Jalapa, Guatemala; Laurence C. Stuart collector]; 1963, p. 37. Hijla euphorbiacea biscriata Lyncli, in Smith, Langebartel, and Williams, 1964, p. 24 [nomcn nudutn]. Diagnosis: This moderately small species having a green dorsum with a dark brown face mask and usually with a pair of brown lines posteriorly on the dorsum has smooth skin, small discs, no webbing between the fingers, and the toes about two-thirds webbed. The presence of uniformly tan posterior sur- faces of the thighs immediately distinguishes icalkeri from euphorbiacea, which has yellow spots on the thighs. Hijla regiUa differs by having a dark interorbital triangular mark, and cadaverina differs by being brown or gray and having tuberculate skin. Most speci- mens of eximia have transverse bars on the thighs (usually absent in icalkeri) and have brown spots and /or more extensive stripes on the dorsum. Hyla plicata is larger (males to 44 mm.) than icalkeri (36 mm.) and has slightly more webbing on the feet; the web extends to the base of the penultimate phalanx of the fifth toe in plicata and only to the distal end of the antepenultimate phalanx in icalk- eri). Description: Males of this moderately small species attain a maximum snout-vent length of 35.9 mm., and females reach 37.8 mm. In a series of 20 males from IS kilometers northwest of Comitan, Chiapas, Mexico, the snout-\ent length is 29.0 to 35.6 (mean, 32.0) 1970 DUELLMAN: HYLID FROGS 511 mm.; the ratio of tibia length to snout-vent length is 0.463 to 0.517 (mean, 0.491); the ratio of foot length to snout-vent length is 0.445 to 0.495 (mean, 0.475); the ratio of head length to snout-vent length is 0.294 to 0..321 (mean, 0.305); the ratio of head width to snout-vent length is 0.308 to 0.361 (mean, 0.332), and the ratio of the diameter of the tympanum to that of the eye is 0.469 to 0.633 (mean, 0.553). Three females from the same localit\- have snout-vent lengths of 30.8 to 32.6 (mean, 31.6) mm., and do not differ from the males significantly in proportions. Specimens from Guatemala exhibit the same range in measurements and proportions, ex- cept that they have slightly smaller tympani. In a series of 40 males from Soloma and San Juan Ixcoy, Departamento Huehuetenango, Guatemala, the ratio of the diameter of the tympanum to that of the eye is 0.444 to 0.552 (mean, 0.497). The head is slightly narrower than the body, and the top of the head is barely con- vex. In dorsal profile the snout is acutely rounded, and in lateral profile it is round. The snout is moderately long, and the slightly protuberant nostrils are situated at a point about two-thirds of the distance from the eyes to the tip of the snout. The canthus is round- ed, and the loreal region is barely concave; the hps are thick and barely flared. A thin dermal fold extends posteriorly from the eye, above the tympanum, and diffuses onto the body above the insertion of the arm. The fold covers the upper edge of the tympanum, and the rest of the tympanic ring is barely discernible. The tympanum is separated from the eye by a distance about two-thirds of the diameter of the tympanum. The arms are moderately long and slender; an axillary membrane is absent. A row of low, indistinct tubercles is present on the ven- trolateral edge of the forearm, and a weak transverse dermal fold is present on the wrist. The fingers are moderately long and slender and bear small discs; the width of the disc on the third finger is equal to about two-thirds of the diameter of the tympanum. The sub- articular tubercles are moderately large and round; none is bifid. The supernumerary tu- bercles are small, indistinct, and present only on the proximal segments of the digits. A small, round palmar tubercle is present. The prepollex is barely enlarged, and in most breeding males does not bear a nuptial ex- crescence. The webbing on the hand is ves- tigial (fig. 243E). The hind limbs are mod- erately long and slender; the heels of the ad- pressed limbs overlap by about one-fourth of the length of the shank. The tibiotarsal articulation extends to the eye. A thin trans- \erse dermal fold is present on the heel, and a distinct tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is small, ovoid, and barely elevated. The outer metatarsal tubercle, if present, is small and subconical. The toes are long and slender and bear discs that are slightly smaller than those on the fingers. The subarticular tuber- cles are moderately small and round, and the supernumerary tubercles are low, indistinct, and present only on the proximal segments of the digits. The toes are about one-half webbed (fig. 244E). The webbing extends from the base of the penultimate phalanx of the first toe to the distal end of the antepen- ultimate phalanx of the second to the base of the antepenultimate phalanx of the third, from the base of the penultimate phalanx of the third to the base of the antepenultimate pha- lanx of the fourth and on to the base of the penultimate phalanx of the fifth toe. The anal opening is directed posteriorly at the upper level of the thighs, and a short anal sheath is evident. The skin on the throat, belly, and proximal posteroventral surfaces of the thighs is granular; elsewhere, the skin is smooth. The tongue is cordiform, shallowly notched behind, and free posteriorly for about one-third of its length. The dentigerous pro- cesses of the prevomers are small posterome- dially inclined elevations between the pos- terior margins of the small, ovoid choanae. Males have three to six teeth on each process and a total of six to 11 (mean, 8.2) prevo- merine teeth; females have three to six teeth on each process and a total of six to 12 ( mean, 8.7) prevomerine teeth. The vocal slits ex- tend from the midlateral base of the tongue to the angles of the jaws. The vocal sac is single, median, and subgular. The general coloration of Hyla ualkeri is bright green or greenish tan above with or without dark brown longitudinal markings 512 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 (pi. 66, figs. 4 and 5). Generally the dorsum is green. There is a dark brown line e.xtend- ing from the snout through the nostril and eye to the midflank or groin; this brown stripe is bordered above by a narrow white line. The dorsum is marked by a dorsolateral brown stripe posteriorly, a row of dorsolateral brown spots, or no markings whatsoever. The dorsal surfaces of the thighs usually are uniform green, but in a few specimens brown flecks or small spots are present. The upper lip is flesh- colored on the margin and separated by a narrow brown line from the yellowish green color from the nostril to the angle of the jaw. The posterior surfaces of the thighs are dull yellowish brown. The venter is pale creamy yellow and vocal sac in breeding males is brown anteriorly and yellow posteriorly. The iris is dull bronze with fine black reticulations. In preservative, the dorsal ground color is pale bluish gray. The dorsal markings arc brown; in some individuals these are narrow- ly outlined with white. The edge of the upper lip, the upper border of the lateral brown stripe, the outer edge of the shank, and the stripe above the anus are white. The posterior surfaces of the thighs are creamy tan, and the venter is creamy white. Tadpoles: A typical tadpole in develop- mental stage 37 has a body length of 16.0 mm. and a total length of 37.2 mm. The body is deep, slightly deeper than wide; the snout in dorsal profile is bluntly rounded, and in lat- eral profile it is round. The eyes are small, widely separated, and directed laterally. The nostrils are directed anterolaterally at a point about midway between the eyes and the tip of the snout. The opening of the sinistral spiracle is directed posterodorsally at a point below the midline about two-thirds of the distance from the tip of the snout to the pos- terior edge of the body. The anal tube is long and dextral. The caudal musculature is mod- erately slender and tapers gradually to the tip of the acutely pointed tail. The caudal fins are moderately deep; at midlcngth of the tail, the depth of the dorsal fin is slightly greater than the depth of the caudal muscu- lature. The dorsal fin extends onto the bod\- (fig. 245E). The tadpoles arc dark brown above with a silvery iridescence on the venter. The cau- dal musculature is creamy tan with dark brown or grayish brown mottling and reticula- tions on the musculature and fins. In pre- servative, the body is dark brown; the caudal musculature is tan and there is a heavy con- centration of dark pigment on the dorsal as- pects of the musculature. The mouth is moderately small and di- rected anteroventrally. Lateral folds are ab- sent from the lips. The median half of the upper lip is bare; the rest of the lip is bor- dered by two rows of small papillae. The beaks are massive and bear short, blunt serra- tions. The upper beak is in the form of a broad arch with long, slender lateral pro- cesses; the lower beak is broadly V-shaped. There are two upper and three lower rows of teeth. The upper rows are long, and the second upper row is broadly interrupted me- dially. The first and second lower rows are about equal in length, but much shorter than the upper rows. The first lower row is nar- rowly interrupted medially in some specimens. The first lower row is noticeably shorter (fig. 246E). Mating Call: The call of Hyla icalkeri consists of groups of four to si.\ short, quickly repeated, low-pitched notes. The call rate varies from 30 to 48 (mean, 38.0) call groups per minute, and the note repetition rate varies from 960 to 1200 (mean, 1090) notes per minute. The duration of the note varies from 0.03 to 0.04 (mean, 0.035) of a second, and the pulse rate is about 120 pulses per second. The fundamental frequency varies from 135 to 184 (mean, 158) cycles per second, and the dominant frequency varies from 1755 to 2175 (mean, 1910) cycles per second (pi. 14, fig. 2). Natural History: Hyla walkeri inhabits pine-fir and pine-oak forests. In the rainy season, males call from temporary grassy ponds, frequently in clearing or meadows. The males call while floating on the water with their hind limbs partly fle.xed or with the hands grasping grass or debris. Some indi- viduals were observed sitting in shallow wa- ter near the shore. All clasping pairs were observed floating in the water. Stuart ( 1954b, p. 168) reported this species calling on June 17-18, 1952, at San Lorenzo, Departamento Jalapa, Guatemala. In June, 1960, I obtained 1970 DUELLMAN: HYLID FROGS 513 calling males from 10 to IS kilometers north- west of Comitan, Chiapas, Mexico. Porter ( 1962, p. 168 ) reported males calling from a meadow at San Cristobal de las Casas, Chia- pas, Mexico, on June 15, 1960. At the same locality, I found adults in rotting pine logs on February 17, 1961. Tadpoles were obtained from grassy ponds at 10 kilometers northwest of Comitan, Chia- pas, on June 17, 1960; tadpoles and metamor- phosing young were obtained at a grassy pond at 2.5 kilometers south of Jitotal, Chiapas on August 5, 1960. Remarks: In most features of its mor- phology, Hyla walkeri is indistinguishable from Hyla eximia and euphorhiacea. In col- oration, it differs from both of these by gen- erally lacking transverse marks on the dorsal surfaces of the thighs. From euphorhiacea it differs by lacking the yellow spots in the groin and on the anterior and posterior sur- faces of the thighs. The calls of Hyla walkeri and euphorhiacea are ahke in consisting of short groups of notes; in this respect, the calls of both of these species differ from eximia, the call of which consists of individual notes not grouped together. Etymology: The specific name is a patro- nym for Charles F. Walker. Distribution: Hyla icalkeri occurs in the central highlands of Chiapas, Mexico, the Sierra de los Cuchumatanes in western Gua- temala, and on the plateaus of central Guate- mala, and in the highlands of southeastern Guatemala (fig. 251). The species is known from elevations between 1450 and 2340 me- ters. See Appendix 1 for the locality records of the 194 specimens examined. The Hyla versicolor Group DEFiNrriON: The frogs in this group are medium-sized species having a tan, gray, or green dorsum with darker irregular blotches or spots; the limbs are barred. The venter is white; the vocal sac is gray, and the palpebral membrane is clear. The flanks and anterior and posterior surfaces of the thighs are uni- formly pale or mottled with black or dark brown. The dorsum is moderately rugose. The fingers are barely webbed, and the feet are one-half to two-thirds webbed. A strong tarsal fold is present, but an axillary mem- brane and dermal appendages on the limbs are absent. Males have a single, median, subgular vocal sac and horny nuptial excres- cences on the prepollices. The skull is only moderately ossified, and a large frontoparietal fontanelle is present (fig. 252). The sphen- ethmoid is not ossified anteriorly, and the nasals are large and not, or barely, separated medially. The squamosal is not in bony con- tact with the crista parotica, and the anterior arm of the squamosal extends no more than half of the distance to the maxillary. The columella is moderately expanded distally. A quadratojugal is present and articulates with the maxillary. The prevomers are moderately well ossified and bear teeth. The medial ramus of the pterygoid does not articulate with the prootic. The tadpoles have moder- ateh' deep fins and anteroventral mouths with two upper and three lower rows of teeth. The mating calls consist of a rattling series of short notes or distinct short, pulsed notes. The haploid number of chromosomes is 12. Composition*: Five species {arenicolor, avivoca, chrysoscelis, femoralis, and versi- color) comprise the group, which is wide- spread in North America east of the Sierra Nevada and the Colorado Desert. Only Hyla arenicolor occurs in Mexico; of that species, 599 preserved frogs from Mexico have been examined. Three lots of tadpoles and eleven skeletons from the United States have been examined. CoMMEXTS: Blair ("1958" [1959]) defined Fig, 252. Dorsal \iew of the skull of Hyla arenicolor, K.U. No. 44441. X 4. 514 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 the versicolor group on the basis of call struc- ture to include arenicolor, phaeocnjpta ( = avivoca), femoralis, versicolor, and baudinii. Starrett (1960b) and Duellman and Trueb (1966) showed that haudinii belonged in Smilisca. The group was modified by Johnson ( 1966) who showed that ''versicolor" was com- prised of two cryptic species, chrysoscelis and versicolor, differing from one another by mat- ing calls and a high degree of hybrid invia- bility. Members of the versicolor group arc very much alike in size, structure, and coloration, except femoralis, which is smaller than the other species and has relatively smooth skin and bold markings on the anterior and pos- terior surfaces of the thighs. According to Blair ("1958" [1959]) the call of femoralis is most like that of arenicolor, the only species occurring in Middle America. On the basis of structure of the adults and tadpoles and on the nature of the mating calls the Hijla versicolor group seems to be most closely related to the Hyla cinerea group of southeastern North America. The versi- color group apparently is not closely related to any of the groups endemic to Mexico and Central America. Hyla arenicolor Cope Hyla affinis Baird, 1854, p. 61 [syntypes, U.S.N.M. No. 11410 (originally 3261) (five specimens) from "northern Sonora," Mexico ( type locality restricted to Santa Rita Mountains, Arizona, by Smith and Taylor (1950, p. 354) and further restricted to Pefia Blanca Springs, 10 miles northwest of Nogales, Santa Cruz County, Arizona, by Gorman (1960, p. 218), who designated U.S.N.M. No. 11410a as the lectotype); John H. Clark collector; preoccupied by Hyla affinis Spi.x, 1824, from Brasil]. Brocchi, 1881, p. 43. Hyla arenicolor Cope, 1886a, p. 84 [replacement name for Hyla affinis Baird, 1854, preoccupied by Hyla affinis Spix, 1824]. Boulenger, 1882a, p. 373. Hyla copii Boulenger, 1887, p. 53 [syntypes, B.M.N.H. Nos. 1947.2.23.26 and 27 from El Paso, Texas, U.S..\.; Alphonso Forrer collector]. Giinther, 1901 (1885-1902), p. 266. Hyla coper Cope, 1888, p. 80 [typographical error for copii]. Hyliola digueti Mocquard, 1889a, p. 165 [syntypes, M. N.H.N. No. 492 (five specimens) from Territory of Tepic, Mexico ( restricted to Tepic, Nayarit, Me.xico, by Smith and Taylor, 1950); Leon Diguet collector]. Hyla arenicolor (part): Kellogg, 1932, p. 156. Smith and Taylor, 1948, p. 89. Di.\r,NOsis: This medium-sized species has tuberculate skin, vestigial webbing on the hands, and a dull gray or brown dorsum marked with irregular darker spots or blotch- es. The posterior surfaces of the thighs are dull yellow or tan, and numerous white flecks arc present in the anal region. Hyla arenicolor resembles cadaverina, which differs by having slightly more webbing, smaller discs, fewer and less distinct supernumerary tubercles, and smaller tympanum; the diameter of the tym- panum is about half of the diameter of the eye is cadaverina and about two-thirds of the diameter of the eye in arenicolor. Other Mid- dle American hylids that might be confused with arenicolor all have webbing between the fingers. DEScmPTiON: Males of this medium-sized species attain a maximum known snout-vent length of 51.2 mm., and females reach 57.1 mm. In a series of 22 males from the vicinity of Guadalajara, Jalisco, Mexico, the snout-vent length is 32.8 to 39.5 (mean, 35.5) mm.; the ratio of tibia length to snout-vent length is 0.454 to 0.518 ( mean, 0.489 ) ; the ratio of foot length to snout-vent length is 0.385 to 0.442 (mean, 0.413); the ratio of head length to snout- vent length is 0.301 to 0.360 (mean, 0.327); the ratio of head width to snout- vent length is 0.354 to 0.398 (mean, 0.377), and the ratio of the diameter of the tympanum to that of the eye is 0.543 to 0.730 (mean, 0.652). Three females from the same area have snout- \ent lengths of 39.5 to 44.4 (mean, 41.6) mm. The size of the specimens from the vicinity of Guadalajara is typical over most of the Mexi- can Plateau. Duellman (1961, p. 46) noted that specimens from higher ele\ations in Michoacan were smaller than those from lower elevations; seven males from elevations above 1400 meters have snout-\ent lengths of 32.3 to 38.4 (mean, 34.7) mm., whereas nine males from elevations below 1400 meters have snout-vent lengths of 44.7 to 51.2 (mean, 49.1) mm. Thirteen males from the vicinity of Chilpancingo, Guerrero, have snout-vent lengths of 44.7 to 48.9 (mean, 45.7) mm. In northern Mexico, the frogs are somewhat larger than they are on the plateau in the southern part of the range; for example, the largest of 36 males from Chihuahua has a snout-vent length of 42.8 mm. It seems as 1970 DUELLMAN: HYLID FROGS 515 though there is a general trend from north to south on the Mexican Plateau for a decrease in size, but that individuals from lower ele- vations in the southern part of the range are by far the largest of the species. There ap- pears to be no significant variation in propor- tions. The head is as wide as the body, and the top of the head is barely convex. In dorsal profile the snout is acutely rounded, and in lateral profile it is bluntly rounded. The snout is short, and the nostrils are barely pro- tuberant at a point about three-fourths of the distance from the eyes to the tip of the snout. The canthus is rounded, and the loreal region is barely concave; the lips are moderately thick and barely flared. A thin dermal fold extends posteriorly from the eye, above the tympanum, and downward to a point just posterior to the angles of the jaw. In some individuals the fold obscures the upper edge of the tympanum, but in most specimens, the tympanum is entirely distinct, situated pos- teroventral to the eye, and separated from the eye by a distance equal to about half of the diameter of the tympanum. The arms are moderately long and slender; an axillary membrane is absent. A row of dis- tinct or partially fused tubercles is present on the ventrolateral edge of the forearm and a weak transverse dermal fold is present on the wrist. The fingers are moderately long and slender and bear small discs; the width of the disc on the third finger is equal to about half of the diameter of the tympanum. The sub- articular tubercles are large and round; the distal tubercle on the fourth finger usually is bifid. Moderately large supernumerary tu- bercles and a large elliptical palmar tubercle are present. The prepollex is moderately en- larged and in breeding males lacks a horny nuptial excrescence. The webbing is vestigial (fig. 253A). The hind limbs are moderately short and robust; the heels of the adprcssed limbs barely overlap. The tibiotarsal articu- lation extends to a point between the eye and nostril. A thin transverse dermal fold is pres- ent on the heel, and an elevated, flap-like tarsal fold extends the full length of the tar- sus. Numerous small tubercles are present on the plantar surface of the tarsus. The inner metatarsal tubercle is moderately large, ele- \ated, and elliptical. A conical outer meta- tarsal tubercle is present. The toes are mod- erately long and slender and bear discs that are about equal in size to those on the fingers. The toes are about one-half webbed (fig. 253B ) . The webbing extends from the base of the penultimate phalanx of the first toe to the distal end of the antepenultimate phalanx of the second, from the middle of the penulti- mate phalanx of the second to the base of the antepenultimate phalanx of the third, from the middle of the penultimate phalanx of the third to the base of the antepenultimate pha- lanx of the fourth and on to the middle of the penultimate phalanx of the fifth toe. The anal opening is directed posteriorly near the upper level of the thighs. The skin on the dorsum is moderately tuberculate; small tubercles are present on the dorsal sur- faces of the hmbs. The skin on the throat, belly, and proximal posteroventral surfaces of the thighs is granular; elsewhere, on the venter, the skin is smooth. The tongue is nar- rowly cordiform, shallowly notched posterior- ly, and barely free behind. The dentigerous processes of the prevomers are short, postero- medially inclined, narrowly separated medial- ly elevations between the moderately small, ovoid choanae. Males have four to six teeth on each process and a total of nine to 12 (mean, 10.4) prevomerine teeth; females have five to seven teeth on each process and a total of 10 to 13 (mean, 11.4) prevomerine teeth. The vocal slits extend a short distance posterolaterally from the midlateral base of the tongue. The vocal sac is single, median, subgular, and moderately distensible. The general coloration of Hyla arenicolor is dull grayish brown with darker brown or gray spots ( pi. 64, fig. 2 ) . The typical colora- tion of an individual from Agua del Obispo, Guerrero, Mexico, in life is grayish brown above with dark brown spots and faint trans- verse bands on the limbs. The groin, anterior and posterior surfaces of the thighs, and ven- tral surfaces of the hind limbs is orange-yel- low. The belly is white, and the vocal sac is purplish brown. The eye is grayish copper. Individuals found by day frequently are quite pallid by comparison with those found at night. For example, an individual found in 516 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fig. 253. Hand (A) and foot (B) of Htjla arenicolor, K.U. No. 86935. x 4. a shady ravine at Chinapa, Michoacan, Me.x- ico, was pale ashy gray (pi. 64, fig. 3). In preservative, the dorsum varies from tan to dull brown or gray. Numerous spots or small blotches, frequently outlined with black flecks are evident on the dorsum in most individuals. In some specimens, the dark spots are present on the flanks, but in most individuals the flanks are dull gray or brown with minute white flecks. Two to four trans- verse bands are present on the thighs and shank, and usually two transverse bands are present on the forearm. The posterior sur- faces of the thighs are faintly mottled in some individuals from the Mexican Plateau, and in all specimens numerous white flecks are pres- ent in the anal region. The throat in breeding males is gray, brown, or black, frequently marked by small white flecks. The tliroat in some females is faintly spotted with brown. Tadpoles: Zweifel (1961) thoroughly de- scribed the tadpoles of this species from the Chiricahua Mountains, Cochise County, Ari- zona. I am unaware of any tadpoles of Hyh arenicolor from Mexico; consequently, the following description is based on Zweifel's material from Arizona. A typical tadpole in developmental stage 37 has a bod>' length of 12.5 mm. and a total length of 31.9 mm. The body is ovoid, no wider than deep. The dorsal profile of the snout is bluntly rounded, and in lateral pro- file the snout slopes gradually from the nos- trils to the tip. The eyes are large, widely separated, and directed laterally. The nostrils are directed anterolaterally at a point about midway between the eyes and the tip of the snout. The opening of the sinistral spiracle is at a point below the midline at about mid- length of the body. The anal tube is short and dextral. The caudal musculature is mod- erately robust and extends nearly to the tip of the bluntly pointed tail. The caudal fins are moderately deep. At midlength of the tail, the depth of the dorsal fin is approxi- mately equal to the depth of the caudal mus- culature. The dorsal fin does not extend onto the body (fig. 254). 1970 DUELLMAN: HYLID FROGS 517 Fig. 254. Tadpole of Hyla arcnicolor, A.M.N.H. No. 64678. X 4. The mouth is of medium size and in a ventral position. Distinct lateral folds are absent. The median part of the upper lip is bare; the rest of the upper lip is bordered by a single row of small papillae, and the lower lip is fringed by two rows of papillae. Additional small papillae arc present laterally in the mouth. The beaks are robust and bear short blunt serrations; the upper beak is in the form of a massive arch with short slender lateral processes, and the lower beak is broad- ly V-shaped. There are two upper and three lower rows of teeth. The upper rows are equal in length and e.xtend to the margins of the lips, whereas the three lower rows are equal in length, but noticeably shorter than the upper rows. The second upper row is narrowly interrupted medially in all speci- mens, and the first upper and first lower rows are narrowly interrupted in some individuals (fig. 255). ■ Zweifel (1961, p. 11) described the col- oration of large tadpoles (stages 36 and 41) as being golden brown with no pattern evi- dent on the body but a patchy distribution of superficial .\anthophores on the caudal muscu- lature, which gives a mottled appearance to Fig. 255. Mouth of tadpole of Hyla arenicoloi A.M.N.H. No. 64678. X 20. the tail. He stated that the ventral surfaces are "dense silver (with a golden sheen when seen at the right angle) which gives way abruptly to a golden brown of the dorsum about midway up the side of the body." In preservative, the dorsal part of the body is dull brown and the venter is transparent. The caudal musculature is creamy tan with a concentration of dark brown pigment on the dorsal edge of the musculature and form- ing faint spots posteriorly. The fins are trans- parent and marked by fine black flecks or reticulations. Mating Call: The call of Hyla arenicolor consists of a series of short, nasal notes "ah- ah-ah-ah." Some individuals call constantly for two or more minutes. An analysis of the calls of four individuals from the Chiricahua Mountains, Cochise County, Arizona, reveals that the note repetition rate varies from 30 to 38 (mean, 33) notes per minute; the dura- tion of the notes is 0.56 to 0.80 (mean, 0.65) of a second and the pulse rate is 28 to 33 (mean, 29) pulses per second. The funda- mental frequency varies from 94 to 113 (mean, 102) cycles per second and the domi- nant frequency varies from 2112 to 2460 (mean, 2329) cycles per second (pi. 12, fig. 3). An analysis of seven recordings obtained in the Peloncillo Mountains, New Mexico, shows that the frogs there differ slightly by having a slower repetition rate, shorter dura- tion of notes, and a slower pulse rate. The frogs produced 26 to 32 (mean, 27) notes per minute having a duration of 0.53 to 0.75 (mean, 0.64) of a second and 22 to 25 (mean, 24) pulses per second. Two individuals re- corded at Amayuca, Morelos, Mexico, pro- duced 64 and 68 notes per minute, and the duration of the notes in each was approxi- mately 0.50 of a second. Natural History: Hyla arenicolor inhab- 518 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 its a variety of vegetational formations. It occurs over a large part of the Mexican Pla- teau, where it is found in mesquite-grassland and scrub forest. In the mountains rising from and bordering the Mexican Plateau, this spe- cies occurs in pine and oak forest, and on the lower slopes of the highlands the frog oc- curs in scrub oak and dense thorn forest. However, throughout its range it is always closely associated with small rocky streams; Hyla arenicolor inhabits ravines and canyons. Males call from rocks or occasionally low bushes surrounding quiet pools in canyons. A few males have been observed to call from shallow water in the pools. CaUing males have been found from June 25 to Juh' 20 in the vicinity of Barranca del Cobre, Chihua- hua, Mexico. Calling males have been ob- tained on the Mexican Plateau in the vicinity of Guadalajara, Jalisco, between May 25 and July 10, at Lombardia, Michoacan, on July 12, and at Agua del Obispo, Guerrero, on June 19. Zweifel (1961, p. 16) noted the presence of tadpoles in the South Fork of Cave Creek in the Chiricahua Mountains, Cochise Coun- ty, Arizona, on June 22, 1958 and surmised that breeding must have commenced at least a month earlier. He also found that the spe- cies bred at that locality in July. Zweifel (1961, p. 17) reported egg laying in pools in a canyon on July 12 or 1.3, 1960 and noted that the period from oviposition to metamorphosis was probably between .50 and 60 days. He gave the snout-vent length of newly metamorphosed young as about 15 mm. Remarks: Kellogg (1932, p. 156) pro- vided a thorough discussion of the synonymy of this species. Most references to Hijla areni- color in Mexico (see Kellogg 1932, and Smith and Taylor 1948) concern not only Hyla are- nicolor but also Hyla cadaverina, a species distinguished from arenicolor by Gorman (1960) who named it Hyhi californiae. The nature of the variation in size and call structure in this species remains unsettled; Jack R. Pierce of Austin College is currently investigating inter-populational variation in this species. Hyla arenicolor is the disjunct southwest- ern representative of the Hyla versicolor group; its present distribution probabh' is a result of continuous more favorable habitats during pluvial periods of the Pleistocene or the post-Wisconsin time and its ability to sur- \i\e in moist pockets in canyons in otherwise highly unfavorable environments. Etymology: The specific name is derived from the Latin arena, meaning sand and the Latin color, meaning color, and refers to the dull brown dorsal ground color of this species. Distribution: Hyla arenicolor occurs in mountainous areas and on high plateaus from soutliern Utah and Colorado southward to include the eastern two-thirds of Arizona, New Mexico, and west Texas in the United States and the Mexican Plateau and associated mountain ranges, southward to Michoacan, Guerrero, and western Oaxaca (fig. 256). The species occurs at ele\'ations between 300 and 3000 meters. See Appendix 1 for the locality records of the 599 specimens examined. Genus Ptychohyla Taylor Ptycholu/la Taylor, 1944a, p. 41 [tv'pe species, Ptycliohyla adipoveniris Taylor, 1944a = Hyla Icon- hardschiiltzci Ahl, 1934]. Smith and Taylor, 1948, p. 91. Stuart, 1963, p. 40. Duellman, 1963c, p. 314. Generotype: Hyla leonhardschultzei Ahl, 1934. Taylor ( 1944a, p. 41 ) proposed the generic name Ptychohyla for a new species, Ptychohyla adipoveniris, described in the same paper (p. 41 ) . Duellman ( 1960c ) com- pared the holotype of P. adipoveniris with that of Hyla leonhardschultzei and concluded that they were representative of the same spe- cies. Ety'mology': The generic name is deri\ed from the Greek ptycho, meaning layer of plate, and Hylas, a character in Greek my- thology. The generic name is in reference to the plate-like ventrolateral glands character- istic of this genus. Definition: Frogs of the genus Ptycho- hyla are small to medium in size and have a uniform green or brown dorsum or one that is marked by darker blotches. The flanks are uniform white or marked by black spots, and the \entcr is white or yellow, with or without spots. The iris is a deep bronze, copper, or red. The palpebral membrane is unmarked. Till- hands are about one-third webbed or ha\e only a \estige of a web between the fin- 1970 DUELLMAN: HYLID FROGS 519 gers. The toes are about two-thirds to three- fourths webbed. Breeding males are char- acterized by a pair of thickened, pigmented ventrolateral glands, which are usually more distinct in preserved than in li\ing specimens. Breeding males of some species have nuptial excrescences. The vocal sac is single, median, and subgular. The skull is broad, flat, and has a large frontoparietal fontanelle. The sphenethmoid is wide and broadly attached to the elongate, rather slender nasals, which are separated medially and lie parallel to the ma.xillaries. The anterior arm of the squa- mosal is short and extends less than half the distance to the maxillary. The quadratojugal usually is reduced to a small spine-shaped 30 ilO° 106° 102° 98° — I — 30' KILOMETERS 18' 110° 106° 102° 98° Fic. 256. Distribution of Hyla arenicolor in Me.vico. 520 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 element posteriorly that does not articulate with the maxillary. Teeth are present on the premaxillaries, maxillaries, and prevomers, but are absent from the palatines and parasphe- noid. The teeth are simple, elongate, and coni- cal. The teeth on the premaxillary and an- terior part of the maxillary are longer, pointed, and terminally curved backwards; whereas posteriorly on the maxillary the teeth become progressively shorter and blunter. The tad- poles are adapted to live in mountain streams; they have stream-lined bodies and long tails bearing low fins. The mouth is large and di- rected ventrally. There are three upper and three lower rows of teeth in a funnel-shaped mouth in the members of one species group and minimally four upper and six lower rows of teeth in a broad marginate mouth in the members of a second group. The mating calls consist of a series of short notes or a single long note. The haploid number of chromo- somes is 12, and the diploid number is 24 (known in P. ignicoJor and Jeonhardschult- zei). Composition of the Genus: Five species are currently recognized; two are polytypic, each containing two subspecies. All known species occur only in Middle America. Of the five species, 404 preserved frogs, 13 skeletons, 57 lots of tadpoles, and one preserved clutch of eggs were examined. Analysis of Characters: The largest .spe- cies is Phjchohijla eutlnjsanoia; the largest specimen examined is a female of the nomi- nate subspecies having a snout-vent length of .5.3. .3 mm. Members of the schinidtorum group {ignicolor and schmidiorum) are notably smaller; the largest male is 32.8 mm. and the largest female, 38.0 mm. In all species the females are 10 to 15 per cent longer than the males. Few difi^erences in proportions exist between the species (table 51), but cer- tain morphological characters are consistently diflerent between species. A vertical fleshy rostral keel is present in leonhardsclndtzei and spinipollex but lacking in the other species. These two species, plus euthijsanota have the fingers about one-third webbed, a nuptial ex- crescence consisting of a cluster of spines, and a weak tarsal fold ("figs. 257 and 258). PUjcho- hijla ignicolor and schmidtorum lack a tarsal fold and nuptial excrescences and have only vestigial webbing between the fingers. The ventrolateral glands distinctive of breeding males are not readily visible in liv- ing individuals of ignicolor and schmidtorum, but in preservative they show as distinctive orange-tan areas. The glands are more dis- tinct in euthijsanota; in some of these the glands are elevated above the surface of the surrounding skin. The extent of the glands is variable (fig. 259), but some of the varia- TABLE 51 Comparison of Sizes and Proportions, with Means in Parentheses, of Males of the Taxa of Ptychohyla. Taxon N P. s. schmidtorum 25 P. s. chamulae 40 P. ignicolor — . 38 P. e. euthijsanota 17 P. e. macrotijmpanum 5 P. leonhardschultzei — 36 P. s))inipollex 32 Snout-vent Length Tibia Length/ S-V L Foot Length/ S-V L Tyinpanum/ Eye 29.0-32.8 (31.0) 26.3-30.5 (28.0) 26.6-30.9 (28.1) 28.9-3S.1 (35.0) 32.0-38.0 (34.9) 28.5-35.6 (31.9) 29.0-41.0 (37.1) 0.453-0.524 (0.481) 0.460-0.519 (0.482) 0.458-0.496 (0.481) 0.444-0.550 (0.487) 0.488-0.520 (0.502) 0.472-0.544 (0.512) 0.469-0.531 (0.490) 0.371-0.409 (0.319) 0.386-0.429 (0.404) 0.380-0.429 (0.406) 0.349-0.405 (0..380) 0.405-0.424 (0.417) 0.386-0.453 (0.420) 0.388-0.426 (0.408) 0.515-0.593 (0.547) 0.482-0.656 (0..549) 0.366-0.531 (0.429) 0.486-0.638 (0.563) 0.500-0.571 (0.541) 0.447-0.619 (0.515) 0.450-0.552 (0.495) 1970 DUELLMAN: HYLID FROGS 521 Fig. 257. Hands of the species of Ptijchohyla. A. P. schmidtorum schmidtorum, K.U. No. 58037. B. P. euthysanota euthysanota, K.U. No. 58010. C. P. ignicolor, K.U. No. 87153. D. P. leonhard- schultzei, K.U. No. 101049. E. P. spinipollex, K.U. No. 58057. X 6. 522 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fig. 258. Feet of the .species of Pttjchohyla. A. P. schmidturum ichmidtorum, K.U. No. 58037. B. P. euthysanota cutht/sonota, K.U. No. 58010. C. P. tgnicolor, K.U. No. 87153. D. P. leonhardschultzei. K.U. No. 101049. E. P. spiuipollcx, K.U. No. 58057. X 6. 1970 DUELLMAN: HYLID FROGS 523 Fig. 259. Extent of ventrolateral glands in three species of Ptijchohyla. A. P. euthysanota euthysanota, K.U. No. 58009. B. P. schmidtorum schmidtorwn, K.U. No. 58033. C. P. ignicolor, K.U. No. 87158. X 2. Note the chin gland and pigmentation in P. ignicolor. tion probably is due to different degrees of development in individual frogs rather than to interspecific differences. Most specimens of P. ignicolor and some of P. schmidtorum chamulae have a small, round glandular area on the chin. The dorsum is green in ignicolor and cham- ulae; in the other species the dorsum is brown, reddish brown, or olive-brown with or with- out darker blotches or reticulation. The ven- ter and flanks are boldly spotted with black in leonhardschultzei and spinipollex; the ven- ter is weakly spotted in euthysanota and im- maculate in schmidtorum whereas small flecks are present in ignicolor. The anterior and pos- terior surfaces of the thighs are brown or orange-brown, e.xcept in ignicolor, which has red or orange-red surfaces of the thighs. Ptijchohyla euthysanota and schmidtorum have a white labial stripe that is continuous onto the flank, and all species have a pale transverse stripe above the anus and white or cream stripes along the outer edges of the forearm and tarsus (pi. 67). The tadpoles of ignicolor and schmidtorum have large funnel-shaped mouths; the teeth are arranged in short rows, three above and three below the beaks, which have long, pointed serrations but lack lateral processes. The tadpoles of euthysanota, leonhard- schultzei, and spinipollex have large mouths with a lateral fold and two rows of labial papillae; the teeth are arranged in long rows, four above and six below the beaks, which have short, peg-like serrations and long lat- eral processes (figs. 260 and 261). Tlie skulls of the various species are nearly alike, except that the quadratojugal-maxillary arch is always incomplete in euthysanota, igni- color, and schmidtorum, whereas in some specimens of leonhardschultzei and spinipol- lex the arch is complete (fig. 262). Further- more, the premaxillaries are longer and bear more teeth in ignicolor and schmidtorum than in the other species. The mating calls of ignicolor and schmid- torum consist of a series of short notes and differ from one another in that the notes are shorter, more slowly pulsed, but higher pitched in schmidtorum. The calls of euthy- sanota, leonhardschultzei, and spinipollex con- sist of one long note and differ in duration, pulse rate, and pitch (table 52, pis. 30 and ,31). Distribution: The combined distributions of the five species of Ptychohyla include the Atlantic and Pacific slopes of the highlands of nuclear Central America and southern Mex- 524 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 ■ ■S«'<«.?. ^'+'-- inipollex Schmidt, 1936, p. 45 [holotype, M.C.Z. No. 21300 from "mountains behind Ceiba," Departamento Atlantidad, Honduras; Raymond E. Stadehiian collector], Plycliohyla spinipollex: Stuart, 1954b, p. 48; 1963, p. 41. Duellman, 1963c, p. 327. Dl\gnosis: This medium-sized species has a tarsal fold, rostral keel, and fingers about one-third webbed. The nuptial spines are moderately large, pointed, and few in num- ber. The eyelid is about as wide as the in- terorbital space, and the snout is rounded abo\e. Ptyclwhyla spinipollex differs from leonhardschidtzei by having fewer and larger nuptial spines, relatively narrower interorbital space, and rounded, instead of an angular snout. Description: This is the largest species in the genus of Ptychohyla. Males attain a maximum snout-\'ent length of 41.2 mm. ( mean, 32 specimens from Finca Los Alpes, Alta Verapaz, Guatemala, 37.1 mm.), and females reach 44.6 mm. (mean, 6 specimens, 42.8 mm.). In the sample of males from 102° 98° 94° [8° 1 s _ 18° 1 0 50 100 200 ^"~^~\_ s • r-^*^ 1 KILOMETERS 1 1 ^'~"^^— ^'^ 102° 98° 94° Fic. 265. Distribution of Ptychohyla leonhardschultzei. 1970 DUELLMAN: HYLID FROGS 545 Finca Los Alpes, the ratio of tibia length to snout-\ent length is 0.469 to 0.531 (mean, 0.490); the ratio of the foot length to snout- vent length is 0.388 to 0.426 (mean, 0.408); the ratio of head length to snout-vent length is 0..30S to 0.335 (mean, 0.321); the ratio of head width to snout-vent length is 0.296 to 0.322 (mean, 0.311), and the ratio of the diameter of the tympanum to that of the eye is 0.4.50 to 0.552 (mean, 0.495). An insuffi- cient number of \vell-preser\'ed specimens are available from other parts of the range in order to determine if there is any geographic variation in size and proportions. The head is as wide as the body; the top of the head is flat. The interorbital distance is only slightly greater than the width of the eyelid; the ratio of the width of the eyelid to that of the interorbital space is 0.878 to 0.923 (mean, 0.905). In dorsal profile the snout is bluntly rounded with a terminal point resulting from the presence of a \ ertical, fleshy rostral keel. In lateral profile the snout is rounded above and truncate. The snout is moderately long; the nostrils are situated at a point about three-fourths the distance from the eye to the tip of the snout. The canthus is angular; the loreal region is barely con- cave, and the lips are thick and moderately flared. A moderately heavy dermal fold ex- tends posteriorly from the posterior corner of the eye abo\e the tympanum to a point above the insertion of the arm. The fold ob- scures the upper edge of the tympanum, which otherwise is distinct. The tv'mpanum is posteroventral to the eye and separated from the eye by a distance equal to the diam- eter of the tympanum. The arm is moderately short and robust. A short axillary membrane is present. A row of low tubercles forms a distinct ridge on the ventrolateral edge of the forearm; a weak transverse fold is present on the \vrist. The fingers are moderately short and stout and bear relati\eh' large discs; the disc on the third finger is equal to the diameter of the tympanum. The subarticular tubercles are large and round; in about two-thirds of the specimens the distal tubercle on the fourth finger is bifid. Small, subconical supernu- merary tubercles are present on the proximal segment of each digit. A low bifid palmar tubercle is present. The prepollex is moder- ately enlarged; in breeding males the nuptial excrescence is in the form of 35 to 66 ( mean, 47.4) sharply pointed spines. The fingers are about one-third webbed (fig. 257E). The web between the first and second fingers is \estigia], but connects the second finger at the middle of the penultimate phalanx to the middle of the antepenultimate phalanx of the third and from the distal end of the ante- penultimate phalanx of the third to the base of the penultimate phalanx of the fourth fin- ger. The hind limbs are relatively short and robust; the adpressed heels barely overlap. The tibiotarsal articulation extends to the middle of the eye. A distinct, but low and rounded, tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is low, flat, ovoid or quadrangular, and visible from above. The outer metatarsal tubercle is low, round, and distinct in most specimens. The toes are long and slender and bear discs that are nearly as large as those on the fin- gers. The subarticular tubercles are moder- ately large and subconical. Numerous small, round supernumerary tubercles are present on the proximal segments of each toe. The toes are about three-fourths webbed (fig. 258E). The web extends from the base of the disc of the first toe to the distal end of the antepenultimate phalanx of the second, from the distal end of the penultimate pha- lanx of the second to the base of the third, and from the distal end of the penultimate phalanx of the third to the base of the penul- timate phalanx of the fourth and onto the base of the disc of the fifth toe. The anal opening is directed posteriorly near the upper level of the thighs; the open- ing is bordered laterally by moderately heavy dermal folds and ventrolaterally by tubercles. The anal sheath is short and thin. The skin on the dorsum and the ventral surfaces of the forelimbs and shank is smooth; that on the throat, belly, and ventral surfaces of the thighs is granular. The \entrolateral glands extend from the axilla for about two-thirds the length of the body; they do not reach the groin and are broadly separated midventral- ly. The tongue is ovoid, marginate or shal- lowly notched posteriorly, and barely free behind. Males have three to seven (mean, 4.9) prevomerine teeth, and females have six to 10 (mean, 7.6) prevomerine teeth situ- 546 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 ated on transverse or posteromedially slanting ridges between the ovoid choanae. The vocal slits extend from the midlateral base of the tongue to the angles of the jaws. The vocal sac is single, median, subgular, and not great- ly distensible. The general coloration of Ptijchohyhi spinipollex is yellowish tan with small irregu- lar brown markings on the dorsum (pi. 67, fig. 7). A typical specimen from northern Guatemala has a yellowish tan dorsum with brown and dark brown markings on the back. In most individuals, the markings consist of small interconnected spots and dark brown flecks. Irregularly shaped dark marks on the limbs tend to form transverse bands. The posterior surfaces of the thighs are creamy tan. The first and second fingers are creamy tan, and the third and fourth fingers and webbing on the hand are grayish brown. The webbing on the feet is dark brown. The axilla is pale pink, and the flanks are buff, becoming yellow in the groin. The flanks are marked by dark brown spots. There is a faint white stripe along the ventrolateral edges of the forearms and tarsi, and a narrow white line above and beside the anal opening. There is no white stripe on the edge of the upper lip. The belly is dusky cream with numerous brown to dark gray flecks. The ventrolateral glands are grayish tan. The iris is dull gray- ish bronze. A typical specimen from Honduras (K.U. No. 103225, from Cerro Uyuca) had in life a tan dorsum with oli\e-gray markings. The flanks were white with dark brown spots. The groin, anterior and posterior surfaces of the thighs were dull yellow, and the belly was immaculate white. The iris was dull gravish bronze. In preservative the dorsum varies from grayish tan to dark brown with darker brown or black reticulations on the head and body and dark brown transverse bars or spots on the limbs. The anterior surfaces of the thighs are reddish tan, and the posterior surfaces are yellowish tan. The white stripes char- acteristic of li\'ing indi\iduals persist in pre- ser\ed specimens. The belly is dull white with scattered brown flecks; the flanks arc grayish white with dark brown spots. The ventrolateral glands are grayish tan. In specimens from Finca Los Alpes, Gua- temala, the color of the dorsum varies from pale tan to dark brown with darker brown markings; the white line above the anus is present in all specimens, but it is indistinct in some. Two individuals have a dark brown dorsum with large pale tan square blotches; in life the blotches were pale tan and the dorsum was dark brown. All Guatemalan specimens ha\e dark brown spots and flecks on the \'enter. Some individuals have only a few flecks on the throat and a few large spots on the flanks. Other specimens have dense spotting over the entire venter. One indi\idual was dark brown with many small white flecks on the dorsum. All Hondura- nean specimens either have an unmarked white venter or only a few small flecks on the edge of the chin. Furthermore, the Hondu- ranean specimens have fewer and smaller dark spots on the flanks, which tend to be paler than the flanks in Guatemalan speci- mens. Tadpoles: A typical tadpole in develop- mental stage 31 from Finca Los Alpes, Alta Verapaz, Guatemala, has a total length of 38.7 mm. and a body length of 14.0 mm. The body is rounded and not depressed; it is as wide as deep and ovoid in dorsal profile. The mouth is directed ventrally. The eyes arc small and directed dorsolaterally. The nostrils arc barely protuberant and are di- rected anterolaterally; the nostrils are slightly closer to the tip of the snout than to the eyes. The spiracle is sinistral and posterovcntral to the eye; the anal tube is dextral. The tail is long, low, and pointed posteriorly. The cau- dal musculature is heavy and nearly extends to the tip of the tail. The dorsal fin barely extends onto the body and reaches its great- est depth at the mid-length of the tail; the \cntral fin has an equal depth throughout most of its length (fig. 260E). The mouth is large and has deep lateral folds in the lips, which are bordered b\' two rows of small papillae; four or five additional rows of papillae are present in the lateral fold. The beaks are robust and bear short, peg-like serrations. The upper beak forms a broad arch with short, slender, round lat- eral processes. There are four upper and six or seven lower rows of teeth. The upper rows are about equal in length, and the fointh row is interrupted medially. The first four lower 1970 DUELLMAN: HYLID FROGS 547 rows are equal in length and only slightly shorter than the upper rows; the first lower row is interrupted medially. The fifth, sixth, and se\"enth ( if present ) lower rows have decreasing lengths. In many tadpoles the sev- enth lo\\er tooth row is absent or fragmentar}' (fig. 261C). The top of the head and the tip of the snout are brown; the venter is cream\' gray. The caudal musculature is tan, and the cau- dal fin is transparent. A faint creamy, nar- row, crescent-shaped mark is present on the posterior edge of the body in most specimens, but it is not bordered posteriorly by a dark broun mark. Dark brown flecks are scattered on the caudal musculature and the caudal fin with the exception of the anterior one-half of the ventral fin. In life the eye is bronze. Matixg Call: The call of PUjchohyla spinipoUex consists of a single note, "wraack," repeated at intervals of 45 seconds to four minutes. Each note has a duration of about 0.46 seconds and about 147 pulses per second. The dominant frequency is 4300 cycles per second (pi. 31, fig. 3). Natural History: This species occurs in cloud forests and mixed pine and broad- leafed forests, where the frogs breed in cas- cading mountain streams. Calling males are found on bushes and trees along the streams, and tadpoles ha\e been found in the streams, where they occur primarily in shallow gravel- bottomed pools or in riffles. Two recently metamorphosed young have snout-vent lengths of 15.0 and 15.5 mm. Remarks: Duellman (1963c) and Lynch and Fugler (1965) mentioned that Guate- malan specimens differ from those in Hon- duras by having a hea\ily spotted venter. When I reviewed the genus PtychoJujIa in 1963, I had not seen living specimens from Honduras and was reluctant to recognize taxonomically the Guatemalan population. I have now seen living indi\'iduals from Cerro Uyuca, Honduras; the differences in colora- tion ha\e been described in the preceding description of this species. Nonetheless, I am still reluctant to recognize taxonomically the Guatemalan specimens, until information is available concerning the tadpoles and the mating call of the frogs in Honduras. Per- haps the Guatemalan populations here re- ferred to as Ptijchohyla spinipoUex actually represent a distinct species. At the present time, only a few specimens from widely scat- tered localities are available from Honduras. Etymology: The specific name spinipol- lex is derived from the Latin spina meaning thorn and the Latin pollex meaning thumb and alludes to the spinous nuptial excres- cences in breeding males. Distribution: Ptychohyla spinipoUex in- habits cloud forests at elevations of 800 to 1850 meters on the Atlantic slopes of the highlands in Nuclear Central America from the Sierra de los Cuchumatanes in western Guatemala southeastward to north-central Nicaragua (fig. 264). See Appendix 1 for the locality records of the 67 specimens examined. Genus Plectrohyla Rrocchi Plectrohijla Brocchi, 1877a, p. 92 [type species by original designation, Plectrolitjla guateinalensis Brocchi, 1877a I . Caiiphias Brocchi, 1877b, p. 129 [substitute name for Plectrohyla Brocchi, 1877a]. Generotype: Plectrohyla guateinalensis Brocchi, 1877a, by original designation. Hart- weg ( 1941, p. 1 ) discussed the generic alloca- tion of Brocchi's names: "The generic and specific descriptions of Plectrohyla guatcmalensis, a batrachian from Patzizia, Guatemala, were formulated by Brocchi ( 1877:92). In the course of his study he discovered another new species which he believed to be closely related to guateinalen- sis. Deciding that the original description of Plectrohyla was not satisfactory for the in- clusion of both species (gtiatemalensis and the new one), he described a new genus, Cauphias, and synonymized Plectrohyla with it (1877:129). In the same article (p. 130) he also described his new species, crassits. The species guatcmalensis is the haplotype [monotype] of Plectrohyla (Brocchi, 1877: 92); Barbour (1927:96) designated Plectro- hyla guatcmalensis as the genotype of Cau- phias. Although it cannot be definitely shown that the actual publication date of the de- scription of Plectrohyla preceded that of Cauphias, it seems best to assume so; I there- fore regard the names Cauphias and Plectro- hyla as synonyms and select Plectrohyla as 548 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 the proper name to be used. Should future researches show that crassus (crasstim), is generically distinct, the name Cauphias may not be resurrected, since it is a synonym of Plectrohyla." Hartweg's assumption that the description of Plectrohyla antedates that of Cauphias is logical, because both names were published in the Bulletin de la Societe Philomathique de Paris (Scries 7, volume 1). The descrip- tion of Plectrohyla is on page 92 in number 2, and that of Cauphias is on page 129 in num- ber 3. Duellman (1964b, p. 488) showed that Cauphias crassus is actually a member of the Hyla bistincta group. Etymology: The generic name is derived from the Greek plektron, meaning spur, and Hylas, a character in Greek mythology. The generic name is in reference to the prepolli- cal spines, characteristic of members of the genus. Definition: Frogs of the genus Plectro- hijla are moderately small to large in size; they are variously colored but usually have a green, gray, or brown dorsum and lack bright markings. The pupil is horizontal, and the palpebral membrane is clear. The fingers and toes are long and bear moderately large discs. The webbing on the hands is vestigial, whereas that on the feet is extensive. Indi- viduals of both sexes have an enlarged pre- pollex that is supported internally by a large bony element, the prepollical spine or pro- cess; the spine protrudes through the skin in some species. Vocal slits and a single, me- dian, subgular vocal sac are present in males of four species and absent in the others. The skin on the dorsum is thick and glandular; it is smooth in some species but tuberculate in most. There is no integumentary-cranial co- ossification. The lips are thickened and the forearms arc hypertrophied in breeding males of some species. The skull is broad and moderately shallow. The skull is characterized by a frontoparietal fontancllc, a well-ossified sphenethmoid, and relatively small nasals, which in most species are separated medially and bear a slender maxillary process that articulates with the well-developed posterior process of the pars facialis of the maxillary (fig. 266). The squa- mosal is robust, and the anterior arm does not extend to the maxillary. The pterygoid is ro- bust, and the median ramus of the pterygoid is in bony contact with the prootic. The quadratojugal is absent or reduced to a small spur posteriorly. The maxillary and premax- illary are robust. An apparently unique condi- tion of the premaxillaries distinguishes this genus. The alary process of the premaxillary is bifurcate posteriorly. The dorsal tip of the alary process lies adjacent to the nasal carti- lages anteroventral to the nasals. The poste- rior ramus of the alary process extends be- neath the anterior part of the sphenethmoid (fig. ISA). Thus, the premaxillaries and alary processes support the entire nasal re- gion and anterior end of the sphenethmoid. Teeth are present on the premaxillaries, max- illaries, and prevomers, whereas the palatines and parasphenoid are edentate. The teeth are blunt and weaklv bifid or long and point- ed (fig. 267). The known tadpoles are stream inhabi- tants with robust bodies and long muscular tails with low fins. The mouth is ventral and completely bordered by papillae but lacks lateral folds. There are two upper and three lower rows of teeth that are relatively short. The mating call consists of a single cjuack-like note or a series of short notes. The haploid number of chromosomes is 12 ( known only in /.v(7 and sagorum). Composition of Genus: Ten monotypic species arc recognized in the genus, which is endemic to the highlands of Nuclear Cen- tral America. Of these species, 584 preserved frogs, 22 skeletons, and 57 lots of tadpoles have been examined. Analysis of Characters: Plectrohyla avia is the largest species and matudai is the small- est; males of the former attain snout-vent lengths of 90 mm., and, of the latter, .36 mm. Females of the small species (/.v//, matudai, quecchi, and sagorum) are somewhat larger than the males, whereas in the larger species the females, if known, are about the same size as the males. The sizes and proportions of the ten species are summarized in table 53. The taxonomically important external cliaractcrs are those of the skin, shape of the snout, and nature of the prepollical spine. Plectrohyla guatcmalcnsis, hartuegi. matudai. pycnochila, and sagorum have numerous and 1970 DUELLMAN: HYLID FROGS 549 Fig. 266. Dorsal (A) and lateral (B) views of the skull of Plectrohyla guatemalensis, K.U. No. 68664. x 6. conspicuous tubercles on the dorsum, where- as the skin in the other species is relatively smooth (some have scattered low tubercles), except in avia, which has tubercles on the head. The snout is bluntly rounded in avia, guatemalensis, hartivegi, pycnochila, and quecchi, truncate in matudai and acuminate in the other species. A narrow, fleshy, verti- cal rostral keel is present in quecchi and sagorum. Adults of both sexes have bony prepollical processes; these usually are better developed in males than in females. In some males, the spine actually protrudes through the skin. The shape of the prepollical process is one of the most useful taxonomic characters in this genus, and the shape varies from a flat plate or elongate, rounded blunt spur to a simple curved spine or bifid spine (fig. 268). The fingers of all species are long and bear moderately large discs. The feet are moderately well webbed and have large sub- articular tubercles (figs. 269-273). A strong inner tarsal fold is present in all species, and a distinct outer tarsal fold is present in some populations of P. glanchdosa. The arms of breeding males are greatly hypertrophied, and in some breeding males the lips are swollen. No histological examina- tion of the lips has been made, but examina- tion of the hypertrophied arms revealed that there was no modification, other than extreme muscular development in several species ( glanduhsa, guatemalensis, ixU, matudai, quecchi, and sagorum), but that in avia the humerus is greatly modified (fig. 274). The humerus is massive with well-developed 550 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fig. 267. Lateral view of maxillary teeth. A. Plectrohyla avia, K.U. No. 106295. B. Plectmlnjla guatemalensis, K.U. No. 68664. x 20. ridges. The crista ventralis begins on the ca- pitulum and continues as a heavy, moderately deep ridge for about 40 per cent of the length of the humerus. The crista mediahs is greatly expanded on the distal half of the humerus. The crista lateralis extends for two-thirds of the length of the bone. A broad, deep, V- shaped depression exists between the crista medialis and the crista lateralis. Linea masculinea are present in P. ixil and nuitudai and apparently absent in other spe- cies. The skin of P. glandulosa contains a layer of melanin; presumably this is an adap- tation protecting the internal organs from solar radiation in this species which frequent- ly basks on rocks and clumps of grass. The teeth in four species (avia, glandu- losa, lacertosa, and sagorum) are pointed; in the others the teeth are barely spatulate and not, or only weakly, bifid (fig. 267). There is considerable variation in the number of teeth (table 54). In most groups of related species of hylids the larger species have more teeth than do the smaller species, but in Plectrolitjia, this is not the case, the smallest species (mattidai) has more teeth than any of the three largest species (avia, guatemalen- sis, and hartwegi). The tadpoles of six species are known. Of these, the tadpoles of P. guatemalensis are different in having two rows of papillae bordering the mouth and blunt peg-like ser- rations on the beaks. The tadpoles of guate- malensis and glandidosa are alike in having robust bodies and heavy wrinkled skin (fig. 275), but glandulosa differs by having one row of fringing papillae; in this respect, it is like the other known tadpoles (table 55). The tadpoles of matudai and ixU are unique by •lljj'vi*-'" WA'il" *HHi' Fig. 268. Prepollical processes of Plccirohylci (palmar \ie\v of right hand). A. P. glandulosa, K.U. No. 59828 (.same in pijcnochila) . B. P. lacertosa, U.I.M.N.H. No. 33693, C. P. ixil, K.U. No. 59834 (same in maliulai, quccchi, and sagorum). D. P. avia, K.U. No. 106295. E. P. guatemalensis, K.U. No. 68664 (same in hartwegi). X 10. 1970 DUELLMAN: HYLID FHOGS 551 CO lO W pa < c a « 5^ .9 R- o o PL, c o c o ir> Mh c o CO t^ -* CO CO CO t^ t-~< ^H CD CO CO . . ■* t^ t-- lO ;;^"^ o t^ S^-r ;irio;^-r ^ '^. S^^ST't o 05 o -* o 35 O ^H O t^ O t^ O CO O (M O o -^in -*: CO ^ CO -* in "^ 4< -* t^-t 6^ CO o O CO O CO O ^ O lO O CD O 05 O ^ O 05 Tf -~'CO -^co ^^ CO --- C<1 ^-^ -* -^ CO ^^OJ ^-^tT o O lO lO o CD C35 CO (35 Tj< ^^c~i ''^co :^co • (M • lO _■ O ■■ CO o 03 ^ ini o ic o CO ^ CO _:^ CO cc ^ c^] CO n- O CO O O O -1" o CO v^CO ^-^CO ^^co ^ ^^ro ^^ N ^ O] CO ID O lO CO CO c^, CO CO O CO O C i< i^ ^ O r^ O -A <^. 6 05 O lO CM ^^CO (M 05 CD 00 ^-^CD ,-^-* ^-^ <^ m ^ oi ^ lo . O m O CO O ^^ o I c^ ■ c^ ■ c^ ■ Tt< '• CD ■■ lO _■ ^ CO C3 00 "^ CO "^ 05 CO ^ CO IC CO lO o ^ o o o5 c-l d 05 CO ^-^ cc ^ (M CO o c^ O CO CO CO CO ' ' — ^ d c^^ d oQ (^ v-^ CO ~r CO ~t c-1 ^ <^ ' ' -^ lO o o <= ' M ^^ CO — ' d d o^d CO d O ^ f- o CO ^ d CD ^-^ ■^ CO O CO d d CO 05 CO "^ oq Ol ,-^CO ^-^ 00 ^ dcodcodcoo2SS CO r^ ^^ ,-^co .^-^ . -J "^ lO '^ lO ■ -^r ■ -J' ,^ ^ ^ ' "^ ' -^ !. "^ SdgdgdSodo^g^g^ a d o ci d d c5 •^ O 05 ^ 05 ^ o o o CD '^ CO 02 O CD CO CO CM CO CD 1—1 -"^ _ O) lO ,-^ O5|3o'2^ 05 O CM O CO OCDOi-^Ot^OOOOcOOCOO ^^ T}< - -lo ^-^-^ — -lO - — -^ — - '^ d d d d d d '< CO "-: Ol -i^ "-: 05 • 0^hO-^050oO lo ^', lo ■* ~;, lo ic 05 O lO ^ o in ^ _CD ,^in^^co^_^. ;,_^-.. i 02 in CO CO CM d CD d ' X< -r X< ^ ^^ -* ^ «? in d lO 00 N- CO -^^ CO T d T c>i T ^ ^ CD" t^ ^.■' CD '7 ^ ^ ^ incoOTcoffico^-^oi't'inin-^rinoocMcDcoin a; S oi g P g ^ • 05 -x t~- 'CD CO CM in oq rt 'o 'O] 'CM in CM 00 in o CM CO 1= o o CJ o o CO O ?^ -S fi, a. 552 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fig. 269. Hands of four species of Plcctrolujla. A. P. matudai, K.U. No. 58869. B. P. ixil, K.U. No. 58854. C. P. sagorum, K.U. No. 103164. D. P. quccchi, K.U. No. 64115. x 4. having unequal serrations on the upper beak. In ixil two or three serrations on either side are noticeably enlarged, whereas in matudai one serration is enlarged and fang-like on either side. The serrations are subequal in the other species (fig. 276). Distribution: Frogs of the genus Plectio- hijla occur at moderate to high elc\'ations (1000 to 3500 meters) in the highlands of northern Central America (Honduras, El Sal- \ador, Guatemala, and the states of Chiapas and Oaxaca, Me.xico). Ecologically the spe- cies inhabit montane meadows, pine-cypress forest, pine-oak forest, and cloud forest. All presumably are stream-breeders. Discussion: The members of the genus fall into two seemingly natural groups: 1) Small species having vocal slits {ixil, matu- dai, quecchi, and sap,oru)n). 2) Medium- sized to large species lacking \ocal slits (avia, glandulosa, guatemalensis, harticegi, la- certosa, and ))tinorhila). 1970 DUELLMAN: HYLID FROGS 553 Fig. 270. Feet of four species of Plectrohyla. A. P. matudai. K.U. No. 58869. B. P. ixil, K.U. No. 58854. C. P. sagonim, K.U. No. 193164. D. P. quecchi, K.U, No. 64115. x 3.5. 554 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fic. 271. Hands of three species of Plectrohyla . B. P. lacertosa, U.I.M.N.H. 33693. C. P. glandulosa, K.U. No. 58708. A. P. pycnochila, T.C.W.C. No. 21459. X 4. 1970 DUELLMAN: HYLID FROGS 555 Fig. 272. Feet of three species of Plcctrohyla. A. P. glandulosa, K.U. No. 58708. B. P. lacertosa, U.I.M.N.H. No. 33693. C. P. pijcnochila, T.C.W.C. No. 21459. X 3. Plectrohijla apparently is closely related to the Hyla bistincta group in the Mexican highlands northwest of the Isthmus of Te- huantepec. Probably Plectrohijla and the Hyla bistincta group e\'olved from a common ancester. The members of both groups show parallel adaptations to the lotic environment; some members of each group have lost the voice. Since the Hyla bistincta group has evolved in the Mexican highlands, it is only logical to assume that the ancestral stock that gave rise to Plectrohyla was isolated in north- ern Central America. The Plectrohyla stock probably was in the area this is now Chiapas and Guatemala in the Miocene prior to the uplift of Nuclear Central America that began in the Pliocene. Con- ceivably, in the course of uplift, the Plectro- hyla stock was separated into a highland component and another component at mod- erate elevations on the slopes. The latter component retained vocal slits and evolved into a group of small species, whereas the highland component evolved into a group of larger species lacking vocal slits. The former group, which for convenience can be called the sagorum group, eventually established populations on the Atlantic and Pacific slopes of the highlands. Possibly, the species now known as quecchi was the orig- inal inhabitant on the Atlantic slopes, whereas matudai was endemic to the Pacific slopes. Through isolation differences in voice, shape of the snout, and mouthparts of the tadpoles developed. Subsequent climatic fluctuation, probably in the Pleistocene, permitted mi- gration southward of the quecchi-s\.ock and northward of the matuclai-stock. Depression of climatic zones and uplift through volcan- ism again resulted in isolation of populations on Atlantic and Pacific slopes, but this time two species were present on each slope. The 556 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Via. 273. Hands and feet ot three species ot Plectrnhi/la. A and D. P. avia. K.U. No. 94016. B and E. P. i^uatcmcilcnsh; K.U. No. 64102. C. and F. P. Iiiirtwciii, U.M.M.Z. No. 94428. X 2. 1970 DUELLMAN: HYLID FROGS 557 TABLE 54 Dentitional Characteristics of the Species of Plectrolnjla. Fig. 274. Posterolateral view of the left humerus of Plectrohyla avia, K.U. No. 106295. CL=crista lat- eralis, CM=crista medialis, CV=crista ventralis, X 3. matudai-stock on the Atlantic slope differen- tiated into ixil, and the qtiecchi-stock on the Pacific slope evolved into sagorum. This close relationship between sagorum and quecchi and between ixil and matticloi are obvious on the basis of morphological characters of the adults and tadpoles and in the similarities of the mating calls. Conceiv- ably, matudai and ixil are subspecifically re- lated, and possibly sagorum and quecchi are conspecific. Because each nominate species possesses a distinctive combination of mor- phological characters, in the absence of bio- logical e\idence to support a closer relation- ship, the four populations are regarded herein as distinct species. The relationships of the species in the highland component {guatemalen.sis group) are more obscure, because tadpoles are un- known for four species, and the absence of a voice precludes the use of that ta.xonomically useful trait. Plectrolnjla glandidosa and pycnochila seem to be the least specialized species; the condition of the prepollical pro- cess in these species probably is relatively unchanged from that of the Plectrohyla pro- totype and is much like that in the Hyla histincta group. It is possible that glandidosa Species Shape P. matudai Spatulate P. ixil Spatulate P. sagorum Pointed P. quecchi Spatulate P. glandulosa Pointed P. pycnochila Spatulate P. lacertosa Pointed P. avia Pointed P. guatemalen-, sis - Spatulate P. hartwegi Spatulate " One side only. Number of Teeth" Prevo- Maxillary- merinc Prema.xillary 3-5 50-61 3-5 41-58 3-4 34-45 3-4 47-53 1-3 23-30 3-5 31-36 2-3 30-31 1-3 27-,33 3-6 32-39 4-5 35-40 developed in the Sierra de Cuchumatanes in Guatemala while pycnochila was isolated in the highlands of central Chiapas. Apparently Plectrohyla avia represents an evolutionary intermediary between the gen- eralized glandulosa-pycnocliila stock and guatemalensis and hartwegi. The prepollical spine is long and pointed in avia (indepen- dently evolved in the sagorum group) and is bifid in guatemalensis and hartwegi. Plectro- hyla avia is endemic to moderately high ele- vations on the Pacific slopes; hartwegi occurs at the same elevations but farther \\'est. Plec- trohyla guatemalensis occurs nearly through- out the geographical (but not the altitudinal) range of the genus. It occurs .sympatrically with avia and possibly with hartwegi. The complicated paleogeography and climatic his- tory of Nuclear Central America undoubt- edly provided several barriers resulting in the isolation of populations which evoKed into the three species and provided the physical basis for their present distributions. Plectrohyla lacertosa is known to science solely by one miserably preserved adult male lacking specific locality data. Indeed, it is unfortunate that the specimen possesses such a distinctive combination of characters that it cannot be relegated to the synonymy of another species. This moderate-sized species with a unicjue prepollical process lacks vocal slits and thereby seems to belong in the 558 MONOGRAPH MUSEUM OF NATURAL HISTORY NO.l #«; ?«> Fig. 275. Tadpoles of six species of Plectwhyla. A. P. matudai, K.U. No. 60036. B. P. !Xi7, K.U. No. 60034. C. P. sanonim. K.U. No. 10419.5. D. P. quecchi, K.U. No. 00038. E. P. Klaudtdosa. K.U. No. 104193. F. P. fiimtcnwlcnsis, K.U. No. 60033. X 3. 1970 DUELLMAN: HYLID FROGS 559 Fig. 276. Mouths of tadpoles of six species of Plectrohyla. A. P. matudai, K.U. No. 60036. B, P. i.r!7, K.U. No. 60034. C. P. sagorum, K.U. No. 104195. D. P. quecchi, K.U. No. 60038. E. P. glandulosa, K.U. No. 104193. F. P. guatemalensis, K.U. No. 60033. X 10. guatemalensis group. Until more material is available nothing further can be ascertained about its relationships. Plectrohyla matudai Hartweg Plectrohyla matudai Harhveg, 1941, p. 5 [holo- t>pe, U.M.M.Z. No. 88863 from Cerro Ovando, Dis- trito Soconusco, Chiapas, Mexico, elevation 1800 meters; Norman Hartweg collector]. Smith and Tay- lor, 1948, p. 73. Lynch and Smith, 1966, p. 62 [synonymized Plectrohyla hrachycephala Taylor, 1949b, with P. matudai Hartweg, 1941]. Plectrohyla hrachycephala Taylor, 1949b, p. 16 [holot>-pe, A.M.N.H. No. 5.3761 from a tributary of the Rio Ostuta, at the foot of the Sierra Madre be- tween Sierra Madre and Cerro Atravesado, Oaxaca, Me.xico; Thomas C. MacDougall collector]. Plectiohyla matudai hrachycephala: Bunizahem and Smith, 1954, p. 62. Plectrohyla matudai matudai: Bumzahem and Smitli, 1954, p. 62. Stuart, 1963, p. 40. Diagnosis: This small species (37 mm. in snout-vent length) has a tuberculate dorsum, a blunt snout, and vocal sHts. The prepoHi- cal spine is long and pointed. The dorsal col- oration is separated from that on the venter by a dark line or irregular row of small spots. Plectrohyla ixil has a less tuberculate or smooth dorsum, an acuminate snout, and usu- ally a lateral light stripe. Plectrohyla quecchi and sagorum have a vertical rostral keel. The other members of the genus are larger and lack vocal slits. 560 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 TABLE 55 Comparison of Certain Features in the Known Tadpoles in Plectrohijla. Rows of Fringing Papillae Lateral Upper Tooth Lower Tooth Serrations Species Papillae to Beaks Rows Rows on Beak P. matudai - One; larger papillae Very few Moderately Third much Pointed; two medially long shorter fang-like on upper P. ixil -,.. One; larger papillae Few, scattered Moderately Third shorter Pointed; some medially long enlarged on upper P. sagorum _ One; larger papillae Few Moderately Third shorter Pointed; medially long subequal on upper P. quecchi One; larger papillae None Moderately Third shorter Pointed; medially long subequal on upper P. glandulosa One; row of larger None Moderately Third much Pointed; papillae medially long shorter subequal on upper P. guatemalensis —Two; row of larger Many Long Subequal Blunt; papillae medially subequal on upper Description: Males of this small species attain a maximum snout-vent length of 46.0 mm., and females reach 49.0 mm. In a series of eight males from Finca La Paz, Departa- mento San Marcos, Guatemala, the snout- vent length is 3L5 to 35.6 (mean, 33.1) mm.; the ratio of tibia length to snout-vent length is 0.496 to 0.540 (mean, 0.509); the ratio" of foot length to snout-vent length is 0.375 to 0.444 (mean, 0.412); the ratio of head length to snout-vent length is 0.334 to 0.382 (mean, 0.355 ) ; the ratio of head width to snout-vent length is 0.341 to 0.460 (mean, 0.392), and the ratio of the diameter of the tympanum to that of the eye is 0.400 to 0.575 (mean, 0.495). Specimens from the western part of the range apparently attain a larger size. Bumzahem and Smith (1954, p. 63) reported that the snout-vent length of nine males from Region de Soconusco, Chiapas, Mexico, was .36 to 46 mm.; Taylor (1949b, p. 19) reported snout- vent lengths of 35.0 to 40.0 mm. in four speci- mens from the Rio Ostuta, Oaxaca, Mexico. The head is as wide as the body, and the top of the head is flat; the snout is truncate in dorsal and lateral profiles; the snout is short, and its length is equal to the diameter of the eye. The nostrils are protuberant, di- rected dorsolaterally, and situated at the terminus of the snout. The canthus is slightly elevated and sharply angular, and the loreal region is nearly flat. The lips are moderately thick and barely flared. A moderately heavy dermal fold extends posteriorly from the eye, abo\e the tympanum, and downward to a point above the insertion of the arm. The fold obscures the upper edge of the tympan- um; the posterior edge of the tympanum is indistinct in most specimens, whereas the an- terior and ventral edges of the tympanum usually are well defined. The tympanum is posteroventral to the eye and separated from the eye by a distance slightly greater than the diameter of the tympanum. The arms are moderately short and ro- bust; they are swollen in some breeding males. A longitudinal row of tubercles is present on the ventrolateral edge of the fore- arm, and a transverse dermal fold is present on the wrist. The fingers are long and slender, and bear moderately large discs; the width of the disc on the third finger is greater than the diameter of the tympanum. The subartic- ular tubercles are large and conical; the distal tubercles on the third and fourth fingers are liifid in some individuals. The supernumerary 1970 DUELLMAN: HYLID FROGS 561 tubercles are large and subconical; usually they are present in a single row on the proxi- mal segment of the first and fourth fingers and in two rows on the proximal segments of the second and third fingers. An elevated, bifid palmar tubercle is present. The prepol- lex is moderately enlarged and terminalh- curved; in some males, the sharp prepollical spine protrudes from the terminus of the pre- pollex. The webbing on the hand is \estigial (fig. 269A). The legs are relatively short; the heels of the adpresscd limbs barely overlap. The tibiotarsal articulation extends to the eye. A few small tubercles are present on the heel, but there is no transverse dermal fold. An elevated tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is ovoid and subconical; no distinct outer metatarsal tubercle is present. The toes are moderately long and slender and bear discs that are noticeably smaller than those on the fingers. The subarticular tubercles are mod- erately small and conical; the supernumerary tubercles are moderately large, subconical, and arranged in a single row on each digit, except proximally the arrangement breaks down so that the tubercles are irregularly placed. The toes are about three-fourths webbed (fig. 270A). The webbing extends from the base of the penultimate phalanx of the first toe to the base of the penultimate phalanx of the second, from the distal end of penultimate phalanx of the second to the base of the penultimate phalanx of the third, from the middle of the penultimate phalanx of the third to the base of the penultimate phalanx of the fourth and on to the distal end of the penultimate phalanx of the fifth toe. The anal opening is directed posteroven- trally at the midlevel of the thighs. A short, broad anal sheath is bordered on either side by a moderately large tubercle. The skin on the dorsum is tuberculate; tubercles are pres- ent on the flanks. The skin on the throat, belly, and ventral surfaces of the thighs is coarsely granular, and that on the other ven- tral surfaces is smooth. The tongue is elon- gately cordiform, distinctly notched posterior- ly, barely free behind, and in some specimens shallowly notched anteriorly. The dentiger- ous processes of the prevomers aie widely separated, transverse elevations between the posterior margins of the moderately small o\'oid choanae. Three to five teeth are pres- ent on each elevation. The number of teeth on the maxillary and premaxillary (one side only) varies from 50 to 61. The \'ocal slit extends from the midlateral base of the tongue to the angle of the jaws. The vocal sac is single, median, subgular, and only mod- erately distensible. The general coloration of Plectrohyla ma- tudai is tan or brown with darker brown ir- regular spots or reticulations on the dorsum (pi. 68, fig. 1). In most individuals, the dor- sum is pale brown with dark brown, olive- brown, or black flecks and/or bold reticula- tions on the back. Narrow, dark bars or series of flecks are present on the dorsal sur- faces of the limbs. Most individuals have a narrow dark vertical bar on the upper lip, below the eye. In most individuals, a dark, irregular stripe, which may be fragmented into a series of dashes or flecks, extends from the supratympanic fold nearly to the groin. The stripe or flecks separate the brown dor- sal color from the creamy tan on the flanks. In some individuals, the narrow dark stripe is absent. The axilla usually is gray or bluish gray, and this color usually is narrowly out- lined with black. A few black flecks are pres- ent on the flanks. The anterior surfaces of the thigh's are creamy tan, and the posterior surfaces of the thighs are pale brown. The venter is creamy white, and the vocal sac is grayish brown. The iris is coppery tan with fine black reticulations. In some specimens, minute, metallic green flecks are scattered on the dorsal surfaces. In preservative, the dorsum is dull brown with darker brown or black markings. The anterior and posterior surfaces of the thighs are pale tan or brown, and the venter is creamy tan or pale grayish brown. Tadpoles: Fi\e tadpoles in de\eIopmen- tal stage 25 from Finca La Paz, Departamento San Marcos, Guatemala, have body lengths of 10.5 to 1.3.1 (mean, 11.6) mm. and total lengths of 29.5 to 38.2 (mean, 33.5) mm. A typical tadpole in developmental stage 28 from the same locality has a body length of 13.7 mm. and a total length of 40.5 mm. In dorsal profile, the body is ovoid; the body is slightly wider than deep and not noticeably 562 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 depressed. In dorsal profile the snout is bluntly rounded and in lateral profile, trun- cate. The eyes are small and directed dorso- laterally. The nostrils are situated about mid- way between the eyes and the tip of the snout and are directed anterolaterally. The opening of the sinistral spiracle is about on the midline about midway on the length of the body. The cloacal tube is long and dex- tral. The caudal musculature is heavy and extends nearly to the tip of the rounded tail. The caudal fins are shallow; the dorsal fin is deepest at a point slightly posterior to the midlength of the tail. Throughout its length, the dorsal fin is deeper than the ventral and the dorsal fin does not extend onto the body. At midlength of the tail, the depth of the caudal musculature is much greater than the depth of either fin (fig. 275A). The body is brown, and the caudal muscu- lature is tan with dark reddish brown flecks. Similarly colored flecks are present on the caudal fins. The iris is dull bronze. In pre- servative, the body is dull brown or grayish brown, and the caudal musculature is pinkish tan with l^rown flecks. The mouth is ventral and moderately large; it is ecjual to two-thirds of the width of the body. The mouth is completely bor- dered by a single row of small papillae. Me- dial to these there is an irregular row of larger papillae. The upper beak is broad and barely arched. Moderately long, pointed ser- rations are present on the beak; one serration on either side is greatly enlarged into a fang- like projection. The lower beak is narrow and forms a broad, curved arch; it bears small, pointed serrations. There are two upper and three lower rows of teeth. The upper rows are long and equal in length; the second up- per row is narrowly interrupted medially. The first and second lower rows are equal in length and noticeably shorter than the upper rows, and the third lower row is shorter than the other lower rows (fig. 276A). Hartweg and Orton ( 1941, p. 2) described and illustrated the tadpole of this species under the name "Form a'' Mating Call: Recordings of the call of Plectrohijla matudai are not available, but I have heard this species calling at Finca La Paz, Dcpartamento San Marcos, Guatemala. The call consists of a single note. Taylor and Smith (1945, p. 597) described the call as "a single, sharp note that sounds very much like two pebbles struck together under water. The note is repeated at intervals of about two minutes." Natural History: Plectrohijla matudai inhabits pine-oak forest and primarily cloud forest, where the species ]i\es along small cascading streams. Indi\iduals ha\e been found on \egetation along the streams both by day and night. Taylor and Smith ( loc. cit. ) reported finding the frogs on vegetation and boulders along a stream on Cerro 0\an- do, Chiapas, Mexico; they noted that one male was calling from the water in the stream. All calling males that I have observed were sitting on \egetation. The tadpoles develop in the streams, where the)' characteristically are found ad- hering to boulders in quiet sections of the streams. One recently metamorphosed indi- vidual having a snout-vent length of 17.9 mm. was found on a small herb at the edge of a stream at Finca La Paz on July 30, 1960. Throughout much of its range, Plectrohijla matudai occurs sympatrically with P. f:ag.orum and iiuatemalensis. The latter species usual- h' inhabits the larger mountain streams, whereas matudai and sagorum occur along small streams and rivulets. Rem.\rks: Taylor (1949b, p. 16) named Plectrohijla hrachijccphala on the basis of four specimens from the Sierra Madre in ex- treme eastern Oaxaca, Mexico. In diagnosing his new species, Taylor utilized the following characters: relati\e concealment of the tym- panum, the elevation of the tarsal fold, the relative height of the snout, the relative pus- tularity of the dorsum, and certain charac- teristics of coloration. Bumzahem and Smith (1954, p. 63), reported on a specimen from Cerro Raul, Oa.xaca (U.I.M.N.H. No. 33835), wliich they considered to be intermediate be- tween matudai and brachijcephala. They stated: "On the whole, the specimen seems closer to brachijcephala, but it cannot be re- garded as typical of either form. Further- more, the present specimen was collected in an area intermediate geographical!)- between those to be occupied by matudai and hrachij- ccphala. These facts seem to indicate that 1970 DUELLMAN: HYLID FROGS 563 tlic specimen from Cerro Baiil is most leason- ably interpreted as an intergrade, and that brachycephaUi should, at least until further data is a\ailable. be considered as a subspe- cies of Plcctrohijla matudai." L\ nch and Smith ( 1966, p. 62 ) reported on 35 specimens from Chiapas and Oaxaca. They concluded that Plectrolnjla brachycepli- ala \\'as unrecognizable, and the\' placed tlie name in the synonymy of Plectwhyla matu- dai. Etymology: The specific name is a patro- nym for Eizi Matuda of Chiapas, Mexico. Distribution- : Plectrolnjla matudai oc- curs at elevations of 1000 to 2300 meters on the Pacific slopes of the Sierra Madre from extreme eastern Oaxaca, Mexico, to central Guatemala; this species also is known from the Grijalva depression in western Guatemala and from the Las Nubes block in central Guatemala (fig. 277). See Appendix 1 for the locality records of the 1.39 specimens examined. Plectrohyla ixil Stuart Plcctrohijla ixil Stuart, 1942, p. 4 [holotype, U.M.M.Z. No. 89092 from Finca San Francisco,' 25 kilometers north of Nebaj, El Quiche, Guatemala, ele- \ation 1175 meters; Laurence C. Stuart collector]; 196.3, p. 39. Diagnosis: This small species (40 mm. in snout- vent length) has vocal slits and a long pointed prepollical spine. The dorsum is weakly tuberculate or smooth, and the snout in dorsal profile is acuminate but lacks a \'ertical keel. A broad light lateral stripe, bor- dered below by a dark line separates the dorsal color from that on the \'enter. Plectro- 14° - • P. matudai o P. ixil 0 100 200 KILOMETERS 94' 90' Fig. 277. Distribution ot Plectrohyla matudai and Plectrohyla ixil. 564 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Iii/Ia matudai differs by having a blunt snout, more tuberculate dorsum, and usually by lacking a lateral light stripe, although a dark line or irregular row of spots is present in most specimens. Plectwhyla quecchi and sagorum have a vertical rostral keel, and all other members of the genus are larger and lack vocal slits. Deschiption: Males of this species attain a maximum snout-vent length of 41.6 mm., and females reach 46.5 mm. In a series of 22 adult males from 6.2 kilometers south of Rayon Mescalapa, Chiapas, Mexico, the snout- vent length is .36.9 to 41.6 (mean, 38.9) mm.; the ratio of tibia length to snout-vent length is 0.468 to 0.529 (mean, 0.499); the ratio of foot length to snout-vent length is 0.412 to 0.462 (mean, 0.4.38); the ratio of head length to snout- vent length is 0.286 to 0.369 (mean, 0..332 ) ; the ratio of head width to snout- vent length is 0.331 to 0.395 (mean, 0.355), and the ratio of the diameter of the tympanum to that of the eye is 0.365 to 0.510 (mean, 0.440). These specimens are from the known western extremity of the range, but three in- dividuals having snout-vent lengths of .39, 40, and 40 mm. from Finca San Francisco, El Quiche, Guatemala (the known eastern ex- tremity of the range), are encompassed within the variation exhibited by the specimens from the area of Rayon Mescalapa. The head is nearly as broad as the body, and the top of the head is flat. In dorsal pro- file, the snout is basically truncate at the level of nostrils but sharply pointed terminal- ly; in lateral profile, the snout is truncate. The snout is short; its length is no longer than tile length of the orbit. The nostrils are pro- tuberant, directed dorsolaterally, and situated near the tip of the snout. The canthus is ele- vated and sliarply rounded; the lorcal region is flat, and the lips are moderately thick and bareh' flared. A moderately heavy dermal fold extends posteriorly from the eye, above the tympanum, to a point above the insertion of the arm. The fold obscures the upper edge of the tympanum, whicli otherwise is distinct in most specimens, but in some the tympanic ring is obscured posteriorly. The tympanum is posterior to the ventral border of the eye and is separated from the eye by a distance slightly greater than the diameter of the tym- panum. The arms are short and moderately ro- bust; in breeding males they are hypertro- phied. A row of low tubercles is present on the ventrolateral edge of the forearm and a weak transverse dermal fold is present on the wrist. The fingers are long and slender and bear moderately large discs; the width of the disc on the third fingers is greater than the diameter of the tympanum. The subar- ticular tubercles are large and subconical; in some individuals, the distal tubercle on the fourth finger is bifid. The supernumerary tu- bercles are small and conical; they are irregu- larly arranged in a single row on the proximal segments of the first and fourth fingers and in one or two rows on the proximal segments of the second finger, and usually in two rows on the proximal segment of the third finger. A flattened, bifid palmar tubercle is present. The prepollex is enlarged and cur\ed distally; in some males the sharp prepollical spine pro- trudes from the distal end of the prepollex. 269B). The legs are moderately short and stout; the heels of the adpressed limbs barely o\'crlap. The tibiotarsal articulation extends to the posterior corner of the eye. A few small tubercles are present on the heel, and in some specimens, a faint transverse dermal fold is present on the heel. A distinct, flap- like tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is small, elliptical, and elevated; no distinct outer metatarsal tubercle is present. The toes are modcrateh' long and slender and bear discs that are noticeably smaller than those on the fingers. The subarticular tubercles are mod- erately large and subconical. The supernu- merary tubercles are large, subconical, and arranged in a single row on the proximal segment of each digit. The supernumerary tubercles are also present on the more dis- tal segments of the third and fourth toes. The toes are about three-fourths webbed (fig. 270B). The webbing extends from the penultimate phalanx of the first toe to the base of the penultimate phalanx of the second, from the middle of the penultimate phalanx of the second to the base of the pen- ultimate phalanx of the third, from the mid- dle of the penultimate phalanx of the third to the base of the penultimate phalanx of the fourth and on to the middle of the penul- timate phalanx of the fifth toe. 1970 DUELLMAN: HYLID FROGS 565 The anal opening is directed posteroxen- trally near the midlevel of the thighs. A short, narrow anal sheath is present. A pair of large tubercles is present below the anal opening. The skin on the dorsum is smooth or has a few small scattered tubercles. The skin on the throat, belly, and ventral surfaces of the thighs is strongly granular, whereas that on the other \entral surfaces is smooth. The tongue is ovoid or cordiform; in most individuals a shallow notch is present both anteriorly and posteriorly, but in some speci- mens there is no anterior notch. The tongue is barely free behind. The dentigerous pro- cesses of the pre\omers are short, transverse elevations between the posterior margins of the small, o\oid choanae. There are three to five teeth on each elevation. The number of maxillary and prema.xillary teeth (one side only) varies from 41 to 58. The vocal slits ex- tend from the midlateral base of the tongue to the angles of the jaw. The vocal sac is single, median, subgular, and moderately dis- tensible. The general coloration of Plectrolujia ixil consists of a brown or olive-tan dorsum with a yellowish orange lateral stripe, bordered below by a dark brown line (pi. 68, fig. 2). The dorsum varies from olive-brown to tan or dull greenish gray. In some individuals, there are scattered brown flecks or small spots on the dorsum; there are no distinctive trans- \erse marks on the limbs. The side of the head is darker brown. A dark brown line ex- tends from the nostril to the eye and thence along the supratympanic fold to a point aboxe the insertion of the arm and then posteroven- trally on the flanks towards the groin. A pale creamy yellow or yellowish orange stripe be- gins just posterior to the eye and extends to the groin. A dark brown or black vertical bar usually is present below the eye. The an- terior and posterior surfaces of the thighs are gray or dark grayish brown, and the \enter is pale gray. The vocal sac is dark gray. The iris is deep bronze reticulated with black. In preser\ative, the dorsum is dull gray or brown with or without darker markings. The pale lateral stripe is tan or creamy gray, and the venter is gray. Tadpoles: Six tadpoles in developmental stage 25 from a stream 6.2 kilometers south of Rayon Mescalapa, Chiapas, Mexico, have body lengths of 11.5 to 13.8 (mean, 12.6) mm. and total lengths of 32.4 to 40.7 (mean, 36.1) mm. Three tadpoles in developmental stage 37 from the same locality have body lengths of 15.5 to 16.5 (mean, 16.0) mm. and total lengths of 43.5 to 47.3 (mean, 45.9) mm. In a typical tadpole in developmental stage 37 the body is ovoid in dorsal \iew; the dorsal profile of the snout is rounded; and in lateral profile the snout slopes gradually to its anterior terminus. The eyes are small and directed dorsolaterally. The nostrils are situated about midway between the eyes and the tip of the snout and are directed antero- laterally. The opening of the sinistral spiracle is directed posterodorsally at a point about on the midline slightly posterior to the mid- length of the body. The anal tube is long and dextral. The caudal musculature is ro- bust and extends nearly to the tip of the rounded tail. The caudal fins are low; at midlength of the tail the depth of the caudal musculature is half again the depth of either the dorsal or ventral fins. The dorsal fin does not extend onto the body (fig. 275B). The body is dark brown dorsally and dull gray ventrally. The caudal musculature is pale brown with dark brown blotches, flecks, and reticulations. In preservative, the dorsum is dull brown and the venter is gray. The cau- dal musculature is pinkish tan, and the caudal fins are translucent. The tail is marked by reddish brown blotches and flecks. The mouth is ventral and large; its width is equal to about two-thirds of the greatest width of the body. There is no lateral fold and the mouth is completely bordered by a single row of small papillae. Medial to this fringing row are scattered larger papillae, especially laterally. The beaks are slender. The upper beak is broad and only slightly curved laterally; it bears pointed serrations, three or four of which, on either side are noticeably enlarged. The lower beak is broadly arched and bears fine serrations. There are two upper and three lower rows of teeth. The upper rows are long and sub- equal in length; in most specimens, the sec- ond upper row is narrowly interrupted me- dially. The first and second lower rows are equal in length, but noticeably shorter than 566 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 the upper rows, whereas the third lower row is somewhat shorter than the other lower rows. In some specimens, the first lower row is narrowly interrupted medially (fig. 276B). Stuart (1942, p. 9) described and illus- trated this tadpole under the name of "Form y" Mating Call: The call of Plectrohyla ixil consists of a single note repeated at short intervals. One recording provides the fol- lowing data. The note repetition rate is seven notes per minute and the duration of notes varies from O.IS to 0.26 of a second. There are approximately 200 pulses per second; the fundamental frequency in this poorly modu- lated note is at about 700 cycles per second, and the dominant frequency is at about 2100 cycles per second (pi. 35, fig. 3). Natural History: Plectrohijla ixil inhab- its cloud forests on the Atlantic slopes of the highlands of Chiapas and Guatemala. My ob- servations on this species have been made on the Atlantic slopes in Chiapas along streams at elevations between 1550 and 1690 meters, above the village of Rayon Mescalapa. Males have been observed calling in February, June, and August. Adults have been found on rocks in the streams, both at night and by day. Calling males were observed only on the stems of vegetation overhanging the stream at night. I found one adult in the axil of an elephant car plant by day, and Smith and Brandon (1968, p. 53) reported two individuals in axils of those plants. Meta- morphosing young were obtained in June and August. Seven young having tail-stubs of 3 to 20 mm. had snout-vent lengths of 17.4 to 20.0 (mean, 18.8) mm. A fully transformed juvenile has a snout-vent length of 25.2 mm. Stuart (1942) obtained this species at Finca San Francisco, Departamento El Quiche, Guatemala, on July 31, 1940. At that time, he obtained two juveniles having tail- stubs of about 10 mm. The juveniles have snout-vent lengths of approximately 15.5 mm. Remarks: The general structure of this species indicates a relationship with Plectro- hyla matudai. Furthermore, the presence of enlarged serrations on the upper beak in the tadpoles seems to ally ixil and matudai. In light of their allopatric distribution, it is con- ceivable that they are subspecifically related. However, differences in coloration, tuberosity, and mouthparts of the tadpoles are of suffi- cient magnitude and constancy within each species that it seems better to recognize them as distinct species until evidence of intergra- dation is found. Smith and Brandon (1968, p. 53) dis- cussed specimens of this species from 25 kilo- meters south of Ixhuatan, Chiapas, Mexico, under the name of Plectrohyla matudai; it is evident from their description of the tadpoles that they had specimens of Plectrohyla ixil. Etymology: The specific name refers to the Ixil Indians, a subgroup of the Mame ethnic group, in northern El Quiche, Guate- mala. The "x" is pronounced like "sh"; hence, "e-shel'." Distribution': Plectrohyla ixil occurs at elevations of 1100 to 1700 meters on the At- lantic slopes of the highlands of Chiapas, Mexico, and western Guatemala (fig. 277). See Appendix 1 for the locality records of the 99 specimens examined. Plectrohyla sagorum Hartweg Plectrohijla sagorum Hartweg, 1941, p. 2 [holo- t\pe, U.M.M.Z. No. 88862 from Cerro Ovando, Dis- trito Socomisco, Chiapas, Mexico, elevation 1800 meters; Norman Harhveg collector]. Smith and Tay- lor, 19-J8, p. 73. Stuart, 1963, p. 40. Diagnosis: This moderately small species (51 mm. in snout-vent length) has vocal slits, a long, pointed prepollical spine, a smooth or weakly tuberculate dorsum, and an acuminate snout with a \ertical rostral keel. This com- bination of characters readily separates sa- gorum from all other species in the genus, except Cjuccchi, the only other species with a vertical rostral keel. In quecchi, the snout is blunt, and the dorsum is strongly tubercular; furthermore, in quecchi the flanks are marked with large brown spots, whereas the flanks are marked with small dark flecks in .'/agorum. The other species having \ ocal slits ( ixil and matudai) lack a vertical rostral keel. The re- maining members of the genus are larger (except lacertosa) and lack vocal slits. Description: Males of this species attain a maximum snout-\ent length of 45.5 mm., and females reach 51.9 mm. In a series of 15 males from Volcan Tacana, Chiapas, Mex- ico, and Granja Lorena, Departamento Quet- 1970 DUELLMAN: HYLID FROGS 567 zaltenango, Guatemala, the snout-\cnt length is 33.6 to 45.5 (mean, 39.3) mm.; the ratio of tibia length to snout-vent length is 0.462 to 0.591 (mean, 0.520); the ratio of foot length to snout-\ent length is 0..392 to 0.484 (mean, 0.438); the ratio of head width to snout-\ent length is 0.285 to 0.346 (mean, 0.315); the ratio of head length to snout-vent length is 0.308 to 0.385 (mean, 0.350), and the ratio of the diameter of the t}'mpanum to that of the eye is 0.333 to 0.719 (mean, 0.496). Four females from the same area have snout-vent lengths of 39.6 to 51.9 (mean, 44.6) mm. The head is as wide as, or slightly broader than the body. The top of the head is flat. In dorsal profile, the snout is pointed; a nar- row, \ertical rostral keel is present. In lateral profile, the snout is truncate. The snout is moderately short; the nostrils are barely pro- tuberant and are situated about two-thirds of the distance from the eyes to the tip of the snout. The canthus is ele\atcd and sharp- ly angular; the loreal region is flat, and the lips are thick and barely flared. A moderately heavy dermal fold extends posteriorly from the eye, abo\e the t>'mpanum, to a point above the insertion of the arm. The fold ob- scures the upper edge of the tympanum, which in most specimens is otherwise dis- tinct, and separated from the eye by a dis- tance equal to about two-thirds of the diam- eter of the tympanum. The arms are short and robust; they are hypertrophied in some breeding males. Nu- merous tubercles are present on the ventro- lateral edge of the forearm, and a distinct transverse fold is present on the wrist. The fingers are long and slender and bear moder- ately large discs; the wddth of the disc on the third finger is slightly greater than the diam- eter of the tympanum. The subarticular tu- bercles are moderately large and subconical; in some indi\iduals the distal tubercle on the fourth finger is barely bifid. The supernu- merary tubercles are large and subconical; they are arranged in a single row on the proximal segments of each digit, except near the palm, where additional tubercles are pres- ent. A large, bifid palmar tubercle is present. The prepollex is moderately enlarged and terminally curved. In some indi\'iduals, a prepollical spine protrudes through the termi- nus of the prepollex. The fingers are webbed only basally (fig. 269C). The legs arc mod- erately short and stout; the heels of the ad- pressed limbs overlap by about one-sixth of the length of the shank. The tibiotarsal ar- ticulation extends to the eye. Numerous small tubercles and a faint transverse dermal fold are present on the heel. An elevated tar- sal fold extends the full length of the tarsus. The inner metatarsal tubercle is moderately large, flat, ovoid, and visible from above. Numerous small tubercles arc present on the tarsus so it is not possible to determine if an outer metasarsal tubercles, as such, is present. The toes are moderately long and slender and bear discs that are somewhat smaller than those on the fingers. The subarticular tubercles are moderately small and subconi- cal, and the supernumerary tubercles are moderately large and subconical. The toes are about two-thirds webbed (fig. 270C). The webbing extends from the base of the penultimate phalanx of the first toe to the distal end of the antepenultimate phalanx of the second, from the middle of the penulti- mate phalanx of the second to the distal end of the antepenultimate phalanx of the third, from the middle of the penultimate phalanx of the third to the middle of the antepenulti- mate phalanx of the fourth and on to the base of the penultimate phalanx of the fifth toe. The anal opening is directed posteroven- tralK' near the midlevel of the thighs. A short, narrow anal sheath is present. The anal sheath is bordered above by a transverse fold and below by distinct tubercles. The skin on the dorsum bears many small tubercles; that on the chin, belly, and ventral surfaces of the thighs is strongly granular, whereas the skin on the other ventral surfaces is smooth. The tongue is ovoid, longer than wide, shallowly notched anteriorly and posteriorly, and barely free behind. The dentigerous processes of the prevomers are small, rounded, widely separated elevations between the moderately large rounded choanae. There are three or four teeth on each elevation. The number of maxillary and premaxillary teeth (one side only) varies from 34 to 45. The vocal slits extend from the midlateral base of the tongue to the angles of the jaws. The vocal sac is 568 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 single, median, subgular, and only moderate- ly distensible. The general coloration of Plectrolujhi sa- gorum is dull brown with small irregular, slightly darker brown spots on the dorsal surfaces (pi. 68, fig. 3). The flanks are tan and are marked by fine dark brown reticula- tions or brown flecks. In some individuals, small cream spots are also present on the flanks. The posterior surfaces of the thighs are dull dark brown, and the belly is gray. The vocal sac is dark grayish brown. The iris is deep bronze with fine black reticulations. In preservative, the dorsum varies from grayish brown to dull brown with faint or distinct dark brown flecks on the body and limbs. The flanks usually are somewhat paler and marked by numerous brown flecks or small spots. The venter is dull creamy tan or pale gray. Tadpoles: A typical tadpole in develop- mental stage 32 has a body length of 13.2 mm. and a total length of 36.9 mm. The body is ovoid, no wider than deep. In dorsal and lateral profiles, the snout is rounded. The eyes are small and directed dorsolaterally. The nostrils are directed anterolaterally at a point slightly closer to the eyes than to the tip of the snout. The opening of the sinistral spiracle is on the midline slightly posterior to the midlength of the body. The anal tube is moderately long and de.xtral. The caudal musculature is robust and extends nearly to the tip of the rounded tail. The fins are nar- row; at midlength of the tail, the depth of the caudal musculature is greater than the depth of either the ventral or dorsal fins; the dorsal fin is shallower than the ventral one and does not e.xtend onto the body (fig. 275C). The body is dark gray brown with some faint darker mottling. The caudal muscula- ture is paler brown and the fins are transpar- ent. The tail is heavily spotted with dark gray. In preservative, the body and caudal musculature is dark brown with darker brown flecks and spots on the caudal musculature and fins. The mouth is ventral; its width is equal to about two-thirds of the greatest width of the body. The mouth lacks lateral folds and is completely bordered by a single row of small papillae. A row of larger papillae is present medial to the fringing row; a few small papillae are present lateral to the beaks. The beaks are well developed and bear long, pointed serrations of equal length. The upper beak is in the form of a broad arch with moderately robust lateral processes; the ven- tral beak is massive and V-shaped. There are two upper and three lower rows of teeth. The upper rows are long and equal in length, and the second upper row is narrowly interrupted medially. The lower rows are somewhat shorter than the upper ones, equal in length, and complete (fig. 276C). This is the tadpole described by Hartweg and Orton (1941, p. 5) as "Form h!' M,\TiNC Call: Recordings of the call of Plectrohyla sagorum are not available. Tay- lor and Smith (1945, p. 598) described the call of this species as a "slightly drawn out, coarsely trilled, nasal qtiaaack." Natural History: Plectrohyla sagorum inhabits cloud forest, where it breeds at night in cascading mountain streams and spends the davs in bromeliads. Taylor and Smith ( 1945, p. 597 ) noted that in April, 1940, both adults and juveniles were found in brome- liads on Cerro Ovando, and that males were calling from bromeliads by day. At Granja Lorena, Guatemala, on July 21, 1966, males were calling from low branches of bushes along a small stream at night. Tadpoles were found in gra\el-bottomed pools in the streams. Tadpoles were found in similar habi- tats at 10.4 kilometers west-southwest of San Martin Sacatepcquez, Guatemala, on July 30, 1960 and on February 19, 1961. Adults were found in bromeliads on Volcan Tacana, Chia- pas, on August IS, 1965. These obser\ations indicate that calling apparently takes place throughout the year and probably breeding also takes place throughout the year. Fur- thermore, this species, perhaps more than any other Plectrohyla, utilizes bromeliads as day- time retreats. Renlxrks: Plectrohyla sagorum is known to occur s\ mpatricalK' with at least four other members of the genus {avia, matuclai, guate- malensis, and hartwegi). Of these, the spe- cies seems to be ecologically most like ma- tuclai, which also inhabits small streams. The other species tend to inhabit the larger 1970 DUELLMAN: HYLID FROGS 569 streams, although at Granja Lorena, Guate- mala, both avia and guatemalensis were found along the same small stream with sagorum. Etymology: The specific name is derived from the Latin saga meaning soothsayer; Hartweg (1941, p. 2) proposed the name "in memory of the few witchcraft-practicing In- dians who inhabit that eerie mountain fCerro 0\ando, Chiapas]." Distribution: Plectwhyla sagonim oc- curs at elevations of 1500 to 2050 meters on the Pacific slopes of the Sierra Madre from south-central Chiapas, Mexico, southeastward to north-central El Salvador (fig. 278). See Appendix 1 for the locality records of the 94 specimens examined. Plectrohyla quecchi Stuart Picctrohyla quecchi Stuart, 1942, p. 1 [holot>-pe, U.M.M.Z, No. 89086 from Barranca Las Palmas, 2 kilometers north of Finca Los Alpes (43 kilometers east and slightly south of Cohan, Alta Verapaz, Guate- mala, elevation 1015 meters; Laurence C. Stuart col- lector]; 1963, p. 40. Diagnosis: This small species (47 mm. in snout- vent length) has vocal slits, a long, pointed prepollical spine, a tuberculate dor- sum, and a blunt snout with a vertical rostral keel. This combination of characters readily separates quecchi from all other species in the genus, except sagorum, which also has a vertical rostral keel. The latter species has an acuminate snout and a smooth or weakly tuberculate dorsum. In quecchi, large brown spots are present on the flanks, whereas in sagorum the flanks are marked with small dark flecks. The other species having vocal slits {ixil and matudai) lack a vertical rostral keel. The remaining members of the genus are larger and lack vocal slits. 94' 90« >u .' '^ i '< f 1 1 ■ i / J — . _ . —■^■'' i i i 1 ^ ! \ \ ( \ j <^^ cd f \ y ^\ / ■" T^s^^v^^ / < i. • • O O O \/ f 1 • P sagorum - o P quecchi -- ( c c 100 200 1 KILOMETERS 1 16°- 14° - 16' 14' 94° 90« Fig. 278. Distribution of Plectrohtjla sagorum and Plectrohyla quecchi. 570 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Description: Males of this species attain a maximum snout-vent length of 44.0 mm., and females reach 46.7 mm. In a series of eight adult males, from Finca Los Alpes, De- partamento Alta Verapaz, Guatemala, the snout-\'cnt length is 40.4 to 4.3.8 (mean, 42.2) mm.; the ratio of tibia length to snout-vent length is 0.514 to 0.554 (mean, 0.531); the ratio of foot length to snout-vent length is 0.460 to 0.493 (mean, 0.480); the ratio of head length to snout-vent length is 0.294 to 0.338 (mean, 0.318); the ratio'of head width to snout-vent length is 0.342 to 0.390 (mean, 0..368), and the ratio of the diameter of the tympanum to that of the eye is 0.346 to 0.463 (mean, 0.411). The single known female from the same locality has a snout-vent length of 46.7 mm. and does not differ from the males in proportions. The head is as wide as the body, and the top of the head is flat. In dorsal profile, the snout is bluntly rounded; in lateral profile, it is truncate. A narrow, vertical rostral keel is evident on the dorsal part of the snout. The snout is short; its length is slightly less than the diameter of the eye. The nostrils are nearly terminal and are directed dorsolat- erally. The canthus is elevated and slightly angular. The loreal region is barely concave and the lips are thick and barely flared. A moderately heavy dermal fold extends pos- teriorly from the eye, above the tympanum, to a point above the insertion of the arm. The fold obscures the upper edge of the tympan- um, which in some specimens is distinct and separated from the eye by a distance slightly less than the diameter of the tympanum, whereas in other specimens, the tympanum is barely evident. The arms are short and robust, especially in breeding males. A row of small tubercles is present on the ventrolateral edge of the forearm, and a distinct transverse dermal fold is present on the wrist. The fingers are long and slender and bear moderately large discs; the width of the disc on the third finger is greater than the diameter of the tympanum. The subarticular tubercles are moderatcK' large and subconical; none is noticeably bifid. The supernumerary tubercles are moderately large and subconical; they are present on the proximal segments of each digit and arranged in a single row on the distal part of the seg- ments, but in some individuals irregularly ar- ranged proximally. The flat, bifid palmar tubercle is present. The prepollex is moder- ately enlarged and distally curved; in some males the tip of the prepollical spine pro- trudes through the distal end of the prepollex. The webbing on the hands is vestigial (fig. 269D). The hind limbs are moderately short and stout; the heels of the adpressed limbs overlap by about one-fifth of the length of the shank. The tibiotarsal articulation ex- tends to the eye. A few small tubercles and a distinct transverse dermal fold is present on the heel. A distinct tarsal fold extends the full length of the tarsus. The inner metatar- sal tubercle is long, elliptical, flat, and visible from above. A small conical outer metatarsal tubercle is present. The toes are long and slender and bear discs that are somewhat smaller than those on the fingers. The sub- articular tubercles are moderately large and subconical, and the supernumerary tubercles are large and conical; they are arranged in a single row on the proximal segments of each digit. The toes are about two-thirds webbed (fig. 270D). The webbing extends from the middle of the penultimate phalanx of the first toe to the distal end of the antepenulti- mate phalanx of the second, from the distal end of the penultimate phalanx of the second to the middle of the antepenultimate phalanx of the third, from the middle of the antepen- ultimate phalanx of the third to the distal end of the antepenultimate phalanx of the fourth and on to the middle of the penulti- mate phalanx of the fifth toe. The anal opening is directed posteroven- trally near the midlevel of the thighs. A short, narrow anal sheath is present. A transverse dermal fold exists above the anal sheath, and vertical dermal folds are present below the anal opening. The skin on the dorsum is tuberculatc and that on the throat, belly, and ventral surfaces of the thighs is strongly granular, whereas the skin on the other ven- tral surfaces is smooth. The tongue is ovoid, longer than wide, shallowly notched anterior- ly and posteriorly, and barely free behind. The dentigerous processes of the prevomers are small, elliptical, widely separated eleva- tions between the moderately large ovoid, 1970 DUELLMAN: HYLID FROGS 571 choanae. Each elevation bears three or four teeth. The number of premaxillary and max- i]h\r\' teetli varies from 47 to 53. The vocal slits extend from the midlatcral base of the tongue to the angles of the jaws. The vocal sac is single, median, subgular and moderate- ly distensible. The general coloration of Plecfrohtjla quecchi is pale tan, olive-tan, or pale grayish bro\\'n above \\'ith dark brown spots on the somewhat paler flanks (pi. 68, fig. 4). A few faint darker spots are present on the dorsum. The posterior surfaces of the thighs are tan or pale brown. The belly is grayish with gray or brown suffusion or mottling and the vocal sac is dull olive-green with cream spots. The iris is deep bronze with fine black reticu- lations. In preservative, the dorsum varies from dull gra\ish tan to dark brown with or with- out faint darker flecks or reticulations. The flanks are marked by bold brown spots, and the venter is mottled with gray or brown, especially on the chest. Tadpoles: A typical tadpole in develop- mental stage .34 has a body length of 15.0 mm. and a total length of 42.0 mm. The body is ovoid, no wider than deep. The snout is bluntly rounded in dorsal and lateral profiles. The eyes are moderately small and directed dorsolaterally; the nostrils are situated about miduay between the eyes and the tip of the snout and are directed anterolaterally. The opening of the sinistral spiracle is about on the midline about midlength of the body. The cloacal tube is long and dextral. The caudal musculature is heavy and extends nearly to the tip of the rounded tail. The fins are shallow, and the dorsal fin does not extend onto the body. At midlength of the tail, the caudal musculature is deeper than either the dorsal or ventral fins, which are of ap- proximately equal depth throughout their length (fig! 275D). In preservative, the body is pale brown; the caudal musculature is tan, and the caudal fins are transparent. Large dark brown blotches are present on the tail. The mouth is ventral; its width is equal to about two-thirds of the greatest width of the body. There is no lateral fold. The mouth is completely bordered by a single row of small papillae; numerous larger pa- pillae are present medially to the fringing papillae. The beaks are well developed and possess long, pointed serrations of ecjual length. The upper beak is in the form of a broad arch and lacks noticeable lateral pro- cesses; the lower beak is moderately robust and broadly V-shaped. There are two upper and three lower rows of teeth. The upper ro\\'s are long and equal in length, and the second upper row is narrowly interrupted medially. The lower rows are shorter than the upper ones; the first and second lower rows arc ecjual in length, whereas the third is shorter (fig. 276D). This is the tadpole described as "Form z" by Stuart (1942, p. 10). Mating Call: Recordings of the call of this species are not available, Stuart (1942, p. 4) stated that the call is a "harsh quack repeated at rather long intervals." Natural History: Plectrohyla quecchi is an inhabitant of cloud forest. The only ob- servation of adults was made at Finca Los Alpes, Departamento Alta Verapaz, Guate- mala. There in February, 1940, Stuart found the adults between boulders and pebbles in the water in a mountain stream. I obtained adults there at night in July, 1961. The frogs were found on bushes and vines overhanging a stream at night, but one male was obtained from a vine by day. The tadpoles have been taken from gravel- bottomed pools in streams. Remarks: This species obviously is closely related to P. sagorum, from which it differs principally in the shape of the snout and in the amount of dark pigmentation on the flanks. The two species seem to represent vicarious populations of a formerly more widespread stock. Etymology: The specific name refers to the Quecchi Indians of Alta Verapaz, Guate- mala. The name is pronounced "kek-chi'." Distribution: Plectrohyla quecchi is known from elevations of 1000 to 1600 meters on the slopes of the Atlantic highlands in central Guatemala (fig. 278). See Appendix 1 for the locality records for the 22 specimens examined. 572 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Plectrohyla glandiilosa (Boulengei) Hyla glandulo.sa Boulenger, 1883, p. 164 [syntvpes, B.M.X.H. Nos. 1947.2.20.40 and 41 from "Guate- mala"; presented by Frederick D. Godman]. Giinther, 1901 (1885-1902), p. 281. Duellman, 1964c, p. 455 [synonymized Plectrohyla cotzicensis Stuart, 1948a, with Hijla glandulosa Boulenger, 1883]. Plectrohyla cotzicensis Stuart, 1948a, p. 17 [holo- type, U.M.M.Z. No. 95902 from the source of the Rio Cuilco, on the slopes of Cerro Cotzic, 2 kilometers northwest of Ixchiguan, Departamento San Marcos, Guatemala; Laurence C. Stuart collector]. Stuart, 1963, p. 39. Diagnosis: This moderate-sized species (50 mm. snout-vent length) has a smooth or weakly tuberculate dorsum. The prepolhcal process is short, flat, and blunt, and the dor- sum is mottled gray and dull green. Males lack \'ocal slits. Of the other species lacking vocal slits, guatemalensis and harticegi each differs by having a bifid prepollical spine and a tuberculate dorsum; avia has a long, pointed prepollical spine and a short, blunt snout, whereas pycnochila has a short, flat, and blunt prepollical spine like that in glandulosa but differs by having a tuberculate dorsum and a round snout. PlectroJiyla lacertosa has an elongate, round, terminally blunt prepollical spine, a short snout, and a brown dorsum. Other species of Plectrohyla have vocal slits in males and curved, terminally pointed pre- pollical spines. Description: Males of this moderately large species attain a snout-vent length of 49.1 mm., and females reach 49.7 mm. In a series of 12 males from 8 kilometers south of Pac^uix, Departamento Huehuetenango, Guatemala, the snout-vent length is 42.2 to 49.1 (mean, 44.6) mm.; the ratio of tibia length to snout-vent length is 0.471 to 0.543 (mean, 0.513); the ratio of foot length to snout-vent length is 0.462 to 0.524 (mean, 0.480); the ratio of head length to snout-vent length is 0.274 to 0.321 (mean, 0.296); the ratio of head width to snout-vent length is 0.323 to 0.384 (mean, 0.352), and the ratio of the diameter of the tympanum to that of the eve is 0.255 to 0.537 (mean, 0.378). Five females from the same locality have snout-vent lengths of 39.3 to 49.7 (mean, 44.3) mm. They do not differ significantly from the males in propor- tions except that the ratio of the diameter of the tympanum to that of the eye is 0.356 to 0.558 (mean, 0.431). The head is slightly narrower than the body, and the top of the head is flat. The snout in dorsal profile is acuminate; in lateral profile it is truncate. The snout is short, its length is about equal to the diameter of the eye. The nostrils are barely protuberant and are situated at a point about three-fourths of the distance from the eyes to the tip of the snout. The canthus is barely elevated and moderately angular. In breeding males, the loreal region is flat, and the lips are thick and swollen. In females, and subadult males, the loreal region is slightly concave, and the lips are moderately thick and barely flared. A moderately heavy dermal fold extends pos- teriorly from the eye, above the tympanum, and downward to a point above the insertion of the arm. The fold obscures the upper edge of the tympanum, which otherwise is barely evident and is separated from the eye by a distance slightly greater than the diameter of the tympanum. The arms are moderately long, slender in females, and robust in males. A row of low tubercles is present on the ventrolateral edge of the forearm, and a distinct transverse der- mal fold is present on the wrist. The fingers are moderately long and slender and bear large discs; the width of the disc on the third finger is noticeably greater than the diameter of the tympanum. The subarticular tubercles are moderately large, round, and flattened; none is bifid. The supernumerary tubercles are small and subconical; they are arranged in a single row on the proximal segments of each digit. A small, diffuse palmar tubercle is present. The prepollex is enlarged and rjuadrangular. The webbing on the hands is vestigial (fig. 271C). The legs are moder- ately long and robust; the heels of the ad- pressed limbs overlap by about one-third of the length of the shank. The tibiotarsal ar- ticulation extends to the eye. A few small tubercles and a transverse dermal fold are present on the heel. In all specimens a dis- tinct inner tarsal fold extends the full length of the tarsus. In specimens from the south- western highlands of Guatemala, there is a distinct outer tarsal fold. In specimens from the highlands of central and southeastern Guatemala, the outer tarsal fold is weak, and in most specimens from the Sierra de los 1970 DUELLMAN: HYLID FROGS 573 Ciichumatanes in northwestern Guatemala, the outer tarsal fold is absent or represented by a row of indistinct tubercles. The inner metatarsal tubercle is elliptical and flat. No outer metatarsal tubercle, as such, exists. The toes are moderately long and slender and bear discs that are somewhat smaller than those on the fingers. The subarticular tuber- cles are small and round, and the supernu- merary tubercles are moderately small and conical. The toes are about two-thirds webbed (fig. 272A). The webbing extends from the middle of the penultimate phalanx of the first toe to the base of the penultimate phalanx of the second, from the middle of the penultimate phalanx of the second to the middle of the penultimate phalanx of the third, from the base of the penultimate pha- lanx of the third to the base of the antepen- ultimate phalanx of the fourth, and from the middle of the antepenultimate phalanx of the fourth to the middle of the penultimate pha- lanx of the fifth toe. The anal opening is directed posteroven- trally near the midlevel of the thighs. A short anal sheath is present, but a distinct trans- verse anal flap and anal tubercles are present. The skin on the dorsum is weakly or moder- ately tubercular; that on the flanks is smooth or possesses a few small scattered tubercles. The skin on the throat, belly, and ventral sur- faces of the thighs is granular, and that on the ventral surfaces of the shanks is smooth. The tongue is nearly round, shallowly notched anteriorly and posteriorly, and barely free behind. The dentigerous processes of the prevomers are small, widely separated, trans- verse ridges between the posterior margins of the moderately small, ovoid choanae. There are one to three long, pointed teeth on each ridge. The number of maxillary and premax- illary teeth (one side only) varies from 23 to 30. Vocal slits and a vocal sac are absent. The general coloration of adult males is usually green or dull olive-green above with irregular olive-brown or dark brown mark- ings (pi. 69, fig. 1). Some individuals are dull olive-brown or grayish brown above with faint darker brown markings. The posterior surfaces of the thighs and webbing of the feet are gray, and the venter is grayish white. Adult females usually are nearly uniform green above (pi. 69, fig. 2). In many individ- uals, a distinct brown or tan stripe is present on the canthus and supratympanic fold. The flanks are creamy tan or pinkish tan, and the venter is creamy white. The posterior sur- faces of the thighs and the webbing is pale tan. There is a creamy white line on the outer edge of the tarsus and a transverse white line above the anal opening. In both sexes, the iris is bronze flecked with black. The intensity of the dorsal pigmentation is subject to change. Some females change to dark olive-brown or gray, whereas some males that were rather pale green with olive- brown markings change to dark olive-brown with dark brown markings. Juveniles are pale green above and have a white line on the tarsus and above the anus. The throat and belly are white and the ventral surfaces of the limbs and the webbing are yellow. In preservative, the dorsum is dull grayish tan or dull brown with darker grayish brown markings. The posterior surfaces of the thighs are grayish tan, and the venter is dirty creamy white. Tadpoles: Large series of tadpoles were obtained from various localities in the Sierra de Los Cuchumatanes in Guatemala. Tad- poles in developmental stage 25 apparently undergo a considerable amount of growth. In a series of 31 specimens in that develop- mental stage the body length varies from 8.6 to 13.0 (mean, 11.1) mm., and the total length varies from 20.5 to 31.5 (mean, 26.5) mm. The largest tadpole examined was in develop- mental stage 37 and had a body length of 21.7 mm. and a total length of 56.8 mm. A typical tadpole in developmental stage 28 has a body length of 17.5 mm., and a total length of 45.0 mm. The body is rather elon- gate and bluntly rounded anteriorly and pos- teriorly. The body is as wide as deep. In dorsal profile, the snout is bluntly rounded, and in lateral profile, it is somewhat more sharply rounded. The eyes are widely sepa- rated, small, and directed dorsolaterals. The nostrils are directed anterolaterally at a point about midway between the eyes and the tip of the snout. The opening of the sinistral spiracle is about on the midline at approxi- mately midlength of the body. The anal tube is moderately long and dextral. The caudal 574 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 musculature is robust and extends nearly to the tip of the rounded tail. At midlength of the tail the depth of the caudal musculature is slightly more than the depth of either the dorsal or ventral fins. The dorsal fin does not extend onto the body (fig. 275E). The body is brown with greenish gold lichenous markings laterally and ventrally. The tail is tan with dark brown flecks and blotches. The iris is pale bronze. In preserva- tive, the body is dark grayish brown, and the caudal musculature is creamy tan. The caudal fins are translucent, and the entire tail is marked with dark brown flecks or small blotches. The mouth is ventral and moderately large; its width is equal to about three-fifths of the greatest width of the body. The mouth lacks a lateral fold, but is completely bor- dered by one row of small papillae. Medial to the fringing row is a row of larger papillae on the anterior and posterior lip. There are no papillae lateral to the beak. The upper beak is moderately slender, lacks lateral pro- cesses, and bears short, pointed serrations that are of approximate equal length. The lower beak is massive, broadly V-shaped, and bears short pointed serrations. There are two upper and three lower rows of teeth. The upper rows are long and about equal in length; the second upper row is narrowly in- terrupted medially. The lower rows are com- plete; the first and second lower rows are equal in length but shorter than the other rows, and the third lower row is noticeably shorter than the others (fig. 276E). The characteristics of these tadpoles agree with those described for this species by Stuart (1951, p. 51). As noted by Stuart, small speci- mens (those less than 10 mm. in develop- mental stage 25) do not have the mouthparts fully developed. The keratinization of the beaks is incomplete, and the formation of the rows of teeth is incomplete. Apparently, the third lower row is the last to develop. Mating Call: The absence of vocal slits and a vocal sac preclude the presence of a voice in this species. Natural History: Plectroliijla gJamlulosa inhabits the pine-cypress forest, fir forest, and montane meadows at high elevations in Gua- temala. Adults are usually found along small rivulets. Stuart (1948a, p. 18) reported finding adults "beneath rocks and clumps of sod in shallow tricklets emerging from springs in the flanks of Cerro Cotzic" [2 kilometers northwest of Ixchiguan, San Marcos, Guate- mala]. In July, 1960, and in March, 1966. I found adults sitting on rocks under banks and small cascades in a small stream in a montane meadow 8 kilometers south of Paquix in the Sierra de Los Cuchumatanes, Guatemala. A few individuals were observed sunning on rocks or on bunch grass in and along the stream. The tadpoles usually are foimd in quiet pools in streams or adhering to the lee-side of rocks in the streams. Stuart (1951, p. 52) stated: "Both tadpoles and adults have been taken in tiny rivulets in the pine-cypress zone and in quiet spring-fed pools, where this spe- cies is associated with Bufo bocourti. Thus, though apparently adapted to life in the swift mountain stream, the species can and does invade the lenitic environment." I obtained both tadpoles and metamorphosing young from the shallow Laguna de Vejcha at an elevation of 3040 meters in the Sierra de Los Cuchumatanes. At that locality the tadpoles were in shallow water having a temperature of 16.5 degrees centigrade. Stuart (1951, p. 51) stated: "This species apparently has an extended breeding season, a condition which seems to obtain in most of the stream salientians of Guatemala. Fe- males with eggs apparently ready for deposi- tion were secured at Ixchiguan on April 23, and one in same condition was taken on Maria Tucum on August 4. Tadpoles as small as 6 mm. and at the transformation stage were secured during early and mid- April at Ixchiguan, while transformed juve- niles were taken on Maria Tucum on August 4." My observations corroborate those of Stuart; tadpoles in various stages of develop- ment and metamorphosing young were ob- tained in the Sierra de Los Cuchumatanes in July, 1960, and in March, 1966. The tadpoles of Plectrolujla glamlulosa develop in extreme- ly cold water; consequently, it is highly prob- able that the duration of the tadpole stage is lengthy and possibly requires more than one year. At lower elevations, such as at Soledad 1970 DUELLMAN: HYLID FROGS 575 Grande, Departamento Japala, Guatemala ( ele\'ation 2500 meters ) , this species has been found in bromeliads. Stuart (1954c, p. 48) re- ported finding 10 juveniles and one adult fe- male with eggs in bromeliads. The habits of the frogs of sitting on rocks or clumps of bunch grass in the sun is unique among Middle American hylids. A well- developed melanin layer in the skin appar- ently protects the animal against the effects of solar radiation. Remarks: This species is best known un- der the name of Plectwhyh cotzicemis Stuart, 194Sa. Duellman (1964c, p. 455) resurrected Boulenger's Hyla glandulosa for this species. Etymology: The specific name is Latin, meaning glandular, and refers to the thick glandular condition of the skin. Distribution: Plectrohijla glandulosa oc- curs at elevations from 2400 to 3500 meters in the highlands of Guatemala and adjacent El Salvador (fig. 279). See Appendix 1 for the locality records of the 192 specimens examined. Plectrohyla pycnochila Rabb Plectrohijla pycnochila Rabb, 1959, p. 45 [holo- type, A.M.N.H. No. 62667 from "Coyanie, Veracruz, Mexico"; Byron Harrell collector]. Diagnosis: This moderate-sized species (60 mm. in snout-vent length) has a tubercu- late dorsum and lacks vocal slits. The snout is blunt, and the prepollical process is short, flat, and blunt. The only other species hav- ing a short, flat, blunt prepollical process is glandidosa, which has a smooth dorsum and an acuminate snout. Plectrohijla lacertosa has an elongate, round, terminally blunt prepolli- 94° 90° ^ ' '. /"" ""> ' I ..._i / " \ L.. ^^J' 1 ^ ■ > <■■-■■' S 1 j < <^:^ cnJ i 16° 1 ^^^^ . 1 16° ^o V • •• o • / { / t • P glandulosa 14° - 0 P. avia ■ P. pycnochila A P lacertosa "N f 1 ~-_..'-~'vC i 14° 0 100 200 KILOMETERS 1 94° 90° Fig. 279. Distribution of Plectrohijla lacertosa, P. glandulosa, P. pycnochila, and P. avia. 576 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 cal spine and an acuminate snout. The other members of the genus that lack vocal slits have bifid prepollical spines {guatemalensis and hartwegi) or a long, pointed prepollical spine (avia). The remaining four species (ixil, matudai, quecchi, and sagorum) are smaller (less than 50 mm. in snout-vent length) and have vocal slits and long, pointed prepollical spines. Description: This species is known from two adult males having snout-vent lengths of 52.5 and 60.5 (mean, 56.5) mm. The ratio of tibia length to snout-vent length is 0.502 to 0.581 (mean, 0.542); the ratio of foot length to snout-vent length is 0.463 to 0.530 (mean, 0.497); the ratio of head length to snout-vent length is 0.288 in both; the ratio of head width to snout-vent length is 0.332 to 0.347 (mean, 0.340), and the ratio of the diameter of the tympanum to that of the eye is 0.464 to 0.484 (mean, 0.474). The head is slightly broader than the body, and the top of the head is flat. In dor- sal profile, the upper part of the snout is truncate, whereas the border of the lips is round; in lateral profile, the snout slopes abruptly from the snout to the lips. The snout is short; the nostrils are slightly pro- tuberant and situated at a point about three- fourths of the distance from the eyes to the tip of the snout. The canthus is slightly ele- vated and angular; the loreal region is barely concave, and the lips are thick and barely flared. A heavy dermal fold extends posterior- ly from the eye, above the tympanum, and to a point above the insertion of the arm. The fold obscures the upper edge of the tym- panum, which otherwise is distinct and sepa- rated from the eye by a distance slightly greater than the diameter of the tympanum. The arms are moderately long and robust. A row of low tubercles is present on the ven- trolateral edge of the forearm, and a distinct transverse dermal fold is present on the wrist. The fingers are long and slender and bear moderately large discs; the width of the disc on the third finger is equal to the diameter of the tympanum. The subarticular tubercles are moderately large and subconical; none is bifid. The supernumerary tubercles are small and conical; they are present in a single row on the proximal segments of each digit ex- cept basally on the third and fourth digits where there are additional tubercles. The palmar tubercle is elevated and bifid. The prepollex is rectangular. The fingers are about one-half webbed (fig. 271A). The webbing is vestigial between the first and second fingers and connects the second finger from the base of the penultimate phalanx to the middle of the antepenultimate phalanx of the third and on to the distal end of the antepenultimate phalanx of the fourth finger. The hind limbs are moderately long and slender; the heels of the adpressed limbs overlap by about one- fourth of the length of the shank. The tibio- tarsal articulation extends to the anterior corner of the eye. A distinct transverse der- mal fold is present on the heel, and a low tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is mod- erately small and elevated. A minute, coni- cal outer metatarsal tubercle is present. The toes are long and slender and bear discs that are only slightly smaller than those on the fingers. The subarticular tubercles are mod- erately small and subconical, and the super- numerary tubercles are small, subconical, and arranged in a single row on each digit. The toes are about three-fourths webbed (fig. 272C). The webbing extends from the middle of the penultimate phalanx of the first toe to the base of the penultimate phalanx of the second, from the base of the disc of the sec- ond to the base of the penultimate phalanx of the third, from the base of the disc of the third to the base of the antepenultimate phalanx of the fourth and on to the middle of the penultimate phalanx of the fifth toe. The anal opening is directed posteroven- trally at the midlevel of the thighs. A short anal sheath is present, and a transverse der- mal fold is present above the anal sheath. The skin on the dorsal surfaces is tubercular. The tubercles are small and scattered but distinct. The skin on the throat and chest in one individual is smooth and weakly granular in the other; in both specimens, the skin on the belly and ventral surfaces of the thighs is granular, whereas that on the ventral sur- faces of the rest of the limbs is smooth. The tongue is nearly round, shallowly notched anteriorly and posteriorly, and barely free be- hind. The dentigerous processes of the pre- 1970 DUELLMAN: HYLID FROGS 577 vomers are narrowly separated, curved ele- vations between the small round choanae. There are three to five teeth on each process. The number of maxillary and premaxillary teeth (one side only) \aries from 31 to 36. There are no vocal slits or a vocal sac. The coloration in life is unknown. In pre- servative, the dorsum is dark gray or grayish brown with a few irregular and scattered bluish tan flecks (pi. 5, fig. 1). The ventral surfaces of the forelimbs and throat are cream\' white; the other ventral surfaces are bluish gray. The axilla is white. T.A.DPOLES: The tadpoles of this species are unknown. M.\TiNG C.\LL: The absence of \ocal slits and a vocal sac precludes the presence of a voice in this species. Natural History: The single specimen of Plectrohijla pycnochila having locality data was obtained at 8 kilometers north-northwest of San Cristobol de las Casas, Chiapas, Mex- ico. Dr. Dilford G. Carter, the collector, ob- tained the frog in a cave in pine-oak forest on a slope well above a small stream. Remarks: Rabb (1959, p. 45), named Plectrohyla pycnochiki on the basis of a single male supposedly collected near Coyame, Veracruz, Mexico, in July, 1954, by Byron Harrell. Subsec^uent collecting in the Los Tuxtlas, that volcanic mountain range in which Coyame is located, failed to reveal the presence of Plectrohyla there. There are few streams in Los Tuxtlas, and the only hylid tadpole that has been found in these streams are those of Hyla miotympanum. Dr. Byron Harrell has informed me that he is not certain that he obtained the type specimen of pycno- chila in Los Tuxtlas. He intimated that the specimen may have originated in the high- lands of central Chiapas. Et^'mology: The specific name is derived from the Greek pyknos, meaning thick, and the Greek cheilos, meaning lip, and refers to the characteristically thick lips found in breeding males of Plectrohyla. Distribution: Plectrohyla pycnochila oc- curs in the highlands of central Chiapas, Mexico; the only definite record is from an elevation of 2400 meters (fig. 279). See Appendix 1 for the locality records of the t\vo specimens examined. Plectrohyla lacertosa Bumzahem and Smith Plectrohyla lacertosa Bumzahem and Smith, 1954, p. 64 [holotype, U. I.M.N. H. No. 33693 from "Region de Socomisco," Chiapas, Mexico; Eizi Matuda collec- tor]. Diagnosis: This moderately small species (47 mm. in snout-vent length) has a smooth dorsum and an elongate, round, terminally blunt prepollical spine; vocal slits are absent. All other species oi Plectrohyla have a pointed or bifid prepollical spine, except glandulosa and pycnochila, which have a relatively short, flat, terminally blunt prepollical process. Plectrohyla lacertosa differs from pycnochila by having a smooth instead of tuberculate dorsum, and from glandulosa by being brown instead of mottled gray and green, and by having a completely concealed tympanum in- stead of a partly covered one. Description: This species is known solely from one adult male having a snout-vent length of 47.8 mm. The ratio of tibia length to snout-vent length is 0.494; the ratio of foot length to snout-vent length is 0.460; the ratio of the head length to snout-vent length is 0.310, and the ratio of head width to snout- vent length is 0.366. The head is as broad as the body, and the top of the head is flat. In dorsal profile, the tip of the snout is pointed, but the leading edges of the lips are round. In lateral profile, the snout is truncate. The snout is moderate- ly short; the nostrils are not protuberant and are situated at a point about two-thirds of the distance from the eyes to the tip of the snout. The canthus is barely evident and rounded; the loreal region is flat and the lips are grotesquely swollen. A thin dermal fold extends posteriorly from the eye to a point above the insertion of the arm. The tympan- um is not visible. The arms are short and greatly hypertro- phied. A dermal fold extends along the ven- trolateral edge of the forearm and an im- mensely heavy dermal is present on the wrist. The fingers are moderately long and slender and bear rather small discs. The subarticular tubercles are moderately small and subconi- cal. No supernumerary tubercles are evident. A flat, seemingly tripartite palmar tubercle is present. The prepollex is much elongated into a terminally blunt process, which is cov- 578 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 ered by a horny nuptial excrescence. The fin- gers are webbed only basally (fig. 271B). The hind limbs are moderately short and stout; the heels of the adpressed limbs barely over- lap. The tibiotarsal articulation extends to the eye. A thin, transverse dermal fold is present on the heel, and a low, rounded tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is elliptical, barely rounded in profile, and not visible from above. There is no outer metatarsal tubercle. The toes are moderately long and slender and bear discs that are only slightly smaller than those on the fingers. The subarticular tubercles are small and conical; low, indistinct supernu- merary tubercles are present on the proximal segments of each digit. The toes are about two-thirds webbed (fig. 272B). The webbing extends from the base of the penultimate pha- lanx of the first toe to the distal end of the antepenultimate phalanx of the second, from the distal end of the penultimate phalanx of the second to the middle of the antepenulti- mate phalanx of the third, from the middle of the penultimate phalanx of the third to the base of the antepenultimate phalanx of the fourth and on to the base of the disc of the fifth toe. The anal opening is directed posteroven- trally at the midlevel of the thighs. A short anal sheath and a transverse dermal fold above the sheath are present. The skin on the dorsum is smooth, except for a few minute tubercles on the head and in the sacral re- gion. The skin on the belly and \entral sur- faces of the thighs is strongly granular, where- as the skin on the other ventral surfaces is smooth. The tongue is round, shallowly notched posteriorly, and barely free behind. The dentigerous processes of the prevomers are widely separated, small transverse ele- vations between the minute choanae. There are two and three teeth on the elevations. There are 30 teeth on the maxillary and pre- maxillary on one side and .31 on the other. Vocal slits and a vocal sac are absent. The color in life is unknown. In preserva- tive, the dorsum is dark brown; the anterior and posterior surfaces of the thighs, the inner surfaces of the shanks, the ventral surfaces of the body and limbs, and the webbing on the feet are dull tan (pi. 4, fig. 3). T.^DPOLES; The tadpoles of this species are unknown. M.\TiNG Call: The apparent absence of vocal slits and a vocal sac is suggestive that this species has no call. Natur.\l History: Nothing is knovim of the natural history of this species. Remarks: Bumzahem and Smith (1954, p. 64) named and described this species on the basis of a single specimen from the "Re- gion de Soconusco, Chiapas, Mexico, collected by Mr. Eizi Matuda between 1944 and 1949." Not only did the authors name a new species on the basis of a specimen lacking precise locality data, they based their description on an extremely poorly preserved specimen. I am unsure as to the status of Plectrohtjla lacertosa. Possibly it is a very distinctive spe- cies in the genus, but on the basis of the single poorly preserved specimen at hand, it is diffi- cult to ascertain the relationships with the other members of the genus. The hideously swollen lips and the enormously hypertro- phied arms are suggestive of a possible dis- ease-ridden frog, perhaps suffering from a form of anuran elephantiasis. Mr. David M. Dennis labored arduously to depict the type specimen in the form of a living frog as shown in Plate 4. Etymology: The specific name is Latin meaning nmscular and refers to the greatly swollen arms. Distribution': Plectrohyla lacertosa is known only from the Region de Soconusco, Chiapas, Mexico; the species is not known from any definite locality. See Appendix 1 for the record of the one specimen examined. Plectrohyla avia Stuart Plectrohyla avia Stuart, 1952, p. 6 [holotype, U.M.M.Z, No. 102280 from Granja Lorena, 10 kilo- meter.s airline northwest of Colomba, Departamento Quetzaltenango, Guatemala; Laurence C. Stuart col- lector]. Stuart, 1963, p. 39. Diagnosis: This large species (90 mm. snout-vent length) has a smooth dorsum, ex- cept for small tubercles on the head. Males have a long, single, pointed prepollical spine and lack vocal slits. The dorsum is uniform green. Plectrohyla avia can be distinguished by the above characters from other species 1970 DUELLMAN: HYLID FROGS 579 lacking vocal slits; of these, guatemalensis and hartwegi have bifid prepollical spines and tuberculate skin on the dorsum. PlectroIiyJa glatuhilosa and pycnochila have blunt pre- pollical processes; the latter has a tuberculate dorsum, and the former has a smooth dorsum. Additionally, gJanduhsa differs from avia by ha\ing a pointed snout and mottled dorsum; in acta, the snout is blunt, and the dorsum is uniform green. PlectrohyUi lacertosa is small- er (47 mm. snout-vent length), is brown above, and has an elongate, round, terminally blunt prepollical spine. Species of Plectro- hyla not mentioned above have vocal slits and are smaller (less than 50 mm. in snout- vent length); none is uniform green above. Descriptiox: This is the largest species in the genus. Males attain a maximum known snout-vent length of 90.4 mm.; the females are unknown. Four adult males have snout- vent lengths of 82.5 to 90.4 (mean, 86.2) mm.; the ratio of tibia length to snout-\'ent length is 0.483 to 0.532 (mean, 0.509); the ratio of foot length to snout-vent length is 0.470 to 0.487 (mean, 0.478); the ratio of head length to snout-vent length is 0.317 to 0.3.35 (mean, 0.323); the ratio of head width to snout-vent length is 0.354 to 0.356 (mean, 0.355), and the ratio of the diameter of the tympanum to that of the eye is 0.397 to 0.543 (mean, 0.468). The head is as broad as the body, and the top of the head is flat. In dorsal profile, the snout is rounded; in lateral profile it is bluntly rounded, nearly truncate. The snout is mod- erately short, and the nostrils are noticeably protuberant and situated at a point about three-fourths of the distance from the eyes to the tip of the snout. The canthus is a fold-like ridge; the loreal region is noticeably concave, and the lips are thick and swollen. An ex- tremely heavy dermal fold extends posteriorly from the eye, above the tympanum, and downward to the point of insertion of the arm. The fold obscures the upper edge of the tympanum, which otherwise is distinct and separated from the eye by a distance equal to half again the diameter of the tympanum. The arms are moderately short and ex- tremely robust. No tubercles are present on the ventrolateral edge of the forearm, but a distinct transverse dermal fold is present on the wrist. The fingers are moderately long and slender and bear large discs; the width of the disc on the third finger is equal to the diameter of the tympanum. The subarticular tubercles are large and subconical; that on the first finger is bifid. The supernumerary tu- bercles are small, conical, and arranged in a single row on the proximal segments of each digit. A diffuse palmar tubercle is present. The prepollex is enlarged and curved and in breeding males has a horny nuptial excres- cence. The fingers are about one-third webbed (fig. 273A). The webbing between the first and second fingers is vestigial; the webbing extends from the distal end of the antepenultimate phalanx of the second to the base of the antepenultimate phalanx of the third, and from the middle of the antepenulti- mate phalanx of the third to the distal end of the antepenultimate phalanx of the fourth finger. The hind limbs are moderately long and robust; the heels of the adpressed limbs o\'erlap by about one-third of the length of the shank. The tibiotarsal articulation ex- tends to the eye. A thin transverse dermal fold is present on the heel, and a low, incon- spicuous tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is large, elliptical, and elevated. The outer metatarsal tubercle is low and elliptical. The toes are long and slender and bear discs that are somewhat smaller than those on the fin- gers. The subarticular tubercles are moder- ately small and conical; the supernumerary tubercles are small, conical, and arranged in a single row on the digits. The toes are about three-fourths webbed (fig. 273D). The web- bing extends from the middle of the penulti- mate phalanx of the first toe to the base of the penultimate phalanx of the second, from the middle of the penultimate phalanx of the second to the distal end of the antepenulti- mate phalanx of the third, from the middle of the penultimate phalanx of the third to the base of the penultimate phalanx of the fourth and on to the middle of the penultimate pha- lanx of the fifth toe. The anal opening is directed posteroven- trally near the midlevel of the thighs. A short anal sheath and a supra-anal flap are present. The skin on the dorsum is smooth, except that on the top of the head and on the sides of the head small tubercles are present. The 580 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 throat, chest and ventral surfaces of the arms and shanks are smooth, whereas the skin on the belly and ventral surfaces of the thighs is granular. The tongue is ovoid, slightly longer than wide, shallowly notched posterior- ly, and barely free behind. The dentigerous processes of the prevomers are narrowly sepa- rated, posteromedially inclined ridges be- tween the moderately small, elliptical cho- anae. There are one to three teeth on each ridge. The number of ma.xillary and pre- maxillary teeth (one side only) varies from 27 to 33. Vocal sUts and a vocal sac are ab- sent. The general coloration of Plectrohyla avia is uniform green (pi. 69, fig. 4). The moder- ate dark green of the dorsum is faded on the sides of the head and flanks. The anterior and posterior surfaces of the thighs are greenish gray, and the venter is grayish white. The iris is bronze. In preservative, the dorsum is dull bluish gray and the venter is creamy white. There is no trace of a pattern. Tadpoles: The tadpoles of this species are unknown. Mating Call: This species lacks vocal slits and apparently lacks a vocal sac; it is pre- sumed that it lacks a voice. Natural History: Stuart (1952, p. 6) obtained the type specimen in scrubby forest on April 21, 1949. I observed an adult sitting on a branch over a small stream at the type locality in July, 1966. Two specimens were obtained in August along a stream on Volcan Tacana, Chiapas. Presumably, this species is like others in this genus and breeds in moun- tain streams. Remarks: The humerus in this species is modified by having extensively developed ridges, presumably for the attachment of the large brachial muscles (fig. 274). Etymology: The specific name is Latin, meaning grandmother, and alludes to the large size of this species. Distribution: Plectrohyla avia is known from cloud forest at elevations of 1700 to 2000 meters on the Pacific slopes of the Sierra Madre from south-central Chiapas, Mexico, to southwestern Guatemala (fig. 279). See Appendix 1 for the locality records of the six specimens examined. Plectrohyla guatemalensis Brocchi Plectrohyla guatemalensis Brocchi, 1877a, p. 92 [syntypes, M.N. H.N. No 6332 (2 specimens), from Pacicilla (=Patzicia), Departamento Chimaltenango, Guatemala; Marie-Firmin Bocourt collector]. Stuart, 1963, p. 39. Cauphias guatemalensis Brocchi, 1877b, p. 130; 1882, p. 62. Hyla guatemalensis: Boulenger, 1882a, p. 396. Gunther, 1901 ( 1885-1902), p. 281. Di.^GNOsis: This large species (76 mm. snout- vent) has a weakly to strongly tubercu- late dorsum. Males have a bifid prepollical spine and lack vocal slits. The only other species with a bifid prepollical spine, hart- xcegi, has dark vertical bars on the flanks and anterior and posterior surfaces of the thighs and dark mottling on the ventral surfaces of the shanks; guatemalensis lacks these bold markings. Of the other species lacking \'ocal slits, avia has a single, terminally pointed pre- pollical spine and a smooth green dorsum, except for small tubercles on the head. Plec- trohyla pyctwchila has a blunt, flat prepollical process, and lacertosa is much smaller (47 mm. snout-vent length) and has an elongate, round, terminafly blunt prepollical spine. The species possessing vocal shts are smaller ( less than 50 mm. snout-vent); each has a single, pointed prepollical spine. Description: Males of this large species attain a maximum known snout-vent length of 76.1 mm., and females reach 73.6 mm. In a series of six adult males from Finca Los Alpes, Departamento Alta Verapaz, Guatema- la, the snout-vent length is 72.1 to 76.1 (mean, 73.4) mm.; the ratio of tibia length to snout- vent length is 0.539 to 0.576 (mean, 0.563); the ratio of foot length to snout-vent length is 0.457 to 0.513 (mean, 0.488); the ratio of head length to snout-vent length is 0.274 to 0.292 (mean, 0.285); the ratio of head width to snout- vent length is 0.333 to 0.347 (mean, 0.343), and the ratio of the diameter of the tympanum to that of the eye is 0.240 to 0.357 (mean, 0.304). Five adult females from the same locality have snout-vent lengths of 68.4 to 73.6 (mean, 70.4) mm. and do not difl^er significantly in proportions from the males. Individuals from the western part of the range, in Chiapas, Mexico, are somewhat smaller than those specimens from Guate- 1970 DUELLMAN: HYLID FROGS 581 mala (the eastern part of the range). Fur- thermore, the specimens from Chiapas have proportionately shorter limbs and a smaller head, but a proportionately larger tympanum. For example, seven adult males from streams above Rayon Mescalapa, Chiapas, Me.xico, have snout-vent lengths of 51.2 to 61.5 (mean, 55.9) mm.; the ratio of tibia length to snout- vent length is 0.486 to 0.558 (mean, 0.518); the ratio of foot length to snout-vent length is 0.407 to 0.464 (mean, 0.446); the ratio of head length to snout-vent length is 0.255 to 0.305 (mean, 0.287); the ratio of head width to snout-vent length is 0.291 to 0..36.3 (mean, 0.327), and the ratio of the diameter of the tympanum to that of the eye is 0.318 to 0.379 (mean, 0.352). The head is nearly as broad as the body, and the top of the head is flat. In dorsal pro- file, the snout is bluntly rounded, and in lat- eral profile it slopes abruptly from the nostrils to the edge of the jaw. The snout is short, its length is equal to the diameter of the eye. The nostrils are barely protuberant and nearly terminal. The canthus is well defined and angular; the loreal region is deeply concave, and the lips are thick and moderately flared. A heavy dermal fold extends posteriorly from the eye, above the tympanum, and downward to a point above the insertion of the arm. One or two dermal folds extend ventrally from this heavy fold. The fold obscures the upper edge of the tympanum, which in most specimens otherwise is distinct and separated from the eye by a distance equal to more than twice the diameter of the tympanum. The arms are short and robust; they are especially heavy in breeding males. A few small tubercles are present on the ventrolat- eral edge of the forearm, and a heavy trans- verse dermal fold is present on the wrist. The fingers are moderately long and slender and bear large discs; the width of the disc on the third finger is more than twice the diameter of the tympanum. The subarticular tubercles are large and subconical; the distal tubercle on the fourth finger is flattened and in some individuals faintly bifid. The super- numerary tubercles are moderately large and conical; they are arranged in a single row on the proximal segments of each digit. A large, diffuse, bifid palmar tubercle is present. The prepollex is large, elongate, and bifid. The webbing in the hands is vestigial (fig. 273B). The hind limbs are moderately long and ro- bust; the heels of the adpressed limbs overlap by about one-third of the length of the shank. The tibiotarsal articulation extends to the pos- terior corner of the eye. A heavy transverse dermal fold is present on the heel, and an elevated tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is large, flat, and elliptical. The outer meta- tarsal tubercle is small and subconical. The toes are moderately long and slender and bear discs that are only slightly smaller than those on the fingers. The subarticular tuber- cles are large and subconical. Moderately large, subconical, supernumerary tubercles are present on the proximal segments of each digit. The toes are about three-fourths webbed (fig. 273E). The webbing extends from the base of the disc of the first toe to the base of the penultimate phalanx of the second, from the distal end of the penulti- mate phalanx of the second to the base of the penultimate phalanx of the third, from the distal end of the penultimate phalanx of the third to the base of the penultimate phalanx of the fourth and onto the base of the disc of the fifth toe. The anal opening is directed posteroven- trally near the midlevel of the thighs. A short, heavy, anal sheath is present; it has a membraneous connection with the skin on the posterior surfaces of the thighs. The skin on the dorsum is smooth or bears small scattered tubercles. The skin on the throat, belly and ventral surfaces of the thighs is granular; that on the ventral surfaces of the arms and shanks is smooth. The tongue is nearly round and barely free behind. The tongue is notched shallowly posteriorly and in all specimens and anteriorly in some specimens. The den- tigerous processes of the prevomers are nar- rowly separated, transverse elevations be- tween the posterior margins of the quad- rangular choanae. There are three to six teeth on each elevation. The number of teeth on the maxillary and premaxillary (one side only) varies from 32 to 39. Vocal slits and a vocal sac are absent. The general coloration of Plectrohijla gtia- temalensis is dull green above variously 582 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 marked, or not, with shades of brown (pi. 69, fig. 3). Individuals from Alta Verapaz, Guatemala, were primarily dull olive-green above with or without tan or brown markings. The venter was grayish white. One individual from 5.6 kilometers south of Rayon Mescala- pa, Chiapas, Me.xico, was dull gray above with olive-green spots. The webbing and venter were gray. One specimen from near Panajachel, Solola, Guatemala, was dark green above with reddish brown markings; the posterior surfaces of the thighs and the webbing \\'ere gray. Another specimen from Granja Lorena, Quetzaltenango, Guatemala, had a dull olive-brown dorsum; the flanks and posterior surfaces of the thighs were pale green, and the venter was gray. In all individuals, the iris was golden bronze with or without fine black reticulations. In preservative, the dorsum is dark brown, bluish black, or dull gray. The venter is dull creamy tan or grayish brown. Tadpoles: A typical tadpole in develop- mental stage 27 has a body length of 15.2 mm. and a total length of 43.0 mm. The body is ovoid, widest posteriorly, and no wider than deep. In dorsal profile the snout is bluntly rounded, and in lateral profile, it is acutely rounded. The eyes are small, widely sepa- rated, and directed dorsolaterally. The nos- trils are directed anterolaterally at a point about midway between the eyes and the tip of the snout. The opening of the sinistral spiracle is about on the midline at a point slightly posterior to the midlength of the body. The cloacal tube is long and dextral. The caudal musculature is moderately robust and does not extend to the tip of the rounded tail. The fins are shallow; at midlength of the tail, the caudal musculature is deeper than either the ventral or dorsal caudal fin. The dorsal fin does not extend onto the body (fig. 275F). The body is dark brown with scattered lichenous markings laterally. The caudal musculature is pale brown, and dark brown flecks and small blotches are present on the musculature and fins. In preservative, the body is dark brown, and the caudal muscu- lature is creamy tan. Faint brown blotches are evident on the musculature and fins. The mouth is ventral and large; its width is equal to about two-thirds the width of the greatest width of the body. There is no lateral fold, and the lips are completely bordered by two rows of small papillae. Medial to the fringing rows is a single row of larger pa- pillae; numerous large papillae are present laterally. The beaks are moderately robust and bear short, blunt serrations. The upper beak is in the form of a broad arch with short lateral processes, and the lower beak is broad- ly V-shaped. There are two upper and three lower rows of teeth. The upper rows are long and equal in length; the second upper row is narrowly interrupted medially. The lower rows are complete, moderately long and of equal length (fig. 276F). Stuart (1942, p. 8) described and illus- trated this tadpole as "Form x." Mating Call: The absence of vocal slits and a \'ocal sac precludes the presence of a mating call in this species. Natural History: Plectrohyla guatema- lensis is an inhabitant of cloud forest and humid pine-oak forest. At Finca Los Alpes, Guatemala, adults were found on vegetation overhanging streams by day and by night. At the same locality, individuals were found sitting on rocks behind the waterfall, in a hole in a cliff behind a waterfall, and on rocks in the streams at night. On cloudy or rainy days, these frogs frequently are active; at these times they can be found perched on rocks or vegetation in or along cascading mountain streams. Tadpoles in various stages of development have been found throughout most of the year. Thus, it seems likeh' that this species lias no definite breeding season. Metamorphosing young were found at Finca Los Alpes on July 31, 1961, and along a stream 6.2 kilometers south of Ravon Mescalapa, Chiapas, on June 16, 1960. Stuart (1954c, p. 48) found three subadults in a bromeliad in a tree overhang- ing a nearly dry stream where tadpoles were present at San Lorenzo, Guatemala, in mid- February. Recently metamorphosed young having snout-vent lengths of 23.0 and 24.4 mm. were pale oli\ e-green abo\'e with pale green blotch- es posterolaterally; the throat and chest were silvery green. Remarks: Plectrolujia guatctualensis oc- 1970 DUELLMAN: HYLID FROGS 583 curs s>'mpatrically with sc\cral other species of the genus {avia, matudai, sagoniin, qiiec- clii, and ixil). Plectwhhja guatemalensis and avia tend to frequent the larger streams than do the smaller species ( matudai, sagonim, quecchi, and ixil). \\hich often inhabit rivu- lets. Etymology: The specific name refers to Guatemala, country of origin of the type specimen. Distribution: Plectrohyla guatemalensis occurs at ele\ations from 1000 to 2800 meters on the Atlantic slope of the highlands of Chiapas and Guatemala eastward to the Si- erra de Nombre de Dios in north-central Honduras; on the Pacific slopes, the species occurs from south-central Chiapas eastward to northern El Salvador (fig. 280). See Appendix 1 for the locaHty records of the 103 specimens examined. Plectrohyla hartwegi Duellman Plectrohyla hartwegi Duellman, 1968a, p. 576 [holot\pe, U.M.M.Z, No. 94428 from Barrejonel, 19 kilometers west of Chicomuselo, Chiapas, Me.xico, 1000 meters; Eizi Matuda collector]. Dl\gnosis: This moderately large species (64 mm. in snout-vent length) has a bifid prepoUex and a tuberculate dorsum. Plectro- hyla haiiicegi differs from all other members of the genus by having bold bars on the flanks and anterior and posterior surfaces of the thighs, and dark mottling on the ventral surfaces of the shanks. Description': Males of this species attain a maximum known snout-vent length of 63.8 mm.; the females are unknown. Three males have snout-vent lengths of 41.8 to 63.8 (mean, 51.3) mm.; the ratio of tibia length to snout- vent length is 0.547 to 0.579 (niean, 0.558); the ratio of foot length to snout-vent length is 0.464 to 0.487 (mean, 0.475); the ratio of head length to snout-vent length is 0.309 to 0.3.39 (mean, 0..3.32); the ratio of head width to snout- vent length is 0.350 to 0.377 (mean, 0.360), and the ratio of the diameter of the tympanum to that of the eye is 0.426 to 0.473 (mean, 0.445). The head is as broad as the body, and the top of the head is flat. In dorsal profile, the snout is bluntly rounded, and in lateral pro- file, it is angular and slopes abruptly from 94° 90° 86° 16° 14° 16° 14° ~. y ■■- ^' • ^.-\ \ L... -. \ I-. i ■» ( i f i V. .' \ f c • 1 X - • P guatemalensis 0 P hartwegi \ ^-' 0 50 20 0 KILOMETERS 1 94° 90° 86° Fig. 280. Distribution of Plectrohyla guatemalensis and Plectrohtjla hartwegi. 584 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 the nostrils to the jaw. The snout is short, and the nostrils are barely protuberant and situated at a point about two-thirds of the distance from the eyes to the tip of the snout. A heavy dermal fold extends from the eye, above the tympanum, and downward to a point above the insertion of the arm. Two thinner folds extend ventrally from the heavy fold and cover the posterior edge of the tym- panum. The anterior and ventral edges of the tympanum are distinct, and the tympan- um is separated from the eye by a distance equal to the diameter of the tympanum. The arms are robust. There is no distinct row of tubercles on the ventrolateral edge of the forearm, but there is a faint transverse dermal fold present on the wrist. The fingers are long and moderately slender and bear large discs, the width of the disc on the third finger is noticeably greater than the diameter of the tympanum. The subarticular tubercles are small and conical; except the distal tu- bercle on the fourth finger, which is some- what flattened (bifid in one specimen). The supernumerary tubercles are small, subconi- cal, and arranged in one row on the proximal segment of the fourth finger and in two rows on the proximal segments of the other fingers. Two small palmar tubercles are present. The prepollex is greatly enlarged, barely bifid, and does not have spines protruding through the skin. The webbing on the hands is vesti- gial (fig. 273C). The hind limbs are mod- erately long and robust; the heels of the ad- pressed limbs overlap by about one-third of the length of the shank. The tibiotarsal ar- ticulation extends slightly beyond the tip of the snout. A heavy transverse dermal fold is present on the heel, and a heav>' tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is high, elliptical, and visible from above. The outer metatarsal tubercle is absent. The toes are long and slender and bear rather small discs. The sub- articular tubercles are small and round; the supernumerary tubercles are small and ar- ranged in a single row on the proximal seg- ment of each digit. The toes are about three- fourths webbed (fig. 273F). The webbing extends from the base of the disc of the first toe to the base of the penultimate pha- lanx of the second, to the base of the disc of the second to the base of the penultimate phalanx of the third, from the base of the disc of the third to the base of the penultimate phalanx of the fourth and on to the base of the disc of the fifth toe. The anal opening is directed posteroven- trally at the midlevel of the thighs. The anal sheath is long and has a membraneous con- nection to the posterior surfaces of the thighs. The skin on the dorsal surfaces is finely tu- berculate; that on the throat, chest, belly, and posteroventral surfaces of the thighs is granular, whereas the skin on the ventral sur- faces of the arms and shanks is smooth. The tongue is nearly round and free posteriorly for about one-fourth of its length; it is margi- nate or barely notched behind. The dentig- erous processes of the prevomers are small elliptical elevations between the quadrangu- lar choanae. There are four or five teeth on each process. The number of maxillary and premaxillary teeth (one side only) varies from 35 to 40. Vocal slits and a vocal sac are absent. The coloration in life is unknown. In preservative, the dorsum is uniformly dull brown. The flanks are brown with creamy yellow mottling and dark brown spots in the groin; the anterior surfaces of the thighs are creamy yellow with broad, vertical, dark- brown bars proximally and narrower dull brown bars distally. The posterior surfaces of the thighs are brown with dark brown vertical bars (pi. 5, fig. 2). The belly and ventral surfaces of the limbs are creamy yel- low; bold brown reticulations are present on the ventral surfaces of the shanks. Tax)Poles: The tadpoles of this species are unknown. Mating Call: The absence of vocal slits and presumably a vocal sac probably pre- cludes the presence of a voice in this species. Natural History: One specimen ob- tained in May on Paraje El Triunfo was found in a rocky stream in the cloud forest at an elevation of 2050 meters. There is no other available information on the natural history of this species. Remarks: On the basis of the general shape of this frog and the presence of a bifid prepollex, it seems logical to associate this species as a relative of Plectrohijla guatema- 1970 DUELLMAN: HYLID FROGS 585 lensis, from which it differs chiefly in color pattern. The geographic ranges of the two species overlap; consequently, it is highly unlikely that Imrttiegi represents a geographic race of ci.uatemalcnsis. Etymology: The specific name is a patro- nym for Norman Hartweg, who first recog- nized the distinctness of this species. Distribution: Plectrohyla harticegi is knowTi from elevations of 1000 to 2050 meters on the Pacific slopes of the Sierra Madre in Chiapas and extreme eastern Oaxaca, Mexico (fig. 280). See Appendix 1 for the locality records of the three specimens examined. Genus Smilisca Cope Smilisca Cope. 1865b, p. 194 [type species, Smilis- ca claiilinia Cope, 1865 = Hyla baudinii Dunieril and Bibron, 1841]. Generotype: Hijla baudinii Dumeril and Bibron, 1841. Cope (1865b, p. 194) in his synopsis of the genera of hylid frogs based the diagnosis of the genus Smilisca on a "skeleton in the private anatomical museum of Hyrtl, Professor of Anatomy in the University of Vienna." Cope referred to the specimen as Smilisca daulinia. Duellman and Trueb (1966, p. 297) suggested that Cope inadver- tently used daulinia (a new name) for bau- dinii just as he later used daudinii for bau- dinii (1871, p. 205). Cope's description of the cranial characters of Hyrtl's specimen leaves no doubt that he had before him a specimen of Smilisca baudinii. Etymology: The generic name is derived from the Greek smile, meaning knife, and the Greek iskos, a diminutive suffix, and means literally "little knife" in reference to the sharp- ly pointed frontoparietal processes of S. bau- dinii used as a diagnostic character of the genus by Cope. DEFixniON: Frogs of the genus Smilisca are medium to large in size and have a blotched or barred dorsal pattern of shades of green or brown. The flanks are mottled, spotted, or venated, and the venter is creamy white, except for dark colored vocal sacs in most species. The pupil is horizontally ellip- tical, and the iris is a bronze color with black flecks or reticulations. The palpebral mem- brane is unmarked. The amount of webbing on the hand is variable, but the toes are at least three-fourths webbed. The first toe is shorter than the second and not opposable to the others. The vocal sacs are paired, sub- gular, and greatly distensible. The skin on the dorsum is smooth; distinct paratoid glands are lacking. The tongue is ovoid, barely free behind, and variously notched or not. Breed- ing males have horny brown nuptial excres- cences on the thumbs. The skull is broad, well ossified, has a minimal amount of carti- lage and/or secondarily ossified cartilage, and lacks dermal co-ossification. An internasal septum and quadratojugals are present. The sphenethmoid is large, and the nasals are moderately slender, separated medially, and separated or not from the sphenethmoid. A frontoparietal fontanelle is present, except in S. phaeota. Extensive, laterally projecting, frontoparietal processes are present in S. bau- dinii and phaeota. A well-developed squa- mosal minimally extends one-fourth of the distance to the maxillary and maximally is in contact with the maxillary. The dentiger- ous processes of the prevomers are short, widely separated, and situated at a slight angle to the midline. Teeth are present on the premaxillaries, maxillaries, and prevo- mers, but absent from the palatines and para- sphenoid. The teeth are spatulate and strong- ly bifid. The depressor mandibulae muscle consists of two parts, one arising from the dorsal fascia and the other from the posterior arm of the squamosal. The adductor mandib- ulae muscle consists of two branches — the posterior subexternus and the externus super- ficialis. The mandibular branch of the tri- gemial nerve passes between the branches of the adducator mandibulae muscle. The tad- poles are generalized and have two upper and three lower rows of teeth, and unspe- cialized beaks. The mouth is partly or com- pletely bordered by one or two rows of pa- pillae, and the lips are infolded laterally. The spiracle is sinistral, and the cloacal tube is dextral. The caudal musculature extends nearly to the tip of the tail. The mating call consists of one or more short, poorly modu- lated, explosive notes. The haploid chromo- some number is 12, and the diploid number is 24. 586 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Composition of Genus: Si.x species are currently recognized. All are considered to be monotypic. All known species occur in Middle America. Of the six species, 4544 pre- served frogs, 95 skeletons, 95 lots of tadpoles, and five preserved clutches of eggs were ex- amined from Middle America. Analysis of Characters: On the basis of size alone the species fall into two groups; baudinii, cyanosticta, and plweota, are large, and puma, sila, and sordida are small. The largest specimen examined is a female haxi- dinii having a snout-vent length of 90 mm. Smilisca puma is the smallest species; the largest male has a snout-vent length of 38 mm. and the largest female, 46 mm. Few sig- nificant differences in proportions exist be- tween the species (table 56). Smilisca bau- dinii is more squat and stocky than the other species and has proportionately shorter hind limbs. Although considerable variation in the size of the tympanum exists within each species, noticeable differences are present be- tween species. Consistent differences exist in relative lengths of the digits, size of the subarticular tubercles, size and number of the supernu- merary tubercles, size and shape of the inner metatarsal tubercle, and the amount of web- bing (figs. 281-283). In the series of large species (bamlinii-pliaeota-cyanosticta) a pro- gressive increase in the amount of webbing in the hand and a decrease in number, and corresponding increase in size, of the super- numerary tubercles are evident. Smilisca puma is unique in the genus by lacking web- bing in the hand and by having large sub- articular tubercles on the hand and a rela- tively small inner metatarsal tubercle. S?7i!- lisca sila and sordida have shorter, more ro- bust fingers than the other species. Both spe- cies have extensive webbing and many small supernumerary tubercles on the feet. The color and pattern are among the most important taxonomic characters in the genus. Especially significant is the coloration of the flanks, which is venated in pliaeota, mottled in baudinii, venated anteriorly and mottled posteriorly in puma, and spotted or flecked in the other species (pis. 70 and 71). Smilisca cyanosticta and phaeota each has a broad white labial stripe, and puma has a narrow stripe. The upper lip is marked with vertical dark bars in baudinii and sila, whereas it is unicolor in sordida. A large dark brown or black postorbital mark is present in baudinii, cyanosticta, and phaeota and absent in the other species. All species have dark trans- verse bands on the limbs. The dorsum of the body in puma is marked by two longitudinal dark stripes that are interconnected in some specimens; the dorsal markings usually con- sist of one or more irregular dark blotches in the other species, but in some specimens of TABLE 56 Comparison of Sizes and Certain Proportions, with Means in Parentheses, of Males of the Species of Smilisca. Species N S. baudinii 140 S. cyanosticta 40 S. pliaeota 50 S. puma 20 S. sila .33 S. sordida 55 Snout-vent Length Tibia Length/ S-VL Tympanum/ Eye 47.3-75.9 0.421-0.536 0.561-0.944 (58.7) 44.6-57.8 (0.478) 0.519-0.597 (0.735) 0.627-0.884 (50.7) 40.8-65.5 (0.560) 0.509-0.602 (0.714) 0.627-0.855 (53.9) 31.9-38.1 (0.555) 0.482-0.531 (0.766) 0.521-0.722 (34.7) 31.6-44.8 (0.513) 0.497-0.581 (0.649) 0.476-0.483 (37.7) 31.9-44.6 (0.548) 0.505-0.571 (0.532) 0.465-0.571 (37.9) (0.5.34) (0.491) 1970 DUELLMAN: HYLID FROGS 587 Fig. 281. Hands of four species of Smilisca. A. S. baudinii, K.U. No. 87182. B. S. ctjauosticta, K.U. No. 87199. C. S. phaeota, K.U. No. 96156. D. S. pwna, K.U. No. 64312. X 4, e.\cept D, which is x 5. 588 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fig. 282. Feet of four species of Smilisca. A. S. baudinii, K.U. No. 87182. B. S. cijanosticta, K.U. No. 87199. C. S. phaeota, K.U. No. 96156. D. S. puma, K.U. No. 64312. x 4, e.xcept D, which is x 5. 1970 DUELLMAN: HYLID FROGS 589 Fig. 283. Hands and feet of two species of Sinilisca. A and C. S. sila, K.U, No. 91867. B and D. S. sordida, K.U. No. 91753. x 5. 590 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 sordida the dark marks form transverse bars. The belly is creamy white in all species; the vocal sac is white in breeding sordida and gray or brown in the other species. The tadpoles of Smilisca sila and sordida live in streams and have ventrally oriented, larger mouths and proportionately longer tails than do the pelagic, pond dwelling tadpoles of the other species (fig. 284). The differ- ences in coloration and in the mouthparts are slight; Smilisca sordida is unique in having two complete rows of labial papillae and long, shallowly S-shaped lateral processes on the upper beak (fig. 285). E.xamination of the skulls- reveals that members of the baudinii group (baitdinii, cij- anosticta, and phaeota) have well ossified skulls that have gently curved lateral margins and relatively large nasals with their long axes parallel to the maxillaries. Anteriorly the nasals are pointed and posteriorly they bear long, delicate palatine processes extend- ing to the maxillaries. The sphenethmoid is fully ossified and extends anteriorly between the nasals. The squamosals are large and ex- tend to the maxillary in baudinii, but not in cyanosticta and phaeota. The prootics are massive. Extensive lateral flanges are present on the frontoparietals in baudinii and pliae- ota (fig. 2S6). The skulls of puma and sor- dida differ from those of the baudinii group by having somewhat angular lateral margins, small bony sphenethmoid that does not ex- tend anteriorly between the nasals, and rela- 'tively small prootics. The moderate-sized nasals are rounded anteriorly and bear rela- tively short maxillary processes; the long axes of the nasals are not parallel to the maxillaries. The squamosals are small and do not extend to the maxillaries (fig. 286). The skull of Smilisca sila is intermediate between these two species groups. The lateral margins are gently curved but have a pronounced angu- larity just anterior to the palatines. The na- sals are moderate in size and have their long axes parallel to the maxillaries. The nasals are slightly pointed anteriorly and bear short, blunt palatine processes posteriorly. The sphenethmoid is extensively ossified but does not extend anteriorly between the nasals. The prootics are relatively large but short. The squamosals arc moderate in size; the anterior arms extend only one-fourth the distance to the maxillary. Duellman and Trueb (1966) discussed the comparative osteology of the species of Smilisca in detail, and Trueb (196Sb) described the internal cranial anat- omy of S. baudinii. The mating calls of all species of Smilisca consist of short, explosive, poorly modulated notes. The calls consist of one "wonk" of series of such notes in baudinii and cyano- sticta, a low growl in phaeota, and a relatively high-pitched rattle in sordida. The calls of puma and sila consist of a low-pitched scjuawk usually followed by one or more rattling sec- ondary notes. Quantitatively, the calls of the species differ in the number of notes, dura- tion of notes, and in pitch (table 57, pis. 32 and 33). Distribution: The combined distribu- tions of the six species of Smilisca include most of the lowlands of Mexico and Central .America, in some places to elevations of near- ly 2000 meters. The range extends from southern Sonora, Mexico, and the Rio Grande Embaymcnt of Texas to South America and includes such continental islands as Isla Co- zumel, Mexico, Isla Roatan, Honduras, and Isla Popa and Isla Cebaco in Panama. In South America one species occurs on the Caribbean lowlands of Colombia and in the \alleys of the Rio Cauca and Rio Magdalena; another species occurs on the Pacific slopes of Colombia and northwestern Ecuador. Discussion: The genus Smilisca has not been consistently recognized by workers in the past twenty years. Except for Cope's \'arious publications dealing with the Neo- tropical herpetofauna, the name was not used in the 1800's. Smith and Taylor (1948) res- urrected the generic name and followed Cope by only including Hyla baudinii Dumeril and Bibron in the genus. Starrctt (1960b) ex- panded the definition of the genus and placed Hyla i:,abbi Cope, Uyhi phaeota Cope, and Hyla icellmanorum Taylor in the genus. Duellman and Trueb (1966) refined the defi- nition of the genus and recognized the six species that are currenth- placed in the genus. Although Smilisca is difficult to define, the six included species seem to form a natural group. The paired subgular vocal sacs are a reliable diagnostic character. Experience 1970 DUELLMAN: HYLID FROGS 591 Fig. 284, Tadpoles of the species of Smilisca. A. S. baudinii, K.U. No. 60018. B. S. cyanosticta, K.U. No. 87652. C. S. phacota, K.U. No. 87683. D. S. puma, K.U. No. 91807. E. S. sila, K.U. No. 80620. F. S. sordida, K.U. No. 68475. x 4.5. 592 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 ^af^^omoi&^,^^_^,^S§6^' Fig. 285, Mouths of tadpoles of the genus Smilisca. A. S. baudinii, K.U. No. 60018. B. S. cyanosticta, K.U. No. 87652. C. S. phaeola, K.U. No. 87683. D. S. puma, K.U. No. 91807. E. S. sila, K.U. No. 90620. F. S. sordida, K.U. No. 68475. All x 25, except F, which is X 17. 1970 DUELLMAN: HYLID FROGS 593 Fig. 286. Dorsal and lateral views of the skulls of Smilisca. and D. S. sordida, K.U. No. 34872. x 3. A and B. S. phaeota, K.U. No. 91827. C TABLE 57 Characteristics of the Mating Calls, with Means in Parentheses, of the Species of Smilisca. Species N S. haudinii 20 S. cyanosticta 10 S. phaeota 10 S. puma 28 S. sila 15 S. sordida _._ 19 Notes per Call Group Duration of Note (seconds) Fundamental Frequency (cps) Major Frequencies ( cps ) Lower Upper 2-15 0.09-0.13 135-190 175-495 2400-2725 (8.0) (0.11) (166.2) (351) (2507) 1-2 0.25-0.45 135-160 480-935 1600-2100 (1.2) (0.38) (145.1) (841) (1894) 1-2 0.10-0.45 110-165 3.30-495 (1.6) (0.31) (143.0) (372) 2-10 0.06-0.35 125-200 495-980 1456-2240 (3.7) (0.13) (145.6) (743) (1868) 1-6 0.06-0.28 90-115 665-1180 1980-2700 (2.4) (0.16) (103.0) (889) (2218) 1-6 0.18-0.45 90-140 1150-1540 2340-2990 (1.7) (0.29) (123.1) (1216) (2694) 594 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 with the frogs in the field substantiates the close relationship of the six species. The mating calls, behavior, and general habitus are sufficiently alike so as to remove doubts about their relationships. By utilizing internal and external mor- phological characters, larval characters, mat- ing calls, and analyses of skin proteins, Duell- man and Trueb (1966) divided the genus into two species groups. The baudinii group contains the three large species {haudinii, cijanosticta, and phaeota), and the sordida group contains the three small species ( puma, sila, and sordida). The reader is referred to Duellman and Trueb (1966) for a detailed discussion of the phylogenetic relationships and a reconstruction of the phylogenetic his- tory. Smilisca baudinii (Dumeril and Bibron) Hijla baudinii Dumeril and Bibron, 1841, p. 564 [holotv-pe, M.N.H.N. No. 4798 from "Mexique"; 13audin collector; type locality restricted to Cordoba, Veracruz, Mexico, elevation 925 meters by Smith and Tavlor (1950)). Brocchi, 1882, p. 29. Boulenger, 1882a, p. .371. Giinther, 1901 (1885-1902), p. 270. Kellogg, 19.32, p. 160. Hijla vanvlietii Baird, 1854, p. 61 [lrolot\'pe, U.S.N.M. No. 3256 from Brownsville, Cameron County, Texas, elevation 15 meters; Captain S. Van Vliet collector]. Htjla vociferans Baird 1859. p. 35 [figures 11-13 on plate 38 are designated "Hyla vociferans, Baird"; the name is not mentioned in the text, nor is a speci- men designated]. Hyla muricohr Cope, 1862, p. 359 [holotype, U.S.N.M. No. 25097 from Hacienda Mirador, Vera- cruz, Mexico, elevation 1020 meters; Charles Sartorius collector]. Smilisca dauliiiia (lapsus for baudinii) Cope, 186.5h, p. 194. Smilisca daudinii (lapsus for baudinii) Cope, 1871, p. 31. Smilisca baudinii: Cope, 1875, p. 31. Taylor, 1952c, p. 794. Stuart, 1963, p. 41. Duellman and Trueb, 1966, p. 289. Hyla pansosana Brocchi, 1877b, p. 125 [holotype, M.N.H.N. No. 6313 from Panzos, Alta Verapaz, Guate- mala, elevation 36 meters; Marie-Firmin Bocourt col- lector). Hyla baudinii { baudinii by fiat): Barbour, 1923, p. 11. Hyla baudinii baudinii: Stejneger and Barbour, 1923, p. 34. Hijk bcltrani Taylor, 1942d, p. 306 [holotype, U.I.M.N.H. No. 25046 from Tapachula, Chiapas, Mexico, elevation 140 meters; A. Magana collector]. Smilisca haudini baudini: Smith, 1947, p. 408. Smith and Taylor, 1948, p. 75. Hyla manisorum Taylor, 19.54b, p. 630 [holot>pe, K.U. No. 34927 from Batan, Limon Province, Costa Rica, elevation 15 meters; Edward H. Taylor collec- tor]. Diagnosis: This large member of the genus is readily discernible from other Smi- lisca by the presence of a large, high, ellipti- cal inner metatarsal tubercle, a short, bluntly rounded snout, relatively short hind limbs (tibia length is less than 55 per cent of the snout- vent length), contrasting dark vertical bars on the upper lip, a broad postorbital dark mark, cream flanks with bold brown or black reticulations in the groin, the posterior surfaces of the thighs brown with cream flecks, and the dorsal surfaces of the limbs marked with dark transverse bands. The dorsum is variously marked with large spots or blotches, and in breeding males the vocal sacs are gray. Other members of the Smilisca baudinii group (cijanosticta and phaeota) have a low, flat, elliptical inner metatarsal tubercle, a more pointed snout, relatively longer hind limbs, and a white labial stripe. Furthermore, the flanks in plmeota are pale cream with a brown or black venated pattern, and the flanks and thighs in cijanosticta are dark brown with pale blue or green spots. The only other Smilisca with a short truncate snout is the much smaller (maximum size of males, 45 mm., of females, 62.2 mm.) S. sila, which has blue spots or flecks on the flanks and posterior surfaces of the thighs. Descriptio.v; Smilisca baudinii is the larg- est species in the genus; males attain a maxi- mum snout-vent length of 76 mm., and fe- males reach 90 mm. The size attained by adults of both sexes varies geographically. The largest specimens are from Sinaloa ( mean snout- vent length of males, 6S.6 mm. ) ; those from the Atlantic lowlands of Alta Vera- paz in Guatemala, Honduras, and Costa Rica are somewhat smaller, whereas those from the Pacific lowlands of Central America are smaller still. The smallest breeding males are from Isla del Carmen, Campeche, Mexico (mean snout-vent length, 50.9 mm.). In a sample of 25 males from Esparta, Puntarenas Province, Costa Rica the snout- vent length is .53.3 to 66.0 (mean, 60.2) mm. The ratio of the tibia length to the snout- 1970 DUELLMAN: HYLID FROGS 595 vent length is 0.451 to 0.520 (mean, 0.482); the ratio of the foot length to snout-vent length is 0.422 to 0.489 (mean, 0.448), the ratio of head length to snout-vent length is 0..300 to 0..342 (mean, 0.320); the ratio of head width to snout-vent length is 0..344 to 0..383 (mean, 0.361), and the ratio of the diameter of the tympanum to that of the eye is 0.679 to 0.911 (mean, 0.777). Considerable \ariation in certain proportions is evident from samples selected from throughout the range, but no geographic trends are apparent. Specimens from Sinaloa in northwestern Mex- ico have the largest tympani (mean tym- panum/eye ratio, 0.878); the next highest ratio (0.794) is in frogs from Managua, Nica- ragua, whereas frogs from intermediate lo- calities have smaller tympani (ratio at Oco- tito, Guerrero, Mexico, 0.746). Similar dis- cordant variation occurs in the relative length of the hind limb. The mean ratio of tibia length to snout-\'ent length is 0.512 and 0.515 in Limon Province, Costa Rica, and in De- partamento Atlantida, Honduras, respective- ly; the ratio is 0.449 in specimens from San Sahador, El Salvador, and from southern Sinaloa, Mexico. The ratios are intermediate in frogs from other localities. See Duellman and Trueb ( 1966 ) for further data on geo- graphic variation in size and proportions. The head is about as wide as the body and is wider than long. The top of the head is flat. In dorsal profile the snout is acutely rounded; in lateral profile the snout is bluntly rounded. The snout is moderately short. The nostrils are slightly protuberant and are situ- ated at about three-fourths the distance from the eyes to the tip of the snout. The canthus is rounded and distinct; the loreal region is noticeably concave, and the lips are moder- ately thick and barely flared. A moderately heavy dermal fold extending posteriorly from the posterior corner of the eye to a point above the insertion of the arm conceals the upper edge of the tympanum in some speci- mens. Otherwise the tympanum is distinct and separated from the eye by a distance slightly less than the diameter of the tym- panum. The arm is moderately long; the upper arm is slender, and the forearm is robust. No axillary membrane is present. A row of low tubercles is present on the ventrolateral edge of the forearm, and a distinct transverse fold is present on the wrist. The fingers are mod- erately long and stout and bear moderately large discs. The width of the disc on the third finger nearly equals the diameter of the tympanum. The subarticular tubercles are small and conical; the distal tubercle on the fourth finger is flattened and in about half of the specimens is bifid. The supernumerary tubercles are small, conical, and distinct. Usually they are in two rows on the proximal segment of each finger, except the third, where they are in three or four rows. A tri- partite palmar tubercle is present. The pre- pollex is moderately enlarged and in breeding males bears a horny nuptial excrescence. The fingers are about one-third webbed (fig. 281A). A trace of web exists between the first and second fingers; the web extends from the base of the penultimate phalanx of the second finger to the base of the antepenulti- mate phalanx of the third and from the mid- dle of the antepenultimate phalanx of the third to the distal end of the antepenultimate phalanx of the fourth finger. The hind limbs are short and heavy; the adpressed heels bare- ly overlap, and the tibotarsal articulation ex- tends to a point between the tympanum and the eye. A heavy tarsal fold extends the length of the tarsus. The inner metatarsal tubercle is large, high, and elliptical. The shape of the tubercle varies from an elongate ellipse with rounded edges to a spade-like structure. The tubercle is most pronounced in specimens from northwestern Mexico, Ta- maulipas, and the Pacific lowlands of Central America. The toes are moderately long and broad; the discs are noticeably smaller than those on the fingers. The subarticular tuber- cles are moderately large and subconical; the supernumerary tubercles are small, conical, and in a single row of each toe. The toes are about three-fourths webbed (fig. 282A). The web extends from the base of the disc of the first toe to the base of the penultimate pha- lanx of the second, from the base of the disc of the second to the distal end of the ante- penultimate phalanx of the third, from the base of the disc of the third to the base of the penultimate phalanx of the fourth and on to the base of the disc of the fifth toe. 596 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 The anal opening is directed posteroven- trally near the upper level of the thighs and is covered by a short anal sheath. The skin is granular on the belly and ventral surfaces of the thighs; other surfaces are smooth. The tongue is cordiform, shallowly notched an- teriorly and posteriorly, and barely free be- hind. There are five to nine (mean, 7.2) pre- vomerine teeth on high transverse ridges be- tween the quadrangular choanae. The vocal sHts extend from the midlateral base of the tongue to the angles of the jaws. The vocal sac is paired, subgular, and greatly distensible. The general coloration of Smilisca hau- dinii is pale green with olive-green markings, olive-green with brown markings, or pale brown with dark brown markings (pi. 70, figs. 4 and 5). The markings on the back consist of irregular spots or blotches. In most specimens a dark interorbital bar is present and usually connected to a large dorsal blotch. The limbs are marked with dark transverse bands, usually three each on the forearm and thigh and three or four on the shank. Trans- verse bands also are present on the tarsi and pro.ximal segments of the fingers and toes. The dorsal markings are usually outlined with black. A dark brown canthal stripe is present. The loreal region and upper lips are pale green or tan; the upper lip usually is boldly marked with vertical dark brown bars. Especially evident is a bar below the eye; the pale area just posterior to this bar is creamy white or ashy gray in some speci- mens. A dark brown or black mark extends from the tympanum to a point above the in- sertion of the arm. In most specimens this mark is broad and distinct, but in some it is restricted to a narrow stripe immediately be- low the posterior part of the supratympanic fold. The flanks are yellow or cream with brown or black mottling; in some specimens the dark mottling encloses pale spots, espe- cially in the groin. The anterior surfaces of the thighs are colored like the flanks, except that the mottling is weaker; the posterior sur- faces of the thighs are brown with small creamy yellow spots. A distinct creamy white anal stripe usually is present. White stripes along the outer edges of the tarsi and fore- arms usually are absent. In most specimens the venter is white, but in specimens from the Atlantic slopes and lo\\lands of Guatemala the belly, especially posteriorly, is yellow. In breeding males the throat is gray. The iris varies from golden bronze to dull bronze with black reticulations. Although considerable variation in color and pattern exists, the variation does not seem to be closely correlated with geography. In specimens from the southern part of the range the dorsal dark markings usually are in the form of small spots, especially on the posterior part of the body. Two specimens from Limon Province, Costa Rica (K.U. No. 34927 from Batan and K.U. No. 36789 from Suretka), lack a dorsal pattern and bands on the limbs. These specimens are nearly uniform brown above with only a few small dark spots on the back. Six specimens (K.U. Nos. 78464, 78466-78470 from 7.3 kilometers southwest of Matatan, Sinaloa, Me.xico) are distinctive in having a uniformly grayish green dorsum with the only dorsal marks be- ing on the tarsi; canthal and post-tympanic dark marks are absent, and a broad white labial stripe is present and interrupted by a single vertical dark mark below the eye. Fur- thermore, a white stripe is present along the outer edge of the foot, and the flanks and posterior surfaces of the thighs are creamy white, boldly marked with black. Throughout most of the range the lips are strongly barred, but in some specimens from southern Nicaragua and Costa Rica the lips are pale, and in a few specimens the vertical bars are indistinct. Two specimens from De- partamento Alta Verapaz, Guatemala (F.M.- N.H. No. 21006 from Coban and U.M.M.Z. No. 90908 from Finca Canihor) differ by hav- ing many narrow trans\erse bands on the limbs and fine reticulations on the flanks. In preservative the dorsum varies from pale bluish gray to brown or tan with darker markings. The yellow spots on the flanks and posterior surfaces of the thighs fade to creamy white. Tadpoles: Ten hatchlings (developmen- tal stage 21) have total lengths of 5.1 to 5.4 (mean, 5.22) mm.; 10 tadpoles in develop- mental stage 38 have total lengths of 35.0 to 37.5 (mean, 35.5) mm. The relative length of the tail to the length of the body increases greatly from the time of hatching until re- 1970 DUELLMAN: HYLID FROGS 597 sorbtion begins at de\elopmental stage 40. The average ratio of tail length to total length in hatchings is 0.495, whereas in stage 38 the ratio is 0.640. A typical tadpole in developmental stage 30 has a total length of 22.3 mm. The body is slightly wider than deep; the snout is round in dorsal and lateral profiles. The nostrils are about midway between the eyes and the tip of the snout. The eyes are widely separated and directed dorsolaterally. The spiracle is sinistral and slightly ventral to the midline, and the spiracular opening is at about mid- length of the body. The mouth is anteroven- tral; the cloacal tube is short and dextral. The caudal musculature is slender, slightly curved upward distally, and does not quite reach the tip of the tail. The dorsal fin extends onto the body and is deepest at about one-third the length of the tail. At midlength of the tail the dorsal fin is slightly deeper than the ventral fin (fig. 284A). The mouth is moderately small and has well-developed lateral folds. The median part of the upper lip is bare; the rest of the mouth is bordered by two rows of labial papillae, except that additional papillae are present in the lateral fold. The upper beak is moderately deep and forms a broad arch with slender lateral processes. The lower beak is more slender and broadly V-shaped; both beaks have blunt serrations. There are two upper and three lower rows of teeth. The two upper rows are about equal in length, and the second row is broadly inter- rupted medially. The three lower rows are complete; the first and second rows are equal in length and slightly shorter than the upper rows, whereas the third lower row is notice- ably shorter. The first upper row usually is sharply curved anteriorlv in the midline (fig. 2S5A). The dorsal part of the body is dark brown with a pale creamy gray, crescent-shaped mark on the posterior edge of the body. The venter is transparent with scattered brown flecks anterolaterally, especially below the eye. The caudal musculature is pale tan with a dark brown longitudinal streak on the mid- dle of the anterior one-third of the tail, dark brown on the dorsal one-third of the tail, and brown flecks and blotches on the rest of the musculature. The fins are transparent with brown flecks and blotches on the entire dor- sal fin and posterior two-thirds of the ventral fin. The iris is bronze. Duellman and Trueb ( 1966 ) noted that the coloration, especially the degree of pig- mentation, is variable in the tadpoles of Smi- lisca baudinii and suggested that the intensity of pigmentation possibly is correlated with the amount of light. Tadpoles from sunlit pools were pallid by comparison with those from shaded forest pools. The authors also noted that the relative length and depth of the tail is variable but could not correlate this variation with geography. Mating Call: The call of Smilisca bau- dinii consists of a series of short, explosive notes, "wonk-wonk-wonk." Two to 15 notes comprise a call group; each note has a dura- tion of 0.09 to 0.13 (mean, 0.11) seconds. Call groups are spaced from 15 seconds to several minutes apart. The notes have 140- 195 (mean, 175), pulses per second and a fundamental frequency of 135 to 190 (mean, 166) cycles per second. Within the frequency spectrum two bands are emphasized; these major frequencies are at about 350 and 2500 cycles per second (pi. 32, fig. 1). Duellman and Trueb (1966) pointed out the existence of an organization in the chorus structure in this species. This was elaborated upon by Duellman ( 1967a ) , who showed that Smilisca baudinii calls in duets; each chorus is made up of several pairs of calling males, and successive choruses apparently are initiated by the same duet. Distress calls that are high pitched and emitted with the mouth open have been heard from both sexes of Smilisca baudinii. Natural History: Throughout most of its range Smilisca baudinii inhabits xeric and subhumid regions having prolonged dry sea- sons. At unfavorable seasons this species takes refuge in bromeliads, in elephant-ear plants, in holes in trees, under bark of trees, and under the outer sheaths of banana plants. Throughout most of its range in Mexico Smilisca baudinii is known to breed from June to October, but on the more humid Caribbean lowlands of Central America it ap- parently has a longer breeding season. Although males call from nearly any body 598 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 of water, including cisterns and buckets, the usual breeding sites arc shallow, temporary pools. Usually the males call from the ground at the edge of the water, but sometimes they sit in shallow water or perch on bushes and trees. Amplexus is axillary. The eggs are spread in a surface film on the water. Each deposition contains several hundred eggs hav- ing a diametei of about 1.3 mm. and encased in a vitelline membrane with a diameter of 1.5 mm. Duellman and Trueb (1966, p. 357) provided counts of 2620, 2940, and 3320 ovu- lated eggs removed from three female frogs. Newly metamorphosed young have snout- vent lengths of 12.0 to 15.5 mm. (mean, 13.4 mm. in 23 specimens). The young usually are white below and dull olive-green above with faint brown transverse bands on the limbs. A white suborbital spot is a distinctive marking on the young of this species. Smilisca baudinii is one of the most abun- dant and conspicuous (by its loud and dis- tincti\e call) of the Middle American hylids. Gadow (1908, p. 76) estimated 45,000 frogs at one breeding site in Veracruz, Mexico, and I have encountered breeding congregations of several hundred, perhaps thousands, of indi- viduals in Mexico, Guatemala, and Costa Rica. Curiously, large numbers of Smilisca baudinii usually are not present at breeding ponds where numerous kinds of other frogs are calling. Instead, calling males of S. bau- dinii usually are at a separate pond. Excep- tions do occur, and large choruses of baudinii have been found with Phnjnohijas venuhsa, Triprion spattdatus reticidatus, Rhinophrynus dorsalis, Engijstomops pustulosus, and Bufo marmoreus. Remarks: Duellman and Trueb (1966, p. 296) discussed the allocation of the various trivial names that are placed in the synonymy of Smilisca baudinii; the type specimens of all of the names proposed have been exam- ined except Hyla vociferans Baird, for which no type was designated. Baird (1859, p. 35) designated figures 11-13 on plate 38 as Hyla vociferans: whether this was a lapsus for Hyla vanvlietii, which he described in 1854, or was intentionally the proposal of a new name can- not be ascertained. The figures quite clearly illustrate the frog now known as Smilisca baudinii. Duellman and Trueb ( 1966, p. 290) erroneously regarded Hyla vociferans Baird as a nomen nudum. However, the rules of zoological nomenclature ( Stoll, 1961, p. 11) clearly state that names based on an illustra- tion, even though not accompanied by a de- scription or designation of a specimen, prior to 1931, are to be regarded as valid. Thus, Duellman and Trueb's designation should be disregarded. Barbour (1923) named Htjla baudini do- lomedes from the Rio Esnape, Darien Prov- ince, Panama. Dunn ( 1931b) first pointed out that the holotype of H. b. dolomedes is a Smilisca phaeota. The cranial osteology of Smilisca baudinii was described bv Duellman and Trueb ( 1966) and by Trueb (' 1968b). Etymology: The specific name baudinii is a patronym for Monseur Baudin, a French commander in Mexico who donated the type specimen to the Museum National d'Histoire Naturelle in Paris. DiSTRiBUTiox : SmUi.sca baudinii has a wide range throughout the lowlands (up to clc\'ations of about 1000 meters) of Middle America from the Rio Grande Embaymcnt of Texas and southern Sonora, Mexico, south- ward to Costa Rica, where on the pacific lowlands the range terminates at the southern limits of the xeric scrub forest in the \icinity of Esparta; on the Caribbean lowlands the distribution apparently is discontinuous south- ward to Surctka (fig. '287). Stuart (1954c, p. 46) recorded the species at ele\'ations up to 1400 meters in southeastern Guatemala, and Duellman and Trueb (1966, p. 298) gave 1600, 1675, and 1925 meters as the highest known ele\'ations for the species in Mexico. Sec Appendix 1 for the locality records of the 3274 specimens examined. Smilisca cyanosticta (Smith) Ilyla phacuta: Smith and Taylor, 1948, p. 88. lUila phaeota cyanosticta Smith, 1953, p. 150 [holotype, U.S.N. M. No. 111147 from Piedras Negras, El Peteii, Guatemala, elevation 100 meters; Hobart M. Smith collector]. Smilsca phaeota (cyanosticta by fiat): Starrett, 1960b, p. 303. Sinih.K-a phaeota cyanosticta: Stuart, 1963, p. 42. SiniU.'^ca cyanosticta: Duellman and Cole, 1965, p. 141. Duellman and Trvieb, 1966, p. 303. 1970 DUELLMAN: HYLID FROGS 599 15° 9° 84^^ ^ 0 200 500 KILOMETERS 27° 21°, 108° 102° 96° 90° 84° Fig. 287. Distribution of Smilisca baudinii. Diagnosis: This moderately large species of Smilisca has a low, flat, elliptical inner metatarsal tubercle, relatively long hind limbs ( the ratio of tibia length to snout-\'cnt length usually is greater than 0.520), and a sloping, moderately long snout. The presence of blue spots on the flanks and posterior surfaces of the thighs, a silvery white labial stripe, and a large brown postorbital mark distinguish Smilisca cyanosticta from all other Middle American hylids. Smilisca sila has blue spots on the flanks and thighs, but it has a short, truncate snout, smaller size (males, 45 mm.; females, 62 mm.), and lacks a white labial stripe and postorbital dark mark. Smilisca phaeota resembles cyanosticta in size, propor- tions, and coloration, except phaeota lacks blue spots. Faded specimens can be identified by probing the lateral edge of the fronto- parietals; large posterolaterally projecting su- praorbital flanges are present in phaeota, whereas the flanges are narrow and not pro- jecting in cyanosticta. Description: Males of this moderately large species attain a maximum snout-vent length of 56 mm., and females reach 70 mm. The largest specimens are from Piedras Ne- gras, EI Peten, Guatemala; seven specimens have snout- vent lengths of 50.1 to 55.7 (mean, 52.5) mm. Specimens from the western part of the range are smaller; the snout-vent length varies from 46.6 to 56.8 (mean, 50.6) mm. in 10 specimens from Los Tuxtlas, Veracruz, and from 44.6 to 55.8 (mean, 50.3) mm. in 23 specimens from northern Oaxaca, Mexico. In a sample of 23 males from between 600 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Yetla and Campamento Vista Hermosa, Oaxa- ca, Mexico, the ratio of tibia length to snout- vent length is 0.519 to 0.597 (mean, 0.563); the ratio of head length to snout-vent length is 0.274 to 0.313 (mean, 0.294), and the ratio of the diameter of the tympanum to that of the eye is 0.644 to 0.788 (mean, 0.718). The aver- age ratio of tibia length to snout-vent length is 0.564 in 10 males from Los Tuxtlas and 0.548 in seven males from Piedras Negras. In these same samples, respectively, the ratio of the diameter of the eye to that of the tym- panum is 0.699 and 0.763. Thus, from west to east there is an increase in snout-vent length and relative size of the tympanum and a de- crease in the relative length of the tibia. Fe- males differ from males by having proportion- ately larger tympani; in four females the ratio of the diameter of the tvmpanum to that of the eye is 0.706 to 0.870 (mean, 0.783). The head is about as wide as the body and is longer than wide; the top of the head is flat. The snout is long and slopes gradually from the eyes to the nostrils, which are about four-fifths the distance from the eyes to the tip of the snout. In lateral profile the snout is acutely rounded, and in dorsal profile it is bluntly rounded. The nostrils are noticeably protuberant. The canthus is round, but dis- tinct; the loreal region is concave, and the lips are moderately thick and flared. A thin su- pratympanic fold extends from the posterior corner of the eye and curves over the upper edge of the tympanum to the insertion of the arm. The tympanum is distinct and separated from the eye by a distance equal to about one-half the diameter of the tympanum. The arms are moderately long; no axillary membrane is present. A row of small tuber- cles is present on the ventrolateral edge of the forearm, and a distinct transverse fold is present on the wrist. The fingers are moder- ately short and broad. The discs are propor- tionately small; the width of the disc on the third finger is equal to about two-thirds the diameter of the tympanum. The subarticular tubercles are large and subconical; the distal tubercle on the fourth finger is flattened and slightly bifid in some specimens. The super- numerary tubercles are large, conical, and usually in one row on the proximal segments of the second, third, and fourth fingers. A large, low, U-shaped outer palmar tubercle is divided into two elongate tubercles in some specimens. The prepollex is moderately en- larged and bears a horny nuptial excrescence in breeding males. The fingers are about one- third webbed (fig. 281B). A trace of web is present between the first and second fingers; the web extends from the base of the penulti- mate phalanx of the second finger to the base of the antepenultimate phalanx of the third to the distal end of the antepenultimate pha- lanx of the fourth finger. The hind limbs are relatively long and slender; the adpressed heels overlap by about one-third the length of the shank, and the tibiotarsal articulation extends to a point between the eye and the tip of the snout. A thin transverse fold is present on the heel. The tarsal fold is thin and extends the full length of the tarsus. The inner metatarsal tubercle is low, flat, and elliptical. The toes are moderately long and slender, and the discs are slightly smaller than those on the hands. The subarticular tubercles are small and round, and the super- numerary tubercles are small, subconical, and in a single row on the proximal segment of each digit. The toes are about three-fourths webbed (fig. 282B). The web extends from the base of the disc of the first toe to the base of the penultimate phalanx of the sec- ond, from the base of the disc of the second to the penultimate phalanx of the third, from the base of the disc of the third to the base of the penultimate phalanx of the fourth and on to the base of the disc of the fifth toe. The anal opening is directed posteroven- trally near the upper level of the thighs and is covered by a short, broad anal sheath. The skin of the belly and posteroventral surfaces of the thighs is granular; the other surfaces are smooth. The tongue is ovoid, barely free behind, and shallowly notched anteriorly and posteriorly. There are four to 11 (mean, 7.1) prevomerine teeth situated on transverse ridges between the small oval choanae. The vocal slits extend from the midlateral base of the tongue to the angles of the jaws. The vocal sac is paired, subgular, and greatly dis- tensible. The general coloration of Smilisca cijano- sticta is pale green or tan with olive-green or dark brown dorsal markings (pi. 70, fig. 3). 1970 DUELLMAN: HYLID FROGS 601 The dorsal markings usually consist of an in- terorbital mark, and a \'-shaped mark in the occipital region with the anterior branches not extending to the eyelids. In many speci- mens this mark is continuous, by means of a narrow middorsal mark, with an in\erted V- shaped mark in the scapular region. In some specimens these dorsal markings are frag- mented into irregular spots, and in some specimens the dorsum is nearly uniform pale green or tan with a few small dark spots. Three or four dark transverse bands arc pres- ent on the thigh and shank, and two or three bands are present on the tarsus. The webbing on the feet is brown. The loreal region is pale green. A dark brown canthal stripe ex- tends from the nostril to the orbit and is bordered above by a narrow bronze-colored stripe, which continues along the edge of the eyelid to a point above the tympanum. The upper lip is silvery or creamy white, and the labial region below the eye is pale green. A broad dark brown mark, extending pos- teriorly from the eye to a point above the insertion of the arm, completely encompasses the tympanum. The flanks are dark brown with many pale blue, round spots, which give the impression of a pale blue ground color with dark brown mottling enclosing spots. The anterior and posterior surfaces of the thighs and the inner surfaces of the shanks and feet are dark brown with many small pale blue spots. Blue spots usually are pres- ent on the proximal segments of the second and third toes. A distinct white stripe is present on the outer edge of the tarsus and fifth toe and on the outer edge of the foreann and fourth finger. The anal region is dark brown and bordered above b\' a narrow trans- verse white stripe. The venter is creamy white; in breeding males the throat is dark grayish brown with white flecks. The iris is golden or bronze above and darker, usually brown, below. Small black flecks are present on the iris, and in some individuals the iris is so heavily flecked so as to appear gray. Although no geographic variation occurs in the dorsal pattern, the pattern on the flanks is variable. Specimens from the eastern part of the range (Piedras Negras and Chinaja, Guatemala) have bold, dark reticulations on the flanks enclosing large pale blue or pale green spots, which fade to tan in preservative. Specimens from Oaxaca and Veracruz, Mex- ico, characteristically have finer dark reticu- lations and smaller blue spots on the flanks; in some of these specimens the ventrolateral spots are smallest and are white. Living individuals have been observed to change from tan to brown, tan to green, pale green to tan, and pale green to dark green. Despite any metachrosis on the dorsum the labial region below the eye remains pale green, and the spots on the flanks and thighs are always present. The ontogenetic change in coloration is striking and proceeds from pale tan flanks and orange yellow thighs, both lacking spots, to pale tan flanks and red thighs, both lacking spots, to dark brown flanks with blue spots and red thighs lacking spots, to dark brown flanks and thighs, both with blue spots. Ju- veniles have a pale tan dorsum with olive- green or dark brown markings; the white la- bial stripe is present. Tadpoles: Ten hatchlings (developmental stage 21) have total lengths of 5.8 to 6.5 (mean, 6.28) mm.; 10 tadpoles in develop- mental stage 36 have total lengths of 27.0 to 30.0 (mean, 28.75) mm. The average ratio of tail length to total length in hatchlings is 0.521, and in stage 36 the ratio is 0.624. A typical tadpole in developmental stage .30 has a total length of 25.0 mm. The body is slightly wider than deep; the snout is rounded laterally and broadly ovoid dorsally. The nostrils are about midway between the eyes and the tip of the snout. The spiracle is sinistral and slightly posterior to the mid- point of the body. The mouth is anteroven- tral, and the cloacal tube is dextral. The caudal musculature is slender, does not ex- tend to the tip of the tail, and is barely curved upward distally. The dorsal fin does not ex- tend onto the body and is slightly deeper than the ventral fin at midlength of the tail (fig. 284B). The mouth is small and has well-devel- oped lateral folds. The median part of the up- per lip is bare, and the rest of the mouth is bordered by one row of bluntly rounded la- bial papillae, except that a few additional pa- pillae are present in the lateral fold. The up- per beak is moderately deep and forms a 602 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 broad arch with slender lateral processes. The lower beak is slender and broadly V-shaped; both beaks are finely serrate. There are two upper and three lower rows of teeth. All of the rows are about equal in length. The sec- ond upper row is broadly interrupted medi- ally; the other rows are complete (fig. 285B). The dorsal part of the body is dark brown; the ventral surfaces are transparent with greenish gold flecks, which disappear in pre- servative. The posterior edge of the body is cream in most specimens. The caudal muscu- lature is gray in life and creamy white with interconnected brown spots in preservative. The caudal fins are transparent with small brown blotches on the dorsal fin and posterior half of the ventral fin. The iris is coppery bronze. Mating Call: The call of Smilisca cyano- sticta consists of one or two moderately short notes, "wonk-wonk." Each note has a dura- tion of 0.25 to 0.45 (mean, 0.38) seconds. Notes are repeated at intervals of about one- half minute to several minutes. The notes have 110 to 180 (mean, 147) pulses per sec- ond and a fundamental frequency of 135 to 160 (mean, 145) cycles per second. Two har- monics are emphasized, one at about 840 cycles per second and another at about 1900 cycles per second (pi. 32, fig. 2). Natural History: Smilisca cyanosticta inhabits humid tropical and lower montane forests. In these moist environments the frogs apparently are active throughout most of the year. Males were calling in Oaxaca in June and July, in Veracruz in June, July, and Au- gust, and in Guatemala in March. Pyburn (1966, p. 2) stated that in Los Tu.xtlas, Vera- cruz, breeding takes place in pools, in the forks of trees, depressions in logs, and in shallow pools. Duellman and Trueb (1966, p. 306) reported males calling from a water- filled depression in a log, in and near springs, in a quiet pool in a stream, and in a rain barrel. The latter authors thought that the eggs were deposited as loose clumps in the water, but Pyburn (1966) reported that the eggs are deposited as a thin surface film. Pyburn (1966, p. 6) stated that the eggs are 1.16 to 1.32 (mean, 1.22) mm. in diameter and are surrounded by a single envelope having a diameter of 1.68 to 2.04 (mean, 1.78) mm. He stated that one captive female laid nine clutches of eggs between September 2, 1962, and October 13, 1963. Six of the clutches (the only ones counted) contained 4.37 to 1844 (mean, 1147) eggs. Duellman and Trueb (1966, p. 357) noted that one female contained 910 ovulated eggs. Pyburn (1966) represented a description of the embryonic and larval development of this species; he found that tadpoles raised in the laboratory required 40 days after hatch- ing to reach metamorphosis at a body length of about 14 mm., the same size given for recentlv metamorphosed young bv Duellman and Trueb (1966, p. 307). Remarks: Smith (1953, p. 150) named cyanosticta as a subspecies of "Hyla phaeota." Superficially the two frogs have much in com- mon, but as demonstrated by Duellman and Trueb (1966) the differences in cranial oste- ology and in the mating calls are highly sug- gestive of specific, rather than subspecific, differences. The most significant cranial chf- ferences are: the presence of a large fonto- parietal fontanelle in cyanosticta and the ab- scence of a fontanelle in phaeota, the pres- ence of large posterolateral-projecting supra- orbital flanges in phaeota as compared with narrow non-projecting flanges in cyanosticta, and the attachment of the nasals to the sphen- ethmoid in cyanosticta and their separation in phaeota. The mating call of cyanosticta has a higher pulse rate and pitch than does that of phaeota. Furthermore, phaeota has only one low emphasized harmonic. Pyburn (1966) discussed this species under the name Hyla phaeota cyanosticta. Etymology: The specific name cyanosticta is derived from the Greek kyanos, meaning dark blue, and stiktos, meaning spotted, and refers to the blue spots on the flanks and thighs. DiSTRiHUTiON: Smilisca cyanosticta inhab- its humid forests on the Atlantic slopes of southern Mexico and northern Central Amer- ica from northern Oaxaca and southern Vera- cruz through northern Chiapas in Mexico and into El Peten and northern Alta Verapaz in Guatemala (fig. 288). The range is discon- tinuous; in southern Mexico the species occurs in humid montane forests at elevations of 830 to 900 meters on the northern slopes of the 1970 DUELLMAN: HYLID FROGS 603 O S. cyanosticta • S- phaeota 12' 0 100 300 I I I I KILOMETERS Fig. 288. Distribution of Smilisca cyanosticta and Smilisca phaeota. Sierra de Juarez and at elevations of 300 to 1200 meters in the Sierra de los Tuxtlas, but it is absent in the intervening lowlands char- acterized by drier forest. The species is known from low elevations in the humid for- ests of El Peten and northern Alta Verapaz, Guatemala, but apparently is absent in the slightly drier forests in the northern part of the Isthmus of Tehuantepec. In addition to the locality records of the 84 specimens examined listed in Appendix 1, Pyburn (1966) reported the species from three localities in the Sierra de Los Tuxtlas, Veracruz, Mexico — 2.7 kilometers south of Coyame, 5 kilometers east of Cuetzalapan, and 4 kilometers south-southwest of Sonte- comapan. Smilisca phaeota (Cope) Hyla phaeota Cope, 1862, p. 358 [holotype, U.S.N.M, No. 4347 from Turbo, Intendencia de Choco, Colombia, sea level; J. Cassin collector]. Boulenger, 1882a, p. 402. Gunther, 1901 (1885-1902), p. 269. Taylor, 1952c, p. 837. Hyla baudini dolomedes Barbour. 1923, p. 11 [holotype, M.C.Z. No. 8539 from Rio Esnape, Sambii X'alley, Darien Province, Panama; Thomas Barbour and Winthrop S. Brooks collectors]. ilyla phaeota phaeota: Smith, 1953, p. 152. SmiUsca phaeota: Starrett, 1960b, p. 303. Duell- man and Trueb, 1966, p. 308. Diagnosis: This large species of Smilisca has a low, flat, elliptical inner metatarsal tu- bercle, relative long hind limbs ( the ratio of tibia length to snout-vent length usually is greater than 0.520), and a sloping, moderately 604 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 long snout. The presence of a white labial stripe and a dark postorbital mark distin- guishes Smilisca phaeota from all other Mid- dle American hylids, except S. cyanosticta. The latter has blue spots on the flanks and on the anterior and posterior surfaces of the thighs, whereas in pluieota the flanks are pale green or tan with fine brown or black vena- tion and the anterior and posterior surfaces of the thighs are pale brown with small cream spots on the posterior surfaces. Smilisca bau- dinii differs from phaeota by having a shorter, more truncate snout, dark, bold mottling on the flanks, and vertical bars on the upper lip. Deschiption: Males of this species attain a maximum snout-vent length of 65 mm.; females reach 78 mm. A considerable dis- crepancy in size occurs in different parts of the range. The average snout-vent length of 10 males from the Canal Zone is 56.5 mm., much the same as that in a sample from the Rio Quesada, Choco, Colombia (56.0 mm.). In equal samples of males from Puerto Viejo, Heredia, Costa Rica, and Bonanza, Zelaya, Nicaragua, the average snout-vent lengths are 51.7 mm. and 43.7 mm. respectively. The largest specimens are from the Golfo Dulce region in Puntarenas Province, Costa Rica, where the average snout-vent length is 61.4 mm. in 10 males. In a sample of 10 males from the Atlantic side of the Canal Zone the ratio of tibia length to snout-vent length is 0.53.3 to 0.598 (mean, 0.578); the ratio of foot-length to snout-vent length is 0.400 to 0.45S (mean, 0.427); the ratio of head length to snout- vent length is 0.323 to 0.367 (mean, 0.349); the ratio of head width to snout-vent length is 0.335 to 0.376 (mean, 0.356), and the ratio of the diameter of the tympanum to that of the eye is 0.651 to 0.855 (mean, 0.749). In a sample of 10 females the only major differ- ence in proportions is that the ratio of the diameter of the tympanum to that of the eye varies from 0.746 to 0.900 (mean, 0.805). The head is about as wide as the body. The snout is moderately long and slopes gradually from the eyes to the nostrils, which arc about four-fifths of the distance from the eyes to the tip of the snout. In lateral profile the snout is acutely rounded, and in dorsal profile it is bluntly rounded. The nostrils are noticeably protuberant. The canthus is round, but distinct; the loreal region is concave, and the lips are moderately thick and flared. A moderately heavy supratympanic fold ob- scures the upper edge of the tympanum and curves downward to the insertion of the arm. The tympanum is distinct and separated from the eye by a distance equal to about one-half the diameter of the tympanum. The arms are moderately long and slender. An axillary membrane is absent. A few small tubercles are present along the ventrolateral edge of the forearm in some specimens; a distinct transverse fold is present on the wrist. The fingers are moderately long and broad. The discs are relatively small; the width of the disc on the third finger is equal to about two-thirds the diameter of the tympanum. The subarticular tubercles ;ire large and round; the distal tubercle on the fourth finger is bifid in some specimens. The supernu- merary tubercles are large and conical. They are in one row on the proximal segment of each digit, except that in some specimens the tubercles are arranged in two irregular rows on the second digit. A flat, tripartite outer palmar tubercle is present. The prepoUex is moderately enlarged and bears a horny nup- tial excrescence in breeding males. The fin- gers are about one-third webbed (fig. 281C). A trace of web is present between the first and second fingers; the web extends from the base of the penultimate phalanx of the second finger to the base of the antepenultimate pha- lanx of the third to the distal end of the ante- penultimate phalanx of the fourth finger. The legs are relatively long and slender; the ad- pressed heels overlap by about one-third the length of the shank, and the tibiotarsal ar- ticulation extends to a point between the eye and the tip of the snout. A thin transverse dermal fold is present on the heel. The tarsal fold is thin and usually extends only about half the length of the tarsus. The inner meta- tarsal tubercle is low, flat, and elliptical. The toes are moderately long and slender; the discs are slightly smaller than those on the fingers. The subarticular tubercles are small and round; the supernumerary tubercles are small, subconical, and in a single row on each toe. The toes are about three-fourths webbed (fig. 282C). The web extends from the base 1970 DUELLMAN: HYLID FROGS 605 of the disc of the first toe to the base of the penultimate phalanx of the second, from the base of the disc of the second to the base of the penultimate phalanx of the third, from the base of the disc of the third to the base of the penultimate phalanx of the fourth and on to the base of the disc of the fifth toe. The anal opening is directed posteroven- trally near the upper level of the thighs and is covered by a short, broad anal sheath. The skin of the belly and posteroventral surfaces of the thighs is granular; the other surfaces are smooth. The tongue is a long ovoid, barely free behind, and not, or only shallovvly notched posteriorly. There are five to nine (mean, 7.3) prevomerine teeth situated on transverse ridges between the small oval choanae. The vocal slits extend from the mid- lateral base of the tongue to the angles of the jaws. The vocal sac is paired, subgular, and greatly distensible. The general coloration of Stnilisca phaeota is pale green or tan with dark olive-green or dark brown dorsal markings (pi. 70, figs. 1 and 2). The dorsal markings usually consist of a dark interorbital mark and a broad blotch extending from the occiput to the sacral region. The dorsal blotch is irregular in shape; in some specimens it is fragmented into an anterior and a posterior blotch or into several spots. In most specimens the mark- ings are bold, but in some the dorsal pattern is so faint as to be barely discernible. Four or five dark transverse bands are present on the thigh, five or six on the shank, and four on the tarsus. Usually two or three narrow bands are present on the proximal part of the fourth toe. The webbing on the feet is brown. The loreal region is pale green and is bordered above by a narrow dark brown canthal stripe extending from the nostril to the orbit. The upper lip is silvery white. A broad dark brown or black mark, extending posteriorly from the eye to a point above the insertion of the arm, completely encompasses the tympanum. The flanks are pale green to creamy tan and are marked with a fine dark brown or black venation. The anterior sur- faces of the thighs are pale brown to grayish tan; in some specimens small darker flecks are present. The posterior surfaces of the thighs are similarly colored with dark flecks present in most specimens and small cream- colored spots present in some individuals. A distinct white stripe is present on the outer edge of the forearm and fourth finger and on the outer edge of the tarsus and fifth toe; tlie latter stripe is bordered below by dark brown on the tarsus. The anal region is dark brown and usually bordered above by a nar- row, transverse creamy white stripe. The venter is creamy white. In breeding males the throat is dark gray. The iris is bronze, darkest medially, and marked with fine black reticulations. The only significant geographical varia- tion in coloration is the presence of a faint tint of pale blue on the flanks in specimens from the Caribbean lowlands of Nicaragua and northeastern Costa Rica. Living individ- uals are capable of changing color from green to brown, or reverse. Recently metamorphosed young usually are pale tan with brown on the sides of the head and on the flanks. The brown is sepa- rated from the dorsal color by a narrow cream stripe, which disappears when indi- viduals reach a snout-vent length of about 20 mm. Also, at that stage of growth the dark pigment of the flanks dissipates into the finely venate pattern of the adults. Tadpoles: Three hatchlings (develop- mental stage 21) have total lengths of 7.9 to 8.6 (mean, 8.21) mm. and an average ratio of tail length to total length of 0.477; in tad- poles in de\elopmental stage 36 the ratio is 0.613. Tadpoles reach their maximum size at stage 39 when they have a body length of 14.0 mm. and a total length of as much as 39.8 mm. A detailed description of larval development was presented by Duellman and Trueb (1966). A typical tadpole in developmental stage 30 has a total length of 22.9 mm. The body is as wide as deep; the snout is round in dor- sal and lateral profiles. The nostrils are about midway between the eyes and the tip of the snout. The eyes are widely separated and directed dorsolaterally. The spiracle is sinis- tral and slightly ventral to the midline, and the spiracular opening is at about the mid- length of the body. The mouth is anteroven- tral; the cloacal tube is short and dextral. The caudal musculature is slender, slightly curved 606 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 upward distally, and does not reach the tip of the tail. The dorsal fin extends onto the body and is deepest at about one-third of the length of the tail. At midlength of the tail the dorsal fin is slightly shallower than, or equal in depth to, the ventral fin (fig. 284C). The mouth is moderately small and has well-developed lateral folds. The median part of the upper lip is bare; the rest of the mouth is bordered by one row of labial papillae. Additional papillae are present in the lateral fold. The upper beak is moderately deep and forms a broad arch with slender lateral processes. The lower beak is more slender and broadly V-shaped; both beaks have blunt serrations. There are two upper and three lower rows of teeth. The second upper row is slightly shorter than the first and broadly interrupted medially. The three lower rows are complete. The rows are about equal in length and slighUy shorter than the second upper row (fig. 285C). The dorsal part of the body is pale brown with a pale cream crescent-shaped mark on the posterior edge of the body. The belly is transparent with scattered brown flecks. The caudal musculature is pale creamy tan with brown spots. The fins are transparent with brown flecks and blotches. The iris is pale bronze. Mating Call: The call of Smilisca phae- ota is a low vibrant growl. Each cafl group consists of one or two notes having a dura- tion of 0.10 to 0.45 (mean, 0.31) seconds. Call groups are repeated at intervals of 20 seconds to several minutes. The notes have 100 to 130 (mean, 116) pulses per second and a fundamental frequency of 110 to 165 (mean, 143) cycles per second. Only one harmonic within the frequency spectrum is emphasized; this dominant frequency is at 330 to 495 (mean, 372) cycles per second (pi. 32, fig. 3). Natural History: Throughout most of its range Smilisca pliaeota inhabits humid lowland tropical forest. Because of rather equable climatic conditions, frogs of this spe- cies are active throughout the year. Although breeding activity is highest in the rainy sea- son, slight showers in the drier parts of the year stimulate males to call. Males usually call from secluded spots at the edge of, or in, shallow temporary pools; occasionally indi- viduals are found at the edges of streams or large ponds. Duellman and Trueb (1966) reported that the eggs are deposited in loose clumps amidst vegetation. Subsequent observations indicate that probably the eggs are normally deposited in a surface film. Three females contained 1665, 1870, and 2010 ovulated eggs (Duell- man and Trueb, 1966). Recently metamor- phosed young have snout-\ent lengths of 12.7 to 16.7 mm. (mean, 14.3 mm. in 11 speci- mens). Smilisca phaeota, although not extremely abundant, is one of the frequently encoun- tered hylids in lower Central America. Its habit of calling throughout the year in small temporary pools, often in the immediate vicinity of human habitation, make it one of the best known frogs to local people. Remarks: Dunn (1931b, p. 413) sug- gested that //(//« baudinii dolomedes Barbour (1923) from Darien Province, Panama, was actually Htjla phaeota. Smith (1953) de- scribed Hyla phaeota cyanosticta from Pie- dras Negras, Guatemala. Duellman and Trueb (1966) concurred with Dunn's assignment of dolomedes but demonstrated that on the basis of cranial osteology and characteristics of the tadpoles and mating calls cyanosticta was not conspecific with phaeota. Wilhelm Peters (1863, p. 463) named Hyla lahialis from "umgegend von Bogota," Cundinamarca, Colombia, but in 1874 he re- garded Hyla lahialis to be identical with Hyla phaeota Cope, 1862. Giinther Peters informed me that the holotype of Hyla lahialis could not be found as of January 5, 1965, but that it was catalogued as number 4913 in the Zoologisches Museum Berlin. In the supposed absence of a type specimen of Hyla lal>ialis, Duellman and Trueb (1966) followed Peters' decision in 1874 that his Hyla lahialis was conspecific with Hyla phaeota Cope. In the summer of 1969 I found the t>'pe of Hyla lahialis Peters in the Zoologisches Museum Berlin. The specimen (ZMB 4913) is not a Smilisca phaeota. The type specimen is the same as the Andean frogs subsequently named Hyla vilsoniana by Cope (1899). Etymology: The specific name phaeota apparently refers to the dark markings on the 1970 DUELLMAN: HYLID FROGS 607 dorsum and is dcri\'cd from the Greek phaios meaning dark or dusky. Distribution: Smilisca pliaeota is wideh' distributed below elevations of about 1000 meters in lower Central America (fig. 288). On the Caribbean lowlands it ranges from northeastern Nicaragua to northwestern Co- lombia and inland in the valleys of the Rio Cauca and Rio Magdalena; the species oc- curs on the Pacific lowlands from south-cen- tral Costa Rica to northwestern Ecuador, ex- clusive of the Panamanian savannas and the Azuero Peninsula. See Appendix 1 for the locality records of the 581 specimens examined. Smilisca puma (Cope) Hyla pinna Cope, 1885b, p. 183 [holotype, U.S.N. M. No. 13735 from "Nicaragua"; J. F. Moser collector]. Gunther, 1901 ( 1885-1902), p. 270. Hyla weUmanorum Taylor, 1952c, p. 843 [holo- t>-pe, K.U. No. 30302 from Batan, Limon Province, Costa Rica, elevation 15 meters; Edward H. Taylor collector]. Smilisca weUmanorum: Starrett, 1960b, p. 303. Smilisca puma: Duellman and Trueb, 1966, p. 314. Diagnosis: This small species is easily distinguished from other members of the genus by the lack of webbing on the hand. The toes are about one-half webbed; the diameter of the tympanum is about two- thirds of that of the eye. A narrow white labial stripe is present. The dorsum is tan with a pair of dark brown (sometimes inter- connected) longitudinal stripes on the back. The subarticular tubercles on the hand are relatively large, and the inner metatarsal tu- bercle is small. No other species of Smilisca has a pattern tending toward longitudinal stripes on the dorsum or has essentially no webbing in the hand. Description: Smilisca puma is the small- est species in the genus; males attain a maxi- mum snout-vent length of 38 mm., and fe- males reach 46 mm. In a sample of 10 males from Puerto Viejo, Heredia Province, Costa Rica, the snout-vent length is 32.5 to 37.9 (mean, 34.8) mm. The ratio of the tibia to the snout-vent length is 0.484 to 0.529 (mean, 0.512); the ratio of foot length to snout-vent length is 0.375 to 0.426 (mean, 0.406); the ratio of the head length to the snout-vent length is 0..355 to 0.386 (mean, 0..373); the ratio of the head width to the snout-vent length is 0.346 to 0.378 (mean, 0.361), and the ratio of the diameter of the tympanum to that of the eye is 0.521 to 0.718 (mean, 0.647). The head is nearly as wide as the body and slightly narrower than wide. The top of the head is flat. In dorsal profile the snout is pointed; in lateral profile the snout is bluntly rounded. The snout is moderately long. The nostrils are noticeably protuberant and are situated at about three-fourths of the distance from the eyes to the tip of the snout. The canthus is rounded but distinct; the loreal region is noticeably concave, and the lips are thin and moderately flared. A thin dermal fold extending posteriorly from the corner of the eye to a point above the insertion of the arm conceals the upper edge of the tym- panum. The tympanum is otherwise distinct and separated from the eye by a distance about equal to the diameter of the tympanum. The arm is moderately short; the upper arm is rather slender, and the forearm is no- ticeably robust. No axillary membrane is present. No distinct row of tubercles or der- mal fold is present on the V'entrolateral edge of the forearm, but a distinct transverse fold is present on the wTist. The fingers are short and stout and bear moderately large discs. The width of the disc on the third finger is about equal to the diameter of the tympan- um. The subarticular tubercles are large and round; in a few individuals the distal tubercle on the fourth finger is slightly bifid. Supernu- merary tubercles are absent except on the proximal part of the third and fourth fingers in some specimens; in these the tubercles are small and indistinct. The palmar tubercle is low, usually flat, and in most specimens rather indistinct. In some individuals the tubercle is bifid, tripartite, or fragmented into three or four small tubercles. The prepollex is barely enlarged, breeding males lack nuptial excres- cences. Webbing is absent between the first and second fingers and vestigial between the others (fig. 281D). The hind limbs are mod- erately short and slender; the adpressed heels overlap by about one-fourth of the length of the shank, and the tibiotarsal articulation extends to the eye. A thin tarsal fold extends from between two-thirds to the full length of the tarsus. The inner metatarsal tubercle 608 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 is small, low, flat, and elliptical. The toes are moderately long and slender; the discs are about the same size as those on the fingers. The subarticular tubercles are moderately small and round; supernumerary tubercles are present only on the basal segments of the fourth and fifth digits. The toes are slightly more than one-half webbed (fig. 282D). The web connects the first and second toes at the bases of the penultimate phalanges; the web extends from the middle of the penultimate phalanx of the second to the middle of the antepenultimate phalanx of the third, from the middle of the penultimate phalanx of the third to the base of the antepenultimate of the fourth and on to the middle of the penul- timate phalanx of the fifth toe. The anal opening is directed posteriorly at the upper level of the thighs and is cov- ered by a short anal sheath. The skin is gran- ular on the belly and the posteroventral sur- faces of the thighs; other surfaces are smooth. The tongue is cordiform, usually shallowly notched anteriorly and deeply notched poste- riorly, and barely free behind. There are four to seven (mean, 5.3) provomerine teeth on high transverse ridges situated at a level between the posterior borders of the small round choanae. The vocal slits extend from the midlateral base of the tongue to the angles of the jaws. The vocal sac is paired, subgular, and greatly distensible. The general coloration of Smilisca puma is yellowish-tan with brown markings on the dorsum (pi. 71, fig. 5). The markings on the back usually consist of a pair of dorsal stripes, variously modified. In some specimens the stripes are discreet and extend from the post- orbital region nearly to the vent, but in some specimens the stripes are connected by a transverse mark in the scapular region and in many others also by a transverse mark in the sacral region. In some specimens the stripes are fragmented posteriorly, and in one individual the dorsal pattern consists of two series of dark longitudinal dashes. Another specimen has two stripes fused middorsally for nearly their entire lengths. A dark brown interorbital mark is present; in most speci- mens this is in the form of an interorbital bar that extends onto the eyelids, but in some .specimens the mark consists of a short V- shaped mark or small spot between the eyes. There is no dark post-tympanic mark, but dark brown pigment forms a venated pattern from the axilla to the midflank. The inguinal region is white, finely mottled with dark brown. Some specimens have scattered me- tallic green flecks on the dorsum. The dorsal surfaces of the hind limbs are colored like the body and have two or three dark brown transverse marks on the thighs, three to five marks on the shanks, and one or two marks or irregularly arranged dark flecks on the tarsi. The posterior surfaces of the thighs are dark brown, and the webbing of the feet is tan to grayish brown. A narrow white stripe is present on the edge of the upper lip, and a transverse white stripe above the anus is invariably present. Narrow white stripes on the outer edges of the tarsi and of the fore- limbs usually are distinct. The belly and xen- tral surfaces of the limbs are creamy white. In breeding males the vocal sac is grayish brown. The iris is a deep bronze. T.A.DPOLES: No recently hatched tadpoles or late developmental stages are available. Four tadpoles in developmental stage 34 have body lengths of 9.0 to 9.5 mm. and total lengths of 23.0 to 24.5 mm. The largest tad- pole examined is in developmental stage 40 and has a total length of 31.0 mm. In a tad- pole in developmental stage 34, the body is about three-fourths as deep as wide; the snout is round in dorsal and lateral profile. The nostrils are about midway between the eyes and the tip of the snout; the eyes arc widely separated and directed dorsolaterally. The spiracle is sinistral and slightly ventral to the midline, and the spiracular opening is at about two-thirds of the length of the body. The mouth is anteroventral; the cloacal tube is short and dextral. The caudal musculature is slender and barely curved upward distally, and does not quite reach the tip of the tail. The dorsal fin extends onto the body and is deepest at about two-thirds of the length of the tail, where its depth is only slightly more than that of the ventral fin (fig. 284D'). The mouth is moderately small and has well-developed lateral folds. The median part of the upper lip is bare; the rest of the mouth is bordered by one or two rows of labial pa- pillae plus additional papillae in the lateral 1970 DUELLMAN: HYLID FROGS 609 fold. The upper beak is shallow and forms a high arch with slender lateral processes. The lower beak is equally slender and broad- ly V-shaped; both beaks are finely serrate. There are two upper and three lower rows of teeth. The two upper rows are about equal in length, and the second row is broadly inter- rupted medially. The lower rows are com- plete. The first and second lower rows are about equal in length and nearly as long as the upper row, whereas the third lower row is noticeably shorter (fig. 285D). The body is olive-brown with silvery green flecks laterally. The caudal musculature is oHve-brown with greenish tan flecks. The fins are pale brown with greenish gold flecks. Dark reticulations are present on the caudal musculature and on the adjacent parts of the fins on the anterior half of the tail. The iris is deep bronze. Mating Call: The call of Smilisca puma consists of a low squawk, usually followed by a series of one or more rattling secondary notes. Call groups are spaced at intervals of five to 55 seconds. The duration of the pri- mary notes varies from 0.06 to 0.35 (mean, 0.13) seconds, and that of the secondary notes is 0.10 to 0.47 seconds. The primary notes have 187 to 240 (mean, 208) pulses per sec- ond and have fundamental frequencies of 125 to 200 (mean, 145) cycles per second. Within the frequency spectrum two bands are emphasized; these major frequencies are at about 740 and 1870 cycles per second (pi. 33, fig. 1). Duellman and Trueb (1966) noted that although individuals of Stnili^ca puma some- times call alone, duets, trios, or quartets were more common. They observed that the chorus is initiated by one individual uttering primary notes until joined by a second, third, and fourth frog. Natural History: Smilisca pinna inhabits humid lowland tropical forest having more or less evenly distributed rainfall throughout the year. Except for periodic dry spells, frogs of this species seem to be active throughout most of the year. Calling males have been collected from February through September, and gravid females have been found in June, July, and August. Males call from shallow water, where they are usually well hidden in the bases on dense clumps of grass. One recently metamorphosed individual has a snout-vent length of 12.4 mm. Remarks: Comparison of the holotype of Hyla weUmanorum Taylor (K.U. No. 30302) with the holotype of Hyla puma Cope (U.S. N.M. No. 13735) leaves no doubt that both of these names apply to the same species. The type specimen of puma was part of a collection received at the United States Na- tional Museum from Lieutenant J. F. Moser from "Nicaragua." Duellman and Trueb (1966, p. 317) noted that on the basis of other species in the collection received from Moser, it is most likely that the holotype of Smilisca puma originated from the Caribbean lowlands of southeastern Nicaragua. How- ever, to this date no specimens bearing spe- cific locality have been received from Nica- ragua, although the species is common in the Caribbean lowlands of Costa Rica. Cochran (1961) Hsted Hyla puma Cope, as a synonym of Hyla molitor O. Schmidt, 1857. On the basis of Schmidt's description of molitor and a supposed syntype (N.M.W. No. 16494) it is inconceivable that puma and molitor are the same. Etymology: The specific name puma seemingly is an Indian name for a cat, from which is derived the \ernacular name for Felis concolor. Possibly Cope used this name in elusion to the tawny dorsal color of the frog, which is not unlike that of the puma. Distribution: This species lives in the wet, forested region of the Caribbean low- lands of Costa Rica and presumably southern Nicaragua (fig. 289). All specimens are from low elevations; the highest record of occur- rence of this frog is 285 meters at Laguna Bonilla. See Appendix 1 for the locahty records of the 65 specimens examined. Smilisca sila Duellman and Trueb Smilisca sila Duellman and Trueb, 1966, p. 318 [holotype, K.U. No, 91852 from El Volcan, Chiriqui Province, Panama, elevation 1280 meters; William E. Duellman collector]. Diagnosis; This moderate-sized member of the genus differs from all other species by having a short truncate snout and in lacking 610 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fig. 289. Distribution of SmiUsca puma. a dark brown or black postorbital mark. The lips are thick and rounded, and the diameter of the tympanum is about one-half that of the eye. The margin of the upper lip is faintly marked by an interrupted white stripe. The flanks and posterior surfaces of the thighs are dark brown or black with pale blue to creamy tan spots or flecks. Blue spots are present on the flanks of S. cyanosticta and some S. sordicla. The former is a larger (males to 56 mm.; females to 70 mm.) species having a longer, more sloping snout, and a dark brown postorbital mark. The snout is low and sloping in S. sordicla. The lips are thin and flaring, and the throat in breeding males is white, whereas the throat in .sila is dark brown. SmiUsca haudinii is the only other genus having a moderately short and truncate snout, but this species is large ( males to 76 mm.; females to 90 mm.); furthermore, haudinii has a dark postorbital mark and has creamy yellow flanks with black or brown mottling. Description: SmiUsca sila is a moderate- sized species of the genus; males attain a maximum snout-vent length of 45 mm., and females reach 62 mm. In a sample of 10 males from Finca La Sumbadora, Panama Province, Panama, the snout-\'ent length is 40.0 to 44.8 (mean, 42.3) mm. The ratio of the tibia length to the snout-vent length is 0.511 to 0.56(S (mean, 0.540); the ratio of the foot length to snout-vent length is 0..376 to 0.4.39 (mean, 0.411); the ratio of head length to snout-\ent lengtii is 0..326 to 0..356 (mean, 0.344); the ratio of head width to snout- vent length is 0.337 to 0.368 (mean, 0..352), and the ratio of the diameter of the tympanum to that of the eye is 0.4S1 to 0.580 (mean, 0.5.32). There is a geographic gradient in size; 1970 DUELLMAN: HYLID FROGS 611 specimens from the western part of tlie range (southern Costa Kica) are smaller than those in the eastern part of the range (eastern Panama). Fixe males from the Pacific low- lands of southern Costa Rica have snout-vent lengths of 31.6 to 38.2 (mean, 34.7) mm.; 10 males from El Volcan, Chiriqui Province, Panama, have snout-vent lengths of 32.6 to 37.9 (mean, 36.4) mm., and eight males from Barro Colorado Island, Canal Zone, have snout-vent lengths of 38.2 to 42.0 (mean, 35.6) mm. These are smaller than the males from Finca La Sumbadora, which is east of the Canal Zone. Ten females from El Volcan have snout-vent lengths of 44.2 to 55.6 ( mean, 49.2) mm., as compared with 56.1 to 62.2 (mean, 58.2) mm. in three females from Finca La Sumbadora. The head is about as long as broad and is as broad as the body. The top of the head is flat. In dorsal and lateral profiles the snout is truncate. The snout is extremely short. The diameter of the eye is nearly equal to the distance from the anterior corner of the eye to the tip of the snout. The nostrils are mod- erately protuberant and are situated at about three-fourths the distance from the anterior corner of the eye to the tip of the snout. The canthus is angular; the loreal region is slightly concave, and the lips are thick and barely flared. A moderately heavy dermal fold ex- tends posteriorly from the posterior corner of the eye, above the tympanum, and curves downward to the place of insertion of the arm. The upper edge of the tympanum is con- cealed beneath the dermal fold. Otherwise the tympanum is distinct and separated from the eye by a distance about equal to the diam- eter of the tympanum. The arm is rather short; the upper arm is slender, and the forearm is moderately ro- bust. No axillary membrane is present. A row of low, indistinct tubercles is present on the ventrolateral edge of the forearm, and an in- distinct transverse fold is present on the wrist. The fingers are moderately long and stout and bear rather small discs. The width of the disc on the third finger is about two-thirds of the diameter of the tympanum. The subartic- ular tubercles are moderately large and coni- cal; none is bifid. The supernumerary tuber- cles are small and indistinct; they are present only on the proximal segments of the second, third, and fourth fingers. A flat, indistinct, triangular shaped palmar tubercle is present. The prepollex is moderately enlarged and in breeding males bears a horny nuptial excres- cence. The fingers are about half webbed (fig. 283A). A trace of web exists between the first and second fingers. The web extends from the middle of the penultimate phalanx of the second finger to the proximal end of the antepenultimate phalanx of the third, and from the distal end of the antepenultimate plialanx of the third to the base of the penul- timate phalanx of the fourth finger. The hind limbs are moderately long and slender; the adpressed heels overlap by about one-fourth of the length of the shank, and the tibiotarsal articulation extends to a point between the eye and the nostril. The tarsal fold is thin and flap-like, and extends the entire length of the tarsus. The inner metatarsal tubercle is low, flat, and elliptical. The toes are moderately long and slender; the discs are slightly smaller than those on the fingers. The subarticular tubercles are large and subconical; the super- numerary tubercles are moderately large, con- ical, and in a single row on the proximal segment of each toe. The toes are about four- fifths webbed (fig. 283C). The web extends from the base of the disc of the first toe to the middle of the penultimate phalanx of the second, from the base of the disc of the sec- ond to the base of the penultimate phalanx of the third, and from the base of the disc of the third to the base of the antepenultimate phalanx of the fourth and on to the base of the disc of the fifth toe. The anal opening is directed posteroven- trally near the upper level of the thighs and is covered by a short anal sheath. Large fe- males from throughout the range and some males from Costa Rica and western Panama have scattered small tubercles on the head and back. In other specimens, the dorsal surfaces are smooth. The belly and postero- \entral surfaces of the thighs are granular. The tongue is broadly cordiform, very shal- lowly notched posteriorly, and barely free behind. There are five to seven (mean, 5.7) pre\'omerine teeth on high, rounded, trans- verse ridges between the posterior margins of the small, ovoid inner naries. The vocal slits 612 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 extend from the midlateral base of the tongue to the angle of the jaws. The \'Ocal sac is paired, subgular, and greatly distensible. The general coloration of Smili.sca sila consists of a gray, tan, or pale reddish brown dorsal ground color and a creamy white ven- ter. The dorsum is marked by dark brown, olive-brown, or dark reddish brown spots or blotches (pi. 71, figs. 3 and 4). Usually the blotches are discreet, but in some individuals they are interconnected and form an irregular dark mark on the dorsum. There is no ten- dency for the blotches to form transverse bars as in some Smili.sca sordida. In some fe- males the dorsal markings are reduced to a few small spots or are nearly absent, whereas in other females the dorsal markings are bold. White, pustular spots or metallic green flecks are present on the dorsal surfaces of many individuals. The dorsal surfaces of the limbs are colored like the body with dark brown transverse bars; usually three or four bars are present on each forearm, thigh, and shank, usually the flanks and posterior surfaces of the thighs have black mottling enclosing pale blue spots and flecks respectively, but the col- oration of the flanks and limbs varies geo- graphically. Specimens from southern Costa Rica and western Panama have distinct bars on the limbs; the posterior surfaces of the thighs have brown reticulations enclosing small blue flecks in specimens from Costa Rica and bolder, black reticulations enclosing large pale blue spots in specimens from western Panama. In specimens from Costa Rica the flanks are brown with pale blue flecks, whereas those from Chiriqui, Panama, the flanks are pale blue with dark brown mottling in the inguinal region. Frogs from El Valle and Cerro La Campana usually have distinct bars on the limbs; the posterior surfaces of the thighs are colored as in frogs from Chiriqui, and the inguinal region is pale blue with coarse brown mottling. Specimens from Barro Colorado Is- land, are marked like those from El Valle and Cerro La Campana, except that on the posterior surfaces the thighs fine black re- ticulations enclosed many pale blue spots. In specimens from Darien and Panama prov- inces, east of the Canal Zone (Altos de Pa- cora, Cerro Jefe, Finca La Sumbadora, and Rio Pacora), the markings on the dorsal sur- faces of the limbs are indistinct or absent in males, but distinct in some females. Intense brown and black pigment forms fine reticula- tions delimiting bold blue spots on the flanks. This coloration extends to the axilla in many specimens. Fine black reticulations enclose many dark blue spots on the posterior sur- faces of the thighs. In living individuals from Costa Rica and western Panama the blue coloration on the flanks and thighs is much less conspicuous than in specimens from east- ern Panama. In females the throat is creamy white; in some specimens scattered brown flecks are present on the chin and throat. In breeding males the anterior part of the throat is dark gray or dark brown. The color of the iris is variable, even in frogs from one locality. The color varies from pale brown to grayish brown with or without a metaUic bronze suf- fusion and dark brown or black reticulations. The labial region is usually indistinctly marked by dark vertical bars separated by paler ground color. The edge of the upper lip is marked by a creamy white stripe which is broadly interrupted by the vertical dark bars. In many specimens the labial stripe is nearl\- indistinguishable. A recently metamorphosed young had, in life, a brown dorsum with darker brown markings, a white spot below the eye and a narrow white labial stripe. The belly was white; the flanks were brown with white spots and the posterior surfaces of the thighs were yellov\ish tan. Tadpoles: Eleven tadpoles in develop- mental stage 25 have total lengths of 25.9 to 31.0 (mean, 28.1) mm.; one tadpole in stage 42 has a total length of 42.0 mm. A typical tadpole in developmental stage 25 has a total length of 28.5 mm. The body is only slightly wider than deep and nearly flat dorsally; the snout is broadly rounded in dorsal \icw and bluntly rounded in lateral view. The nostrils are slightly closer to the eyes than to the tip of the snout. The eyes are widely separated and directed dorsolaterally. The mouth is ventral; the cloacal tube is short and dextral. The spiracle is sinistral and slightly ventral to the midline, and the spiracular opening is at about two-thirds of the distance from the snout to the posterior edge of the body. The 1970 DUELLMAN: HYLID FROGS 613 caudal musculature is moderately heavy and straight; the musculature extends to the tip of the tail. The dorsal fin extends onto the body; the fins are deepest at about two-fifths of the length of the tail, where the depth of the caudal musculature is about equal to the depth of the dorsal and the depth of the ven- tral fin (fig. 284E). The mouth is moderately large and has extensive lateral folds. The median part of the upper lip is bare; the rest of the upper lip is bordered by one row of labial papillae; and the lower lip is bordered by one or two rows of labial papillae. Many small papillae are present in the lateral folds. The upper beak is moderately massive, and its inner sur- face forms a continuous arch with the short lateral processes. The lower beak is less ro- bust and is broadly V-shaped; both beaks bear blunt serrations. There are two upper and three lower rows of teeth. The upper rows are about equal in length and broadly V-shaped. The second upper row is narrowly interrupted medially. The lower rows are complete and about equal in length, but slightly shorter than the upper rows (fig. 285E). In preservative the dorsal part of the body is dark grayish brown with dark brown spots dorsally and white flecks laterally; the venter is pale grayish tan. The caudal musculature is pale tan with brown flecks over the entire surface and dark brown streaks on the pos- terior half of the ventral fin and on all of the dorsal fin. M.\TiNG Call: The call of Smilisca sila consists of a low squawk, usually followed by a series of one or more rattling secondary notes. Call groups are repeated at intervals of four to 20 seconds. The duration of the primary notes is 0.06 to 0.28 (mean, 0.16) seconds, and of the secondary notes, 0.14 to 0.48 sec- onds. The primary notes have 97 to 120 (mean, 108) pulses per second and a funda- mental frequency of 90 to 115 (mean, 103) cycles per second. Two bands are empha- sized within the frequency spectrum; these major frequencies are at about 900 and 2200 cycles per second (pi. 33, fig. 2). N.^TURAL History: Smilisca sila is an in- habitant of shallow rocky streams. The breed- ing season seems to be correlated with the time of the year when the water is clear and at a low level; consequently, the major breed- ing activity takes place in the dry season. Males call from the edges of small, shallow streams, from rocks in the stream or less frequently from vegetation overhanging the streams. Females are most frequently found on the banks of streams, and clasping pairs are usually in shallow pools in streams. Tad- poles have been found in pools in clear streams; some tadpoles have been observed to cling by their mouths to rocks in the streams; others were found on the bottom where they seek refuge among the pebbles or under rocks or leaves. Metamorphosing young have been found on vegetation at the edges of streams and have been raised at the laboratory. Seven recently metamorphosed young have snout- vent lengths of 13.6 and 15.6 (mean, 14.6) mm. Remarks: Duellman and Trueb (1966) demonstrated that this species is distinct from Smilisca sordida; both species had been con- fused under the name of Hyla (Smilisca) gabbi. With the exception of the type de- scription of Smilisca sordida, all references to "Hyla sordida" and "Hyla gabbi" in Pana- ma are based on Smilisca sila. Etymology: The specific name sila refers to the blunt snout and is derived from the Latin silus meaning "pug-nosed." Distribution: Smilisca sila ranges along the Pacific slopes and lowlands of Costa Rica and Panama at elevations from sea level to about 1300 meters; in eastern Panama and northern South America the species occurs on the Caribbean slopes and in the valleys of the northward draining rivers of Colombia (fig. 290). See Appendix 1 for the locality records of the 270 specimens examined. Smilisca sordida (Peters) Hyla sordida Peters, 1863, p. 460 [syntypes, Z.M.B. 3141 (2 specimens) from "Veragiias," Panama; J. von Warzewicz collector]. Brocchi, 1882, p. 42. Boulen- ger, 1882a, p. 393. Giinther, 1901 (1885-1902), p. 273. Hyla gabbi Cope, 1876, p. 103 [synty-pes, U.S.N.M. Nos. 30658 and 30659 from "near Sipurio," Limon Pro\ince, Costa Rica, elevation 60 meters; William M. 614 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 10° 82° 78° 10° v. 1 y-^'\ t, • N , y? \ r •-. - 8° 0 50 100 vl, A . ■> M'" KILOMETERS 82° 78° Fig. 290. Distribution of Sinilisca sila. Gabb collector], Brocchi, 1882, p. 37. Boulenger, 1882a, p. 372. Gunther, 1901 (1885-1902), p. 274. Taylor, 1952c, p. 840. Hyla nigripes Cope, 1876, p. 104 [syntypes, U.S.N.M. Ncs. 30685 and 30686 from "Pico Blanco," Limon Province, Costa Rica; William M. Gabb collec- tor]. Brocchi, 1882, p. 38. Boulenger, 1882a, p. 394. Gunther 1901 (1885-1902). p. 278. Taylor, 1952c, p. 853. Htjla salcit^i Boulenger, 1882a, p. 372 [syntypes B.M.N.H. No. 1947.2.24.12 from Costa Rica; Osbert Salvin collector; B.M.N.H. No. and 1947.2.24.13 from Cartage Province, Costa Rica, elevation 1470 meters; Jansen collector]. Smilisca gabbi: Starrett, 1960b, p. 303. Smilisca sordida; Duellman and Trueb, 1966, p. 323. Diagnosis: This moderate-sized member of the genus is distinguished from the other species by the presence of a white vocal sac in breeding males. The diameter of the tym- panum is about one-half that of the eye, and the lips are thin and flaring. The inner meta- tarsal tubercle is long, low, flat, and elliptical. The fingers are about one-half webbed; the toes are four-fifths webbed. The dorsum is variously marked with dark gray, dark brown, reddish-brown, or olive-green spots or blotch- es. The flanks and posterior surfaces of the thighs are dark brown with bluish white and creamy tan flecks respectively. There is no white labial stripe. Blue spots are present on the flanks of S. cijanosticta and sila. The for- mer is a larger species (males to 56 mm.; females to 70 mm.) having a white labial stripe and a large dark brown postorbital mark. In S. sih the snout is short and trun- cate, the lips are thick and not flaring, and the throat in breeding males is dark gray or brown. Description: Males of Smilisca sordida attain a maximum snout-vent length of 45 mm., and females, 64 mm. In a sample of 10 adult males from 15 to 20 kilometers west- southwest of San Isidro el General, San Jose Province, Costa Rica, the snout-vent length is 38.1 to 42.6 (mean, 40.5) mm.; the ratio of tibia length to snout-vent length is 0.505 to 0.5.38 (niean, 0.523); the ratio of foot length to snout-vent length is 0.406 to 0.440 (mean, 0.426 ) ; the ratio of head length to snout-vent length is 0.329 to 0.351 (mean, 0.343); the ratio of head width to snout-vent length is 0.291 to 0.322 (mean, 0.313), and thc^ ratio of the diameter of the tympanum to that of the eye is 0.449 to 0.571 (mean, 0.489). Speci- mens from the Pacific slopes of Costa Rica are larger than those from the Meseta Central 1970 DUELLMAN: HYLID FROGS 615 TABLE 5S Comparison of Snout-\ent Lengths, with Means in Parentheses, in six Samples of Males of Smilisca sordida from Costa Rica. Locality N Snout-vent Length Puntarenas : Golfito 10 38.4-44.6 (41.8) Puntarenas: Rincon de Osa 20 38.3-42.1(40.8) San Jose: San Isidro el General 10 38.1-42.6 (40.5) San Jose: Escazii and Rio Jorco 10 .34.3-37.6 (36.0) Alajuela : La Fortuna 10 31.9-36.0 (34.4) Limon: Pandora 10 33.8-37.6 (35.9) and from the Caribbean lowlands (table 58). The only noticeable differences in proportions between males and females is in the ratio of the tympanum to that of the eye; for ex- ample, the mean ratio in 10 males from the Meseta Central is 0.493 and in eight females, is 0.614. The head is about as wide as the body and slightly longer than wide. The top of the head is flat. In dorsal profile the snout is acutely rounded. In lateral profile the shape of the snout varies geographically and sexu- ally. Specimens from the Caribbean low- lands have blunt snouts; those from the Pa- cific lowlands have longer, more slender snouts that are pointed in lateral view, and those from the Meseta Central are interme- diate in snout shape between the two low- land populations. These differences in the shape of the snout are dependent on the na- ture of the underlying cranial bones, princi- pally the maxillary and nasals. In specimens from the Caribbean lowlands, the nasals are long, wide, and barely separated from the sphenethmoid; the anterior edge is just pos- terior to the nostril. The maxillary flanges are nearly vertical. In specimens from the Pacific lowlands the nasals are relatively short- er, narrower, and rather widely separated from the sphenethmoid; the anterior edges of the nasals do not extend so far forward as the specimens from the Caribbean lowlands. The maxillary phalanges slant medially. In these cranial characters, specimens from the Meseta Central are intermediate between the two lowland populations. Superimposed on this geographic variation are ontogenetic changes, which are most noticeable in males. In smaller, and presumably younger, speci- mens the snouts are more pointed than in larger specimens; consequently, some small males from the Caribbean lowlands resemble larger males from the Pacific lowlands, since the nasals and maxillaries of the former are not fully ossified. In addition, in small breed- ing males the sphenethmoid is only about one- half ossified, a large frontoparietal fontanelle is present, the anterior arm of the squamosal extends only about one-fourth the distance to the maxillary (two-thirds the distance in larger specimens), and the prootics are short, as compared with the long, thin elements in larger specimens. The nostrils are slightly protuberant and are situated at about three- fourths of the distance from the eyes to the tip of the snout. The canthus is slightly angular; the loreal region is noticeably con- cave, and the lips are thin and flaring. A moderately heavy dermal fold extends pos- teriorly from the posterior corner of the eye to a point above the insertion of the arm; the fold obscures the upper edge of the tympan- um, which is otherwise distinct and separated from the eye by a distance equal to about two-thirds of the diameter of the tympanum. The arm is moderately long; the upper arm is slender, and the forearm is somewhat more robust. A distinct axillary membrane is absent. A row of low tubercles forms a scal- loped dermal ridge along the \entrolateral edge of the forearm, and a faint transverse fold is present on the wrist. The fingers are rather short and stout and bear large discs. The width of the disc on the third finger is equal to the diameter of the tympanum. The subarticular tubercles are large and round; the distal tubercle on the fourth finger is flattened and in about one-third of the speci- mens is bifid. The supernumerary tubercles are moderately small, conical, and usually present in a single row on the proximal seg- ments of each digit. No distinct palmar tu- bercle is present, although a cluster of small tubercles sometimes is fused on the palm. The prepollex is noticeably enlarged and in 616 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 breeding males bears an extensive horny nup- tial excrescence. The fingers are about one- half webbed (fig. 28.3B). The webbing is vestigial between the first and second fingers, and extends from the distal end of the penul- timate phalaax of the second to the middle of the antepenultimate phalanx of the third, and from the base of the penultimate pha- lanx of the third to the distal end of the penultimate phalanx of the fourth finger. The hind limbs are moderately short and slender; the adpressed heels overlap by about one- fourth of the length of the shank, and the tibiotarsal articulation extends to a point between the eye and the tip of the snout. The tarsal fold is thin and extends the full length of the tarsus. The inner metatarsal tubercle is long, low, flat and elliptical. The toes are long and relatively slender; the discs are slightly smaller than those on the fingers. The subarticular tubercles are moderately large, round, and subconieal; the supernumerary tubercles are small, conical, and widely dis- persed in a single row on the proximal seg- ments of each toe. The toes are about four- fifths webbed (fig. 2S3D). The web connects the first and second toes at the bases of the discs; the web extends from the base of the disc of the second toe to the middle of the penultimate phalanx of the third, from the base of the disc of the third to the middle of the penultimate phalanx of the fourth and on to the disc of the fifth toe. The anal opening is directed posteroven- trally near the upper level of the thighs and is covered by a short anal sheath. The skin is granular on the belly and the posteroven- tral surfaces of the thighs; the other surfaces are smooth. The tongue is broadly cordiform, usually shallowly notched anteriorly and pos- teriorly, and barely free behind. There are four to six (mean, 5.2) prevomerine teeth on small transverse ridges between the ovoid ehoanae. The vocal slit extends from the mid- lateral base of the tongue to the angles of the jaws. The vocal sac is bilobate and not greatly distensible. The dorsal ground color of Smilisca sor- dicla is gray, pale tan, or reddish brown; the venter is white. The dorsum is variously marked with dark gray, dark brown, reddish brown, or olive-green spots or blotches (pi. 71, figs. 1 and 2). The limbs are banded with dark brown, or olive-green. The flanks are dark brown with cream, greenish gray, or bluish gray mottling. The posterior surfaces of the thighs are dark brown with pale blue, pale green, or tan flecks. The iris varies from creamy silver to grayish yellow or bronze with a variable amount of black reticulations. A dark interorbital bar usually is present. Dorsal markings on the body usually consist of a blotch, or two or more spots, on the occiput, in the scapular region, and in the sacral region. In many specimens, especially females, these markings are in the form of broad transverse bars. A few individuals lack dorsal markings or have scattered dark flecks on the back. Some individuals have scattered small white spots on the dorsum. White la- bial stripes and anal stripes are absent in all specimens. The transverse bars on the limbs are indistinct in some specimens from the Meseta Central and the Caribbean lowlands, whereas the bands are distinct in all speci- mens from the Pacific lowlands. Specimens from the Caribbean lowlands have two to six bars on each shank, whereas specimens from the Pacific slopes ha\e four to six bars on the shank, and specimens from the Meseta Cen- tral has as many as eight bars on each shank. The flanks and the posterior surfaces of the thighs are usually marked by bluish white or creamy tan flecks, respectively, but this coloration varies considerably. In specimens from the Caribbean lowlands a small amount of flecking is present in the inguinal region; on the posterior surfaces of the thighs flecks are few or absent. In specimens from the Meseta Central, numerous large flecks or small round spots (pale bluish white in life) are present on the posterior half of the flanks; small flecks are present on the posterior sur- faces of the thighs. Specimens from the Pa- cific slopes and the lo\\lands of southern Costa Rica (Puntarenas and San Jose prov- inces) have bold mottling of black and bluish white on the flanks and many bluish white flecks on the posterior surfaces of the thighs. In specimens from the Pacific slopes of Guan- acaste in northwestern Costa Rica, flecks arc present in the inguinal region; indistinct flecks are present on the posterior surfaces of the thighs. The throat is immaculate in 1970 DUELLMAN: HYLID FROGS 617 specimens from the Caribbean lowlands in Limon Province; the throats are dusky later- ally in most other specimens except some from the Meseta Central, in which the throats are heavily flecked with black. This variation occurs in males and females. Tadpoles: Eight tadpoles in developmen- tal stage 36 have body lengths of 10.2 to 11.7 (mean, 10.8) mm. and total lengths of 29.5 to 34.5 (mean, 32.3) mm. A typical tadpole in this stage has a body length of 11.7 mm. and a total length of 34.5 mm. The body is about three-fourths as deep as wide; the snout is broadh' rounded in dorsal view, slop- ing and rounded in lateral view. The nostrils are shghtly closer to the eyes than to the tip of the snout. The eyes are widely separated and directed dorsolaterally. The spiracle is sinistral and ventral to the midline; the spirac- ular opening is directed dorsolaterally at a point about two-thirds the length of the body. The mouth is ventral; the cloacal tube is short and dextral. The caudal musculature is hcaxy and straight. The dorsal fin does not extend onto the body and is deepest at about the midlength of the tail. At that point the depth of the dorsal and ventral fin is about equal (fig. 284F). The mouth is large and has well-developed lateral folds. The entire upper and lower lips are bordered by two rows of small papillae; additional papillae are present in the lateral fold. The upper beak is robust; the inner sur- face is curved so as not to form a continuous arch with the slender lateral processes. The lower beak is robust; both beaks bear blunt ser- rations. There are two upper and three lower rows of teeth. The two upper rows are about equal in length, and the second row is nar- rowly interrupted medially. The three lower rows are complete and nearly as long as the upper rows. Usually the lower rows are deeply indented medially (fig. 285F). The body is tan; in some indixiduals there is an olive-tan tinge. The caudal musculature is tan with dull red or reddish brown flecks and dashes, which tend to form a crossbar pattern on the dorsal surface of the caudal musculature. Bluish green flecks are present on the sides of the body in some individuals. Usually the belly is pale tan with a silvery white tint, but in some specimens the belly is silvery golden. The iris is bronze. Mating Call: The call of Smilisca sor- dida consists of one to six moderately short, rather high-pitched notes repeated at inter- vals of 12 seconds to several minutes. The duration of each note is 0.18 to 0.45 (mean, 0.29) seconds. Each note is a vibrant rattle having 78 to 135 (mean, 105) pulses per second. The fundamental frequency is 90 to 140 (mean, 123) cycles per second. Two fre- quency bands are emphasized; these major frequencies are at about 1215 and 2695 cycles per second (pi. 33, fig. 3). Natural History: Smilisca sordida lives in the vicinity of rocky streams having low gradients. Breeding takes place primarily in the dry season, when the water in the streams is clear and at a low level. Through- out most of the range of S. sordida, showers or even short heavy rains, occur in the dry season. After such rains the breeding activity is maximum. Breeding congregations have been found from December to April. Males usually call from rocks or gravel bars in, or at the edge of, streams. Some individuals perch in low bushes overhanging the streams, and some sit in shallow pools in the streams. Clasping pairs have been found on the banks of streams and shallow water in streams. The tadpoles live in shallow parts of the stream, where they cling to the surfaces of small rocks and hide beneath leaves and rocks. Nine recently metamorphosed young have snout- vent lengths of 13.1 and 15.7 (mean, 14.9) mm. Remarks: Duellman and Trueb (1966, p. 328) discussed the systematic status of the various names that have been applied to the frogs here called Smilisca sordida. Most ref- erences to this Costa Rican species are found under the name of Hijla gabhii. Etymology: The specific name sordida is derived from the Latin sordidus meaning "dirty" and apparently refers to the dull, gray- ish brown dorsal color of many preserved specimens of this species. Distribution: Smilisca sordida occurs along the Pacific slopes and lowlands from Guanacaste, Costa Rica, southeastward to ex- treme western Panama. It occurs to eleva- tions of about 1200 meters on the Meseta Gen- 618 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fig. 291. Distribution of Smilisca sordida. tral in Costa Rica and on the Caribbean slopes and lowlands of Costa Rica and prob- ably adjacent Panama (fig. 291). One speci- men reportedly comes from "Rio Grande, Nicaragua." See Appendi.x 1 for the locality records of the 465 examined. Genus Pternohyla Boulenger Pterni)liijla Boulenger, 1882b, p. 326 |type species, Pternohyla fodiens Boulenger, 1882a, by monotypy]. Generotype: Pternohyla fodiens Boulen- ger, 1882b. Etymology: The generic name is derived from the Greek pterna, meaning heel, and Hijlas, a character in Greek mythology. The generic name is in reference to the spade-like inner metatarsal tubercle. Definition: Frogs of the genus Pterno- hyla are medium in size and have a pale brown dorsum with dark brown dorsal mark- ings. The pupil is horizontal, and the palpe- bral membrane is not reticulated. The limbs are short; the fingers lack webbing, and the toes are less than half webbed. The terminal discs are small, and the inner metatarsal tu- bercle is large. The \ocal sac is subgular and paired. Breeding males ha\'e horny nuptial excrescences on the thumbs. The skin of the head is partly co-ossified with the underlying cranial elements. Tlie canthal ridges are pronounced, and the maxillaries are expanded laterally to form a labial shelf. The skull is as wide as, or slightly wider than, long. A prenasal bone is absent, but an internasal is present in one species. The squamosal-maxil- lary arch is complete, and quadratojugals are present. The palatines are robust and articu- 1970 DUELLMAN: HYLID FROGS 619 late with the sphenethmoid. The medial ra- mus of the pterygoid is reduced and does not articulate with the prootie. Bifid, spatulate teeth are present on the premaxillaries, inaxil- laries, and prevomers; the palatines and para- sphenoid are edentate. The tadpoles are short- tailed pelagic types with an anteroventral mouth ha\ing robust beaks and large papillae laterally and \entrally. The long, pointed teeth are arranged in two upper and three lower rows. The mating call consists of a series of short notes resembling the quacking of a duck. The haploid number of chromo- somes is 12 (known only in fodiens). Composition of Genus: Two monotypic species are recognized; both occur in western Mexico. I have examined 6.30 preserved frogs, 13 skeletons, and three lots of tadpoles of Pternohyla from Mexico and one frog from Arizona. Analysis of Characters: The two specie's differ from one another in several external characters. Pternohyla fodiens has propor- tionately longer legs and feet and a propor- tionately large head (see ratios given in the accounts of the species). The head is about as wide as long in dentate and wider than long in fodiens. The latter has a proportion- ately smaller tympanum than dentata. The fingers and toes of dentata are robust and lack expanded discs. The inner metatarsal tubercle is large, elliptical, and rounded in section in dentata, whereas the tubercle is larger, ovoid, flattened in section, and has an elevated outer edge in fodiens (fig. 292). Both species have dark vocal sacs. Those of fodiens are dark gray or black with the tips of granules white, thereby gi\'ing a white- speckled appearance. The sacs of dentata are grayish brown. In breeding males the two hah'es of the vocal sac are closely approxi- mated in fodiens and broadly separated by granular skin in dentata (fig. 293). Integumentary-cranial co-ossification is in- complete. The nasal at the anterior edge of the orbit, the sphenethmoid, and the dorsal part of the prootie are not co-ossified in either species. The dermal roofing bones are more extensi\'ely ossified in fodiens than in dentata. In the former the more greatly expanded la- bial flanges result in a proportionately broad- er skull, and the larger nasals and fronto- parietals result in less of the sphenethmoid being exposed dorsally than in dentata. The premaxillaries are more robust and are in- \o]\'ed in eo-ossification in fodiens, and the snout region is further modified by the pres- ence of an internasal (Trueb, 1970a), a der- mal bone medial to the external nares and the anterior tips of the nasals. The dorsal surface of the internasal is involved in integumentary- cranial co-ossification (fig. 294). Distribution: Frogs of the genus Pterno- hyla occur in xeric environments from south- central Arizona in the United States south- ward through western Mexico to the Tepal- catepec Valley in Michoacan. Discussion: The adaptive trends in Pter- nohyla have been towards a fossorial exis- tence, as illustrated by the modifications in the limbs and head and by the squat, toad-like form of the body. Although both species have these modifications, certain specializa- tions have been carried farther in one species than in the other. For example, the limbs are proportionately shorter and the terminal discs on the digits are further reduced in dentata than in fodiens. In the latter the integumen- tary-cranial co-ossification of the skull is more nearly complete, and the inner metatarsal tubercle is more specialized than in dentata. The bony internasal ridge in fodiens seems to be analagous to the boss on the snout in some species of Biifo and Scaphioptis. Like- wise, the inner metatarsal tubercle in fodiens is spatulate like that in Scaphioptis and some Biifo, although in fodiens the tubercle is not horny. Trueb ( 1970a ) showed the cranial char- acters of Pternohyla could be derived from those of Sniilisca baudinii and that osteologi- cally P. dentata is somewhat intermediate be- tween S. baudinii and P. fodiens. Smilisca baudinii has paired, subgular vocal sacs and an enlarged inner metatarsal tubercle. The tadpoles and mating call of Pternohyla are not greatly different from those of Smilisca bau- dinii. Furthermore, Starrett (1960b) sug- gested a close relationship of the two genera on the basis of the identical jaw musculature. Thus, it seems likely that Pternohyla e\'olved from a Smilisca baudinii stock and that the evolutionary trends were towards adaptation for a fossorial existence in xeric environments. 620 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fig. 292. Hands and feet of Ptemohyla. A and C. P. dcniata. K.U. No. 60081. B and D. P. fodiens, K.U. No. 78463. X 6. 1970 DUELLMAN: HYLID FROGS 621 Fig. 293. Ventral views of throats of breeding males of Ptemohyla. A. P. dcntata, K.U. No. 60083. B. P. fodietis, K.U. No. 78463. X 2. Duellman and Trueb (1966) hypothesized that the species of Smilisca probably had differentiated from one another by the end of the Pliocene, at which time S. haudinii inhabited the Pacific lowlands of Mexico. In- creasing aridity throughout the Pleistocene probably was the environmental impetus that resulted in the differentiation of a fossorial stock which gave rise to Ptemohyla. Appar- ently P. dentata represents a population of the Ptemohyla stock that was formerly iso- lated in the upper Rio Santiago Basin on the Mexican Plateau. Ptemohyla dentata Smith Ptemohyla dentata Smith, 19.57, p. 1 [holotype, U.I.M.N.H. No. 40551 from 8 miles northeast of Lagos de Moreno, Jalisco, Mexico; W. H. Davis, W. Z. Lidicker, and John R. Winkelmann collectors]. Diagnosis: This moderate-sized, casque- headed frog is characterized by incomplete integumentary-cranial co-ossification and the absence of an internasal. It is readily distin- guished from Ptemohyla fodiens by having a rounded inner metatarsal tubercle and narrow- tips of the digits, whereas fodicm has a spade- like inner metatarsal tubercle and distinct terminal discs on the digits. Furthermore, fodiens has the two halves of the vocal sac connected medially and an internasal ridge resulting in an acutely rounded snout; den- tata has the two halves of the vocal sac broad- ly separated medially and lacks an internasal ridge, thereby having a bluntly rounded snout. Of the other Middle American casque- headed hylids, Anotheca lacks labial flanges and has long cranial spines, and Triprion has a broad labial flange, a large prenasal bone, and large terminal discs on the digits. Description: In a series of 25 males from Aguascalientes, Mexico, the snout-vent length is 47.6 to 62.1 (mean, 52.4) mm.; the ratio of tibia length to snout-\ent length is 0.311 to 0.360 (mean, 0..344); the ratio of foot length to snout-vent length is 0.320 to 0.370 (mean, 0.351); the ratio of head length to snout-vent length is 0.275 to 0.313 (mean, 0.294); the ratio of head width to snout-vent length is 0.268 to 0.310 (mean, 0.291), and the ratio of the diameter of the tympanum to that of the eye is 0.604 to 0.767 (mean, 0.660). Three females from the same locality have snout-vent lengths of 52.7 to 54.0 (mean, 53.4) mm. They do not differ significantly from the males in any proportions. The head is about as wide as long and noticeably narrower than the body. In dorsal profile the snout is bluntly rounded; in lateral profile it is acutely rounded and protruding beyond the leading edge of the lower jaw. The snout is moderately long and somewhat spatulate. The nostrils are protuberant, di- rected dorsally, and situated at a point about three-fourths of the distance from the eyes to the tip of the snout. The canthal ridge is elevated and terminates just posterior to the nostrils. The loreal region is deeply concave, and the lips are broad and flared. The entire labial region is moderately expanded; the ex- panded lips extend posteriorly to the tympan- um. A bony ridge extends posteriorly from the orbit, abo\'e the tympanum, and continues as a dermal fold to a point above the inser- tion of the arm. The skin on the skull is co- ossified with the underlying dermal bones, 622 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fig. 294. Dorsal views of the skulls of Pternohi/la. A. P. dentata, K.U. No. 106291. B. P. jodiens, K.U. No. 86615. X 5. 1970 DUELLMAN: HYLID FROGS 623 except in the region of the frontoparietal fon- tanelle, the sphenethmoid, the nasals immedi- ately anterior to the orbit, and the outer edges of the maxillaries. There is no distinct inter- nasal ridge extending to the tip of the snout. The upper edge of the tympanum is concealed by the bony supratympanic ridge, and the posterior edge is concealed by granular skin in some specimens; otherwise the tympanum is distinct and separated from the eye by a distance equal to about one-half of the diam- eter of the tympanum. The arms are short and robust. There are no tubercles along the ventrolateral edge of the forearm, but a thin transverse dermal fold is present on the wrist. The fingers are short, robust, and lack terminal discs. The tips of the fingers are bluntly rounded. The subarticular tubercles are large and round; none is bifid. Faint supernumerary tubercles are present on the proximal segments of the third and fourth fingers in some specimens. A broad, diffuse palmar tubercle is present; the prepollex is moderately enlarged and, in males, bears a thin nuptial excrescence. The thumb is nearly as long as the second finger, and webbing between the fingers is absent (fig. 292A). The legs are short. The heels of the adpressed limbs overlap by about one- fourth of the length of the shank; the tibio- tarsal articulation extends to the axilla. A distinct transverse dermal fold is present on the heel, and a distinct, elevated tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is elongate, ellipti- cal, and round in section. The outer meta- tarsal tubercle is small and subconical. The toes are moderately short, slender, and lack terminal discs. The subarticular tubercles are moderately large and round; small, usually indistinct supernumerary tubercles are pres- ent on the proximal segment of each digit. The toes are webbed only basally ( fig. 292C ) . The anal opening is directed posteriorly at the level of the upper edge of the thighs; no anal flap is present. The skin on the dorsal surfaces of the body is weakly granular, and that on the dorsal surfaces of the limbs and the ventral surfaces of the forelimbs, shanks, and feet is smooth. The skin on the belly and ventral surfaces of the thighs is hea\'ily granular. The tongue is broadly cordiform. shallowly notched behind, and free posterior- ly for about one-third of its length. The dentigcrous processes of the prevomcrs are small transverse elevations between the small round choanae. There are four to six teeth on each process, and the total number of prevomerine teeth is eight to 12 (mean, 10.3). The vocal slits extend from the posterolat- eral base of the tongue nearly to the angles of the jaws. The vocal sac is paired and sub- gular; the halves of the sac are connected by a narrow tube, but there is a broad separa- tion of granular skin between the two halves of the sac. The color in life is unknown. In preserva- tive the dorsal surfaces of the body and limbs are grayish brown to pale reddish brown with dark brown to reddish brown spots and longitudinal markings (pi. 2, fig. 1). In those individuals having reddish brown blotches, the blotches are narrowly outlined by dark brown or black. Most individuals have dark spots on the upper eyelids and a dark dash on the head anterior to the eyes. Dark bars are present on the upper lips. The dorsal markings are either discrete spots ir- regularly arranged in about four longitudinal rows or consist of fused spots which form broad longitudinal stripes. The most common pattern of fused spots consists of a pair of paravertebral stripes and a row of dorsolat- eral spots. The flanks are creamy tan with dark brown spots. Brown blotches or trans- verse bars are present on the limbs. There are two or three such bars on each shank and thigh, and usually two on the foot and fore- arm. The posterior surfaces of the thighs are creamy white with brown flecks and dashes. The venter is creamy yellow, and the vocal sacs are browaiish gray. Tadpoles: The tadpoles of Pternohyla dentata are unknown. Mating Call: Recordings of the call of this species are not available; consequently it can not be described and compared with that of P. fodiens. Natural History: Little is known about the natural history of this species. According to Smith (1957, p. 4) the holotype was found in a temporary road-side pond in high plateau country characterized by short-grass plain with scattered xeric shrubs. Chrapliwy, Wil- 624 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Hams, and Smith (1961, p. 87) reported on the large series from Aguascalientes and stated: "All were taken on the night of July 21 from a flooded field with rain water level varying from one to four inches in depth. Rain had fallen intermittently in the day and evening. The frogs, in chorus, were not wary and often continued to call after being picked up. Several pairs were obser\ed in amplexus." Remarks: Smith (1957) in his description of Pternohyla clentata diagnosed this species as having no bony labial fringe and by pos- sessing parasphenoid teeth. Smith apparently referred to the outer edge of the lips as the "labial fringe"; the outer edge of the lips are not involved in integumentary-cranial co-ossi- fication, but the dorsal surfaces of the ex- panded maxillaries are involved in co-ossifica- tion. Despite the statement of Smith, para- sphenoid "teeth" are absent in this species. Furthermore, Smith (1957, p. 3) stated: "The species P. dentata possesses both parasphe- noid and palatal 'teeth,' like Diaglena and Triprion whereas P. fodiens lacks them." Odontoids are present on the palatines in Triprion spatidottis and on the parasphenoid in both species of Triprion. The palatines and parasphenoid are edentate in both species of Pternohyla. Etymology: The specific name is Latin meaning tooth and refers to the supposed presence of parasphenoid teeth in this species. Distribution: Pternohyla dentata is known from the upper Rio Santiago Basin in southern Aguascalientes and northern Ja- lisco, Mexico, at elevations of 1800 to 1900 meters (fig. 295). See Appendix 1 for the locality records of the 148 specimens examined. Pternohyla fodiens Boulenger Pternohyla fodiens Boulenger, 1882b, p. 326 [holo- type, B.M.N.H. No. 1947.2.24.26 from Presidio, Sina- loa, Mexico; Alphonso Forrer collector]. Giinther, 1901 (188.5-1902), p. 292. Kellogg, 1932, p. 135 [.synon\ niizfd Hyla rudi.T-" \ 1 ,^^^^- ■ I , >>? 627 Fig. 296. Tadpole ot Ptcrnolnjla fodiens, K.U. No. 104193. X 4. Tadpoles: A typical tadpole in develop- mental stage 26 has a total length of 29.2 mm. and a body length of 12.3 mm. The body is as wide as deep. In dorsal profile the snout is bluntly rounded, and in lateral profile, more acutely rounded. The nostrils arc directed an- tcrolaterally and situated about midway be- tween the eyes and the tip of the snout. The eyes are moderately small, situated dorsolat- erally and directed laterally. The spiracular opening is on the level of the midline about two-thirds of the distance from the snout to the posterior end of the body. The anal tube is dextral and moderately long. The caudal musculature is slender and terminates just short of the tip of the caudal fins. The dorsal fin does not extend onto the body; at mid- length of the tail the depth of the caudal musculature is slightly less than the depth of either the dorsal or ventral fin (fig. 296). In life both the body and tail are dull tan \\'ith olive-brown mottling. The belly is dusty white. In preser\'ative the body is dark brown and the caudal musculature is creamy tan with dark brown flecks, which are also pres- ent on the caudal fin. Fig. 297. Mouth of tadpole of Ptenwhyla fodiens, K.U. No. 104196. X 25. The mouth is small, anteroventral in posi- tion and direction. The median part of the upper lip is bare; large, partially fused labial papillae are present in a single row midven- tralh' and anterolaterally, and in two rows laterally. The lips are folded laterally. The beaks are robust and bear small, pointed ser- rations. There are two upper and three lower rows of teeth. All of the teeth are moderately long and pointed. The second upper row is broadly interrupted medially. The lower rows are complete; the first and second lower rows arc nearly as long as the first upper row, whereas the third lower row is noticeably shorter (fig. 297). Webb ( 1963 ) described small tadpoles and later developmental stages from Sinaloa; he mentioned metamorphosing incli\iduals having lengths of IS to 24 mm. (including tail stubs). Mating Call: The call of Ptemohijla fo- diens consists of a series of low-pitched notes, resembling the quacking of a duck. The notes are cjuickly repeated; the note repetition rate is SI to 115 (mean, 95) notes per minute. The notes have a duration of 0.21 to 0.2S (mean, 0.25) of a second and a pulse rate of 118 to 125 (mean, 122) pulses per second. The fundamental frequency is 122 to 134 (mean, 126) cycles per second and the dominant frequency is 2200 to 2278 (mean, 2230) cycles per second (pi. 34, fig. 1). N.ATLiRAL History: Ptemohijla fodiens in- habits arid tropical scrub forests where it breeds in temporary pools formed by rains which fall in the months of June through September. Males usually call near temporary ponds, but not at the edge of the water. The frogs call from secluded places such as under the edge of a rock, at the bases of bushes, or in clumps of grass. 628 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Hardy and McDiarmid (1969) reported on the tadpoles of this species from La Cruz, Sinaloa. They stated: "Thousands of tadpoles, all of which appeared to be newly hatched and about 10 mm. in length, were all that remained from the previous night's breeding activity. Most of the larvae were floating in clusters with their tail pointing downward from the surface of the pond. When a cluster was disturbed, the larvae would disperse, some swimming away and others sinking to the bottom. Jelly envelopes, some containing undeveloped eggs were scattered over the bot- tom of the pond. The tadpoles clustered at the surface may have been feeding on surface scum. Three large series were collected and allowed to develop. Three days later the larvae lost their external gills." I obtained larvae of the species in a shal- low, grassy pond, 34 kilometers north-north- west of Tepic, Nayarit. Remarks: Firschein (1951) noted the phragmotic behavior and "Unken refle.x" in this species. I have observed the same be- havior in this frog. Individuals when dis- turbed, flex the head downward and elevate the limbs so as to rest on the belly. The fossorial habits of Ptemohijla indicate that the phragmotic behavior is not for the purpose of closing holes in trees as it is for Triprion, but more likely the head is used for closing burrows in the ground. Pternohyla fodiens was only recently dis- covered in the United States ( Chrapliwv and Williams, 1957). Kellogg ( 1932 ) showed that the type of Hyla rudis Mocquard (1899a) is a young individual of Pternohyla fodiens. As noted previously, the coloration of the juveniles is noticeably different from that of the adults. Furthermore, juveniles lack integumentary- cranial co-ossification. Etymology: The specific name fodiens is the genitive of the Latin fodio, meaning to dig or to dig up and apparently refers to the supposed digging adaptations of the spade- like inner metatarsal tubercles. Distribution: Pternohyla fodiens inhab- its xeric regions from south-central Arizona in the United States southward through west- ern Sonora and the coastal regions of Sinaloa, and thence into the foothills of the Pacific slopes of the Sierra Madre Occidental in Na- yarit and southward onto the Mexican Plateau in Jalisco. This species also occurs on the Colima Plateau and in the Tepalcatepec Val- ley in Michoacan, Mexico (fig. 295). This species occurs at elevations from sea level to about 1500 meters. See Appendix 1 for the locality records of the 498 specimens examined. Genus Triprion Cope PhanjnRodon Cope, 1865b, p. 193 [type species Phanjngodon petasatus Cope, 1865b, by monotypy; preoccupied by Phanjngodon Die.sing, 1861 ( Nematli- elminthes)]. Triprion Cope, 1866a, p. 127 [replacement name for Phanjngodon Cope, 1865b, preoccupied]. Diaglena Cope, 1887, p. 12 [type species, Triprion spatulatus Giinther, 1882, by monotypy]. Generotype: Pharyngodon {=:Triprion) petasatus Cope, 1865b. Etymology: The generic name is derived from the Greek trion, meaning three and the Greek prion, meaning saw, and is in reference to the serrate labial fringes anteriorly and laterally. Definition: The frogs in this genus are moderately large to large and are character- ized by integumentary-cranial co-ossification and a casqued head that is longer than broad (fig. 298). The dorsum is olive-green to yellowish tan and uniformly colored or marked with blotches or reticulations. The pupil is horizontally elliptical, and the palpe- bral membrane is clear. The fingers are webbed basally, and the toes are about two- thirds webbed. Moderately large terminal discs are present on the digits, and a large. Fig. 298. Lateral \ie\v of the head of Triprion petasatus, K.U. No. 71503, showing casque head. X 3. 1970 DUELLMAN: HYLID FROGS 629 elliptical inner metatarsal tubercle is present. The vocal sac is single, median or paired, and subgular or bilobate and situated posteriorly on the throat. The tongue is round. Breeding males have horny nuptial excrescences on the thumbs. The skin is completely co-ossified with the underlying cranial elements. A large prcnasal and the laterally expanded ma.xil- laries form a broad, serrate, labial shelf (fig. 299). The premaxillaries are partly hidden by the prenasal, and the alary processes of the premaxillaries are rotated anteriorly. The skull is completely roofed; a dermal sphen- ethmoid is present or absent. The squamosals are in bony contact with the maxillaries, and the quadratojugals are well developed. The palatine is slender, and the medial ramus of the pterygoid is reduced and attached to the prootic only by connective tissue. Teeth are present on the premaxillaries, maxillaries, and pre\'omers, and odontoids are present on the parasphenoid and present or absent on the palatines. The teeth are spatulate and bifid. The tadpoles are pelagic types with antero- ventral mouths and deep caudal fins. The mating call consists of a single, low-pitched note. The chromosome number is n = 12, 2n = 24 (known only in petasatus). Composition of the Genus: Two species [peiasatus and spatidatus), the latter with two subspecies, comprise the genus; both are Middle American endemics. Of the two spe- cies, 791 preserved frogs, 28 skeletons, nine lots of tadpoles, and four preserved clutches of eggs have been examined. Analysis of Characters: The principal specific characters of the frogs in the genus Thprion are those of the casque head. A der- mal sphenethmoid is present in petasatus and absent in spatulatus; in the former the canthal ridges are nearly perpendicular to the body axis, whereas in spatulatus the ridges are inclined anteromedially. The snout is up- turned in petasatus and nearly straight in spatulatus, and the latter has slender palatines that bear odontoids, whereas in petasatus the palatines are greatly reduced. The vocal sac is single and median in spatulatus and paired in petasatus. In both species the vocal sac is situated on the posterior part of the throat. The structure of the hands and feet in the two species is nearly identical (fig. 300). Distribution: The species of Triprion in- habit xeric areas on the Pacific lowlands of Mexico from central Sinaloa to the Isthmus of Tehuantepec and in the Yucatan Peninsula southward to central El Peten, Guatemala. Discussion: The phylogenetic relation- ships of the casque-headed hylids were dis- cussed by Trueb (1970a), who provided evi- dence that Diaglena and Triprion were con- generic. Furthermore, she showed that the South American casque-headed hylid genera, Aparasphenodon, Corythomantis, Osteocepha- lus, Trachycephalus, and Tctraprion were not related to Triprion. Likewise, the Mexican genus Pternohyla, although probably phylo- genetically closer to Triprion than are the South American genera, represents a phyletic line that is less advanced in adaptive cranial modifications. Triprion evidently was more widespread in Mexico and northern Central America prior to the Pleistocene. The present distribution of the species is a relictual pattern that is com- mon among xeric restricted species and is the result of isolation due to changing environ- mental conditions in the Pleistocene ( Duell- man, 1960b and 1966c). Triprion petasatus is more highly specialized than spatulatus; this specialization is evident in the more high- ly modified skull — presence of a dermal sphenethmoid, reduction of palatines, and higher canthal ridges. Triprion spatulatus Giinther Triprion spatulatus Giinther, 1882, p. 279 [syn- types B.M.N.H. Nos. 1947.2.25.79-1947.2.25.81 from Presidio de Mazatlan, Sinaloa, Mexico; Alphonso Forrer collector]. Diagnosis: This is a large species (males to 87 mm.; females to 101 mm. ) that is readily distinguished from other Middle American casque-headed hylids by having a large pre- nasal, greatly expanded maxillaries, odontoids on the palatines, no spines on top of the head and no dermal sphenethmoid. Triprion peta- satus differs by having a dermal spheneth- moid, the tip of the snout upturned and by lacking odontoids on the palatines. Pterno- hyla has only moderate labial flanges, and lacks a dermal sphenethmoid and prenasal. Furthermore, Pternohyla is squat and toad- like in appearance and has a spade-like inner 630 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fig. 299. Skulls of Triprion. A. Dorsal and B. Ventral of T. spatnlahis. K.U. No. 84904; C. Dor- sal and D. Ventral of T. petasatm, K.U. No. 71780. x 3. 1970 DUELLMAN: HYLID FROGS 631 Fig. 300. Hands and feet of Triprion. A and B. T. spatulatus, U.M.M.Z. No. 115322. C and D. T, petasatvs, K.U. No. 71503. x 3.5. 632 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 metatarsal tubercle. Anotheca lacks labial flanges and has spines on the supratympanic and occipital ridges. Content; Two subspecies arc recognized: T. spatulatus spatulatus Giinther and T. spat- ulatus reticulatus Taylor. Remarks: Although minor differences in proportions exist, the subspecies are most readily distinguished on coloration. The amount of dark pigmentation is greater in the southern subspecies reticulatus than in the northern spatulatus (Duellman, 1968c, p. 198). Specimens from Sinaloa are more nearly uniformly colored than are those from Colima and southward. Thirty-two per cent of the specimens from Sinaloa lack dorsal markings, and 45 per cent have small dark flecks on the dorsum, whereas the others have dark dashes or fine reticulations. No specimens from Co- lima and southward lack dorsal markings; eight per cent of the specimens from Colima have flecks or dashes on the dorsum. The other specimens from Colima and all of those from Michoacan, Guerrero, and Oaxaca have dark reticulations or spots on the dorsum ( fig. 301 ) . The color patterns of T. spatulatus were assigned values and coded numerically; 0 — no dorsal markings 1 — small flecks 2 — dashes 3 — fine reticulations 4 — bold reticulations 5 — reticulations and spots All individuals in each of fixe geographic samples were coded; ranges and means for each sample were calculated (table 59). The sample from Sinaloa (mean color value, 0.96) is distinctly different from the others, which have much higher color values. Specimens from Colima and southward have a mean color value of 3.67. Taylor (1942) described Diaglena reticu- lata on the basis of one specimen and com- pared his type with the only two specimens of Triprion spatulatus in the United Stutes at that time. In addition to the obvious differ- ences in coloration, he noted that reticulatus had a proportionately shorter, broader head, with the canthal ridges uniting farther for- ward than in spatulatus and that the skin was granular on the dorsum, as opposed to smooth in spatulatus. The differences in cranial struc- ture apparently are correlated with age and the amount of ossification. Apparent granu- lation of the skin on the dorsum is due princi- pally to different modes of preservation. Distribution: Triprion spatulatus occurs on the Pacific coastal lowlands in central Sinaloa and from Colima to the Isthmus of Tehuantepec, Mexico, and in the Balsas Basin to elevations of about 350 meters (fig. 302). Triprion spatulatus spatulatus Giinther Triprion spatulatun Guiither, 18S2, p. 279 [syn- types, B.M.N.H. Nos. 1947.2.25.79-1947.2.25.81 from Presidio de Mazatlan, Sinaloa, Me.xico; Alphonso Forrer collector]. Gunther, 1901 ( 1885-1902), p. 293. Fig. 301. Diagrammatic representation of dorsal color patterns in Triprion spatulatus. A. Value 1, K.U. No. 75275. B. Value 2, U.M.M.Z. No. 115322. C. Value 3, U.M.M.Z. No. 104418. D. Value 4, U.M.M.Z. No. 115321. E. Value 5, K.U. No. 86904. The values are tliose assigned for coding purposes (table 59); the plain pattern (Value 0) is not .shown. 1970 DUELLMAN: HYLID FROGS 633 TABLE 59 Geographic Variation in Size and Color Pattern in Triprion spaitdaiiis. (Sample Size in First Column for Measurements, in Fourth Column for Color Pattern) Locality Sinaloa Colima Michoacan Guerrero — Oaxaca N Snout-vent Length Males Females 37 i, 6 9 69.1-85.9 79.6-101,0 35 ^ S- /''' y ! i 0 7" s spafulatus 1 ^ 'v^-i S 1 ^ \ ' • T s reticulatus Y" V^.J-V'7 '\ ,/, '-''' ~'" '-.._.- -XT, I 22° ^<'~/; ,--' '' O ^^\ y > ^ \ 22° 18" ^"^^X^ X" 1 ' - 18° 0 100 300 ^\--__^ ,;■ • / KILOMETERS %*^a,^^^ 106° 100° °l^ ° Fig. 302. Distribution of the subspecies of Triprion spatulatus. 6.34 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Diaglena spatulaia: Cope, 1887, p. 12 [designation of Triprion spatulatiis Giinther, 1882, as the type species of Diatllena Cope, 1887]. Kellogg, 1932, p. 137. Taylor, 1942b, p. .58. Smith and Taylor, 1948, p. 69. Diaglena spatulaia spatulaia: Duellman, 1968c, p. 200. Triprion spatulatus spatulatus: Trueb, 1970a, p. 602 Isynonyniized Diaglena Cope, 1887, with Triprion Cope, 1866a]. Diagnosis: This subspecies is distin- guished from T. s. reticiilahis by having a uni- formly yellowish tan to dull olive-green dor- sum or by ha\ing small dark flecks or dashes dorsally. The other subspecies has bold retic- ulations and/ or spots on the dorsum. Description : In a series of 37 males from the vicinity of Villa Union, Sinaloa, Mexico, the snout-vent length is 69.1 to 85.9 (mean, 75.0) mm.; the ratio of tibia length to snout- vent length is 0.324 to 0.392 (mean, 0.365); the ratio of foot length to snout-vent length is 0.272 to 0.456 (mean, 0.320); the ratio of head length to snout-vent length is 0.304 to 0.386 (mean, 0.353), and the ratio of head width to snout-vent length is 0.171 to 0.261 (mean, 0.218). Six females from the same lo- cality have snout-vent lengths of 79.6 to 101.0 (mean, 86.1) mm. and do not differ significant- ly from the males in proportions. The head is moderately small and modi- fied in the form of a bony casque with the skin completely co-ossified with the skull. The maxillaries and the prenasal are greatly ex- panded and form a broad labial shelf. The snout protrudes far beyond the leading edge of the lower jaw, and the tip of the snout is pointed and not upturned. The edge of the labial shelf is finely serrate. The nostrils are directed laterally at a point about three-fifths of the distance from the eyes to the tip of the snout. Bony supraorbital and preorbital ridges are present. At their juncture in large females, a bony preorbital knob is developed. A sharp canthal ridge extends anteromedially from the preorbital knob and fuses with its counterpart just posterior to the nasal; from this point a distinct nasal ridge extends an- teriorly to the tip of the snout. The loreal region is deeply concave. The labial flange is upturned just anterior to the preorbital ridge; posterior to this point the labial shelf is reduced to a narrow ridge. The eyes are moderately large, protuberant, and directed anterolatcrally. A bony postorbital ridge ex- tends from the orbit to the posterior edge of the skull; the ridge overhangs the upper edge of the tympanum. The posterior edge of the skull is delimited by a low, smooth, transverse bony ridge, which is continuous in all speci- mens. The bony labial ridge posteriorly ob- literates the lower edge of the tympanum in some specimens; likewise, the anterior, down- curved part of the postorbital ridge conceals the anterior edge of the tympanum in some incli\iduals. Consequently, measurements of the tympani arc difficult or impossible. The diameter of the tympanum is equal to about half that of the eye. The upper arms are slender, and the fore- arms are robust. An axillary membrane and tubercles on the ventrolateral edge of the forearm are absent, but a distinct transverse dermal fold is present on the wrist. The fin- gers are moderately long and robust and bear large discs; the diameter of the disc on the third finger is equal to the diameter of the tympanum. The subarticular tubercles are moderately large and subconical; none is bi- fid. The supernumerary tubercles are low, round, and indistinct. A large, flat, elliptical palmar tubercle is present. The prepollex is moderately enlarged and in breeding males is covered with a horny nuptial excrescence \\hich in most incUviduals extends along the inner edge of the thumb to the base of the disc. Webbing is lacking between the first and second fingers and is rudimentary be- tween the others (fig. 300A). The legs are short; the heels of the adpressed limbs over- lap by about one-sixth of the length of the shank. The tibiotarsal articulation extends to the point of the insertion of the arm. A heavy tarsal fold extends the full length of the tar- sus. The inner metatarsal tubercle is mod- erately large, flat, and elliptical. The outer metatarsal tubercle is low, round, indistinct, or absent. The toes are moderately long and bear discs that are slightly smaller than those on the fingers. The subarticular tubercles are moderately small and subconical; the super- numerar\' tubercles are small, low, and incon- spicuous. The toes are about two-thirds webbed (fig. 300B). The webbing extends from the base of the penultimate phalanx 1970 DUELLMAN: HYLID FROGS 635 of the first toe to the distal end of the ante- penultimate phalanx of the second, from the middle of the penultimate phalanx of the second to the middle of the antepenultimate phalanx of the third, from the base of the penultimate phalanx of the third to the base of the antepenultimate phalanx of the fourth and on to the middle of the penultimate pha- lanx of the fifth toe. The anal opening is directed posteriorly at the level of the upper surfaces of the thighs. No anal flap is present, but the area below the anus is covered by moderately large tubercles. The skin is smooth or finely granular on the dorsal surfaces of the body, and somewhat more strongly granular on the flanks and belly. The throat and \'entral surfaces of the thighs are weakly granular and the skin on the rest of the venter is smooth. The tongue is ovoid, usually wider anteriorly than posteriorly, shallowly notched behind, and barely free posteriorly. The den- tigerous processes of the prevomers are small, transverse, narrowly separated, and situated just posteriorly to the posterior margin of the moderately small ovoid choanae. There are four to nine teeth on each process and a total of ten to 16 (mean, 12.0) prevomerine teeth. The vocal slits extend from the posterolateral edge of the tongue to the angles of the jaws. The vocal sac is single, median, subgular, and situated posteriorly on the throat. The general coloration of Triprion spatti- lattis spatiilatus is pale green or yellowish tan with green to yellow flecks (pi. 72, fig. 3). The dorsum varies from a pale grayish green to a dull oli\'e-green or yellowish tan. The head is always somewhat darker than the body. The flanks have a yellowish cast even in those individuals which otherwise are olive- green. The venter is white, except for gray- ish brown flecks on the vocal sac in breeding males. Dark flecks or dashes tend to form indistinct transverse markings on the dorsal surfaces of the thighs and shanks. The iris is dull bronze with black flecks. The dorsal coloration varies from an ab- sence of dark markings on the body to a pattern of dark brown or black reticulations. The color pattern was noted in 105 speci- mens; of these 34 lacked markings, 47 had dark flecks on the dorsum, 17 had dark dashes, six had fine reticulations, and one had bold reticulations (see fig. 301 for a diagrammatic representation of these patterns). In all in- dividuals, the head is more heavily marked than the body; usually the markings on the head consist of short dashes or reticulations. In preservative the dorsum is grayish Ijrown to creamy tan with or without dull brown markings. The flanks are somewhat lighter. The venter is creamy white, and the \ocal sac in breeding males is flecked with grayish brown. T.\DPOLES: No tadpoles of this subspecies ha\e been collected. Mating Call: The call of Triprion spaiii- laiiis spatidatiis consists of a single, moderate- ly long, low-pitched note, "braaa." Record- ings of a chorus of these frogs contain so much background noise that useful audiospectro- grams were impossible to obtain. Natural History: This subspecies inhab- its the xeric, thorn-scrub forest on the coastal lowlands of Sinaloa, where it breeds in tem- porary ponds that are formed in the rainy season, which usually extends from June until early November. On August 14, 1956, I en- countered a breeding chorus at a small, tem- porary pond 31 kilometers north-northwest of Mazatlan, Sinaloa. The frogs were found im- mediately after a torrential rain; males were calling from bare earth banks at the edge of the pond. Hardy and McDiarmid (1969) reported finding several hundred individuals in a pond at La Cruz, Sinaloa on August 20. They stated that the males were calling in full force and stationed about 35 cm. abo\e the water or on rocks in the water. They observed several amplexing pairs swimming in the wa- ter. These authors suggested that although the exact ecological stimulus for breeding is unknown, the combination of sufficient rain, cool weather, and overcast sky may initiate reproductive acti\'ity. Remarks: Duellman and Klaas (1964) reported observations of phragmotic beha\ior of this frog in the laboratory. Indi\iduals were observed to back into holes in a log. The hole was too large to be plugged with the head; instead the frogs pressed themselves tightly against the inner wall of the cavity below the hole and slightly flexed the head 636 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 so that the labial shelf was flush against the wood. Etymology: The specific name is derived from the Latin spatula, meaning spoon, and is in reference to the broad labial flanges, which gives the head a spoon shape. DisTRiBUTiox: Triprion spatitlatus spatu- latus inhabits the Pacific coastal lowlands of southern Sinaloa, Mexico (fig. .302). See Appendi.v 1 for the locality records of the 127 specimens examined. Triprion spatulatus reticulatus (Taylor) Diaglena reticulata Taylor, 1942b, p. 60 [holotvpe, U.S.N. M. No. 115500 from Cerro Arena], Oaxaca, Mexico; Thomas MacDougall collector]. Smith and Taylor, 1948, p. 69. Diaglena spatulata reticulata: Diiellman, 196Sc, p. 200. Triprion spatulatus reticulatus: Trueb, 1970a, p. 602 [synonymized Diaglena Cope, 1887, with Triprion Cope, 1866a]. Diagnosis: This subspecies is distin- guished from the nominate subspecies by hav- ing bold dark brown or black reticulations, and in some indi\'iduals spots also, on a yellowish tan to olive-green dorsum. The nominate subspecies lacks dark dorsal mark- ings or has only small dark flecks or dashes. Description: Males of this .subspecies attain a maximum snout-vent length of 80.7 mm., and females reach 101.4 mm. In a series of 16 males from El Zapote, Guerrero, Mexico, the snout-vent length is 6S.3 to 80.7 (mean, 75.8) mm.; the ratio of tibia length to snout- vent length is 0.372 to 0.417 (inean, 0.393); the ratio of foot length to snout-vent length is 0.326 to 0.360 (mean, 0..345); the ratio" of head length to snout-vent length is 0.343 to 0..379 (mean, 0.360), and the ratio of head width to snout-vent length is 0.201 to 0.248 (mean, 0.233). Fourteen females from the same locality have snout- vent lengths of 83.1 to 101.4 (mean, 89.3) mm. and do not difl^er from the males significantly in proportions. There is a geographic trend from north ( Go- lima ) to south ( Oaxaca ) in snout-vent lengths; both males and females from Oaxaca are noticeably larger than arc those from Colima (table 59). Structurally this subspecies is like the nominate subspecies, except that the edge of the labial flange tends to be slightly more serrate, and the fusion of the canthal ridges in large specimens is at a point between the nostrils, farther anteriorly than in the nomi- nate subspecies. The number of prcvomerine teeth on each process in reticulatus is five to eight, and a total number of prevomerine teeth is 11 to 16 (mean, 13.6). The general coloration of Triprion spatu- latus reticulatus is pale yellowish tan or pale olive-green with dark brown or black reticu- lations and spots on the dorsum (pi. 72, fig. 2). At night individuals from Tehuantepec, Oaxaca, Mexico, had a pale yellowish green dorsum fading to yellow on the flanks. The head was olive-brown. The dorsal reticula- tions were dark brown, and the venter, in- cluding the vocal sac was white. The iris was pale gold flecked with black. Specimens from EI Zapote, Guerrero, Mexico, tended to be more greenish tan with dark brown reticula- tions. Some specimens from the northern pait of the range (Golima) have dark markings consisting of dashes or flecks, and approxi- mately one-third of the specimens have rather fine reticulations on the back. The other specimens from Golima and all of those from farther south have a dorsal pattern consisting of bold reticulations or of reticulations and spots ( see fig. 301 for examples of these color patterns). A slight, but continual, cline exists for an increase in the amount of dark pigmen- tation on the dorsum fiom north to south (table 59). In preser\ati\e the dorsum is creamy tan to pale grayish brown with dark brown or black markings. The markings on the dorsal surfaces of the limbs tend to form bold re- ticulations rather than transverse bands. The venter is uniformly creamy white. Tadpoles: The only a\'ailable tadpoles are recent hatchlings that arc unsuitable for a diagnostic description. Mating Gall: The call of Triprion spatu- latus reticulatus consists of a single, low- pitched note "braaa." The note repetition rate varies from 10 to 17 (mean, 13.1) notes per minute; individual notes have a duration of 0.76 to 0.93 (mean, 0.85) of a second. The pulse rate is 88 to 114 (mean, 99.0) pulses per second. The fundamental frequency varies from 89 to 134 (mean, 103) cycles per 1970 DUELLMAN: HYLID FROGS 637 second, and the dominant frequency varies from 15S9 to 1869 (mean, 1745) cycles per second. There are no definitely emphasized harmonics above the dominant frequencv (pi. 34, fig. 2). Natural History: This subspecies inhab- its tropical scrub forests where the rainy sea- son is restricted to the months of June to Sep- tember. Peters (1955) found a breeding chorus at Ostula, Michoacan, on July 14, 1950; I found the species breeding near Te- huantepec, Oaxaca, on July 5, 1956, near Salina Cruz, Oaxaca, on July 6, 1958, and near El Zapote, Guerrero, on June 12, 1964. In each instance, heaw rains preceded the congregation of the frogs at the breeding sites. At the localities in Oaxaca, males were calling from bare mud or gravel banks near temporary ponds, although a few males called from distances of three meters from the water. At El Zapote, Guerrero, some males were ob- served calling from shallow water at the edge of the pond, but most were calling from barren ground near the ponds and up to dis- tances of 10 meters from the water. Several amplectant pairs have been observed on land, but none has been seen in the water. Taylor (1942b) reported that the type specimen of reticidatus was found in a brome- liad. Another specimen from near Tehuante- pec was found inside a rotting log. Remarks: The minor differences in size and structure between Triphon spatulattis as known in Sinaloa, and those populations to the south, previously referred to the species reticiilafus do not seem to be taxonomically significant. The major differences between northern and southern populations are in the color pattern. Although genetic interchange between the populations herein referred to reticidatus and the northern spatidatus can not be demonstrated at this time, the two populations are considered to be subspecifi- cally related, because of their general struc- ture similarities and because of my desire to emphasize the similarities, rather than to place the two populations on a status equal to that accorded to petasatiis and the species spaiu- latus. Etymology: The specific name is Latin, meaning made like a net, and refers to the dorsal coloration. Distribution: Triprioi^ spatulattis reticu- latus inhabits coastal lowlands of low foothills to cle\ations to about .350 meters from Colima southeastward to the Isthmus of Tehuante- pec, Oaxaca, Mexico; this species also occurs in the Balsas Basin in Michoacan (fig. 302). See Appendix 1 for the locality records of the 432 specimens examined. Triprion petasatus (Cope) Fhanjngodon petasatus Cope, 1865b, p. 193 [holo- type, U.S.N.M. No. 12287 from Cenote Tamache (17 kilometers north of Merida, Yucatan, Mexico; Arthur Schott collector]. Triprion petasatus Cope, 1866a, p. 127. Boulenger, lS82a, p. 431. Gunther, 1901 (188.5-1902), p. 293, Kellogg, 19.32, p. 138. Smith and Taylor, 1948, p. 70. Stuart, 1963, p. 42. Trueb, 1970a, p. 602. Diagnosis: This is a moderately large spe- cies (males to 60.8 mm.; females to 74.2 mm.) that is readily distinguished from other Mid- dle American casc^ue-headed hylids by having a large, upturned prenasal, which with the expanded maxillaries forms a broad labial shelf. Furthermore, petasatus has a large dermal sphenethmoid, paired \'ocal sac, and lacks odontoids on the palatines. Triprion spattdaiiis has a single, median vocal sac, odontoids on the palatines, a prenasal that is not upturned, and no dermal sphenethmoid. Pternohyla has only moderate labial flanges and lacks a dermal sphenethmoid and pre- nasal. Furthermore, Pternohyla is squat and toad-like in appearance and has a large spade- like inner metatarsal tubercle. Anotheca lacks labial flanges and has spines on the supra- tympanic and occipital ridges. Description: Males attain a maximum snout-\'ent length of 60.8 mm., and females reach 74.2 mm. In a series of 20 males from Chichen Itza, Yucatan, Mexico, the snout-vent length is 48.1 to 60.8 (mean, 54.6) mm.; the ratio of tibia length to snout-vent length is 0.374 to 0.414 (mean, 0..393); the ratio of foot length to snout-vent length is 0.305 to 0..351 (mean, 0..3.32); the ratio of head length snout-vent lengths of 65.0 to 74.2 (mean, 0.334); the ratio of head width to snout- vent length is 0.255 to 0..302 (mean, 0.274), and the ratio of the diameter of the tympanum to that of the eye is 0.473 to 0.605 (mean, 0.548). The ten females from the same locality have snout-vent lengths of 65.0 to 75.2 (mean, 70.7) mm. and show no significant differences 638 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 in proportions from the males. Specimens from the southern part of the range ( La Lib- ertad, El Peten, Guatemala) are smaller (mean snout-vent length in 20 males, 52.1 mm.) and have proportionately longer legs and smaller heads (see Duellman and Klaas, 1964, p. 312). The head is large and modified in the form of a bony casque with the skin com- pletely co-ossified with the skull (fig. 298). The maxillaries and the prenasal are greatly expanded and form a broad labial shelf. The snout protrudes far beyond the leading edge of the lower jaw, and the tip of the snout is upturned. The edge of the labial shelf is serrate. The nostrils are directed dorsally at a point about two-thirds of the distance from the eyes to the tip of the snout. A bony pre- orbital knob is present at the anterior edge of the orbit; in some individuals, especially large females, the knob is greatly enlarged so as to overhang the anterior edge of the orbit. A sharp canthal ridge extends from the preorbital knob to a point just posterior to the nostril. From the point of confluence of the canthal ridges a low bony ridge extends an- teriorly between the nostrils to the tip of the snout. The loreal region is deeply concave. A bony preorbital ridge forms the anterior border of the orbit and extends \cntrall\' from the preorbital knob to the labial flange, which is narrow posterior to the preorbital ridge. The eyes are large, protuberant, and directed anterolaterally. A bony supratympanic ridge extends from the posterior edge of the orbit to the posterior edge of the skull; the ridge overhangs the upper edge of the tympanum, which otherwise is distinct. The posterior edge of the skull is delimited by a finely ser- rate transverse bony ridge, which is continu- ous in some specimens, but notched medially in most individuals. The upper arms are slender, and the fore- arms are robust. An axillary membrane and tubercles on the ventrolateral edge of the forearm are absent, but a distinct transverse dermal fold is present on the wrist. The fingers are moderately long and robust and bear large discs; the diameter of that on the third finger is about equal to the diameter of the tympanum. The subarticular tubercles are large and round; none is bifid. The super- numerary tubercles are moderately large and round. A large, flat palmar tubercle is pres- ent. The prepollex is moderately enlarged and in breeding males is covered with a horny nuptial excrescence, which in most individ- uals extends along the inner edge of the thumb to the disc. Webbing is lacking be- tween the first and second fingers and is rudi- mentary between the others ( fig. 300C ) . The legs are short; the adpressed heels barely overlap. The tibiotarsal articulation extends to the posterior edge of the tympanum. A well-defined tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is large, flat, and elliptical; the outer metatar- sal tubercle is minute and round. The toes are long and bear discs that are slightly smaller than those on the fingers. The sub- articular tubercles are round and somewhat larger than the small, round supernumerar}' tubercles. The toes are about two-thirds webbed (fig. 300D). The webbing connects the first and second toes at the bases of the penultimate phalanges and extends from the middle of the penultimate phalanx of the second to the middle of the antepenultimate phalanx of the third, from the base of the disc of the third to the base of the penultimate phalanx of the fourth and on to the base of the disc of the fifth toe. The anal opening is directed posteriorly at the upper level of the thighs. No anal flap is present, but a dermal fold extends postero- ventrally from a point on either side of the anal opening. The skin is smooth on the dorsum (except head), chin, and \'entral sur- faces of the limbs (except thighs); it is granu- lar on the flanks, belly, and \entral surfaces of the limbs. The tongue is round, slightly wider in front than behind, and barely free posteriorly; it is shallowly notched posteriorly in most individuals and marginate in some specimens. In most specimens the dentigerous processes of the prevomers are transverse or slightly curved, whereas in some specimens the processes are inclined posteromedially. The processes He between the moderately large ovoid or longitudinally elliptical cho- anae. Males have a total of 8 to 15 (mean, 11.6) prevomerine teeth, and females have 14 to 20 (mean, 16.1). The vocal slits extend from the posterolateral edge of the tongue 1970 DUELLMAN: HYLID FROGS 639 to the angles of the jaws. The \ocal sac is subgular, paired, and situated posteriorly on the throat. The general coloration of Triprion peta- satus is olive-green or tan with dark brown or black markings on the dorsum (pi. 72, fig. 1). In most males the dorsum is olive- green with dark brown or black irregularly shaped blotches, spots, or numerous flecks on the back. The dorsal surfaces of the limbs are colored like the body and have distinct dark brown or black transverse bands on the shanks and forelimbs; the bands are indis- tinct or lacking on the thighs and the feet in some specimens. The flanks are oli\e-green or yellowish green. Most females are pale tan, and some are olive-brown; all have dark brown or black markings. In specimens of both sexes the posterior surfaces of the thighs are dark brown or reddish brown, and the anterior surfaces are pale brown. The head is colored like the body but lacks dark mark- ings. In some individuals silvery gray flecks are present on the dorsum; these are most apparent on the head. The belly is white, and the \entral surfaces of the shanks and feet are tan. In breeding males the vocal sac is yellow with brown flecks. The iris is golden bronze with fine black reticulations. In preservative the dorsum varies from grayish tan to olive-brown with dark brown markings. Small white flecks are present on the dorsal surfaces of the head, body, and limbs in some individuals. A few specimens lack dorsal markings. The posterior surfaces of the thighs are dark brown, and the anterior surfaces are pale brown. The ventral surfaces of the forearms and thighs are creamy tan and those of the shanks and feet are brown. The throat and belly are creamy white with some brown pigment posterolaterally on the throat in some females and in most males; in tlie other males the entire throat is brown. T.^Di'OLES: The embryonic and larval de- velopment were described in detail by Duell- man and Klaas ( 1964 ) , who noted that the oral suckers persisted into developmental stage 24 and that the teeth were not fully developed until stage 30. Measurements of the tadpoles showed that there is a gradual increase in the length of the tail relative to body length through stage 41. Duellman and Klaas (1964) noted that a great variation in size occurred in developmental stage 25 and suggested that the rate of growth is more rapid in that stage or that the duration of the stage is longer than that of other stages. Throughout development the head and body become darker; the amount of pigment in- creases in the ventral fin, and the pattern of pigmentation of the fins changes from flecks to reticulation and finally to Ncnation. A typical tadpole in developmental stage 30 has a body length of 12.3 mm. and a total length of 27.0 mm. The body is ovoid and slightly wider than deep. In dorsal profile the snout is bluntly rounded and in lateral profile acutely rounded. The nostrils are dorsal in position about two-thirds of the dis- tance from the eyes to the tip of the snout and directed dorsolaterally. The eyes are moderately small and dorsolateral. The long, sinistral spiracle has its opening just below the midline at a point about midlength on the body. The anal tube is short and dextral. The tail is moderately deep and pointed ter- minally. The caudal musculature is moder- ately heavy and does not extend to the tip of the tail. At the midlength of the tail the depth of the musculature is equal to the depth of either fin. The dorsal fin extends onto the body and is deepest at midlength of the tail; the ventral fin is deepest at about one-third of the length of the tail (fig. 303). Fig. 303. Tadpole of Triprion petasattis, K.U. No. 71731. x 4. 640 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 In life the tadpoles are dull grayish brown with creamy tan caudal musculature and transparent fins with brown reticulations; the iris is pale bronze. In preservative the dor- sum is dark brown, and the venter is pale brown. The caudal musculature is creamy gray. The ventral fin lacks pigment, and the dorsal fin is venated. The mouth is moderately small and antcro- ventral. Lateral folds and a shallow ventral fold are present in the lips, which are bor- dered by one row of small papillae, except on the median part of the upper lip, which is bare. Small papillae are present in the lateral folds. The beaks are moderately heavy and bear small, pointed serrations. The upper beak is in the form of a high arch and has long, slender lateral processes. The lower beak is broadly V-shaped. There are two up- per and three lower rows of teeth. The upper rows are about equal in length, and the sec- ond lower row is narrowly interrupted medi- ally. The lower rows are complete; the first lower row is nearly as long as the upper rows, and the other lower rows are progressively shorter (fig. 304). In tadpoles in developmental stage 34 the canthal ridges are apparent, and those in stage 41 have a weak occipital ridge. In stage 45 the tadpoles have obvious canthal and occipital ridges and have an acutely angular snout that projects well beyond the leading edge of the lower jaw. Mating Call: The call of Triprion peta- satiis consists of a single, low-pitched note. The notes are quickly repeated, so that the call sounds like the quacking of a duck. The Fic. 304. Mouth of tadpole of Triitrion pctasalus K.U. No. 71731. X 25. frogs normally produce 33 to 54 (mean, 41) notes in succession. The note repetition rate is 45 to 52 ( mean, 48.7 ) notes per minute. Each note has a duration of 0.26 to 0.39 ( mean, 0.30 ) of a second and a pulse rate of 80 to 90 (mean, 84.7) pulses per second. The fundamental frequency varies from 210 to 350 ( mean, 287 ) cycles per second, and the domi- nant frequency varies from 1900 to 2450 (mean, 2096) cycles per second. Usually five harmonics above the dominant frequency are emphasized with decreasing force from the lowest to the highest (pi. 34, fig. 3). Natural History: The following account is excerpted from Duellman and Klaas ( 1964, pp. 312-315); the reader is referred to their account for more details on habitat and life history. Triprion petasatus inhabits low, xe- rophilous forest or savannas in areas charac- terized by shallow soils and a low amount of rainfall that is highly seasonal in distribution. Breeding activit)' follows rains which pro- vide water in solution pits, sink holes, and aguadas. Stuart (1935, p. .37) found the spe- cies breeding in an intermittent agxiada at La Libcrtad, El Peten, Guatemala between May 23 and 30, 1933. My own observations on breeding activity of Triprion petasatus were made in July, the month in which most other persons have observed the species (Gaige, 1936, p. 290, and Maslin, 1963, p. 3). On July 22, 1962, T. petasatus was breed- ing at localities 9 and 12 kilometers east of Chichen Itza, Yucatan, Mexico. At the first locality males were calling from branches of low trees and bushes around two small solu- tion basins. At the latter locality males and clasping pairs were on the ground at the edge of a water-filled earthen pit. On the same night a large breeding congregation was ■^ound at a locality 3.5 kilometers east of Yokdzonot, Yucatan, where males were call- ing from branches of dense trees and bushes around a small solution pit; amplexing pairs were on branches to heights of 2.5 meters above the ground. Because most calling males and many clasping pairs were observed in trees, probably the frogs spend the days and the dry season in trees. Stuart (1935, p. 37) found individuals in holes in trees around an aguada in which the species was breeding at La Libertad, Guatemala. Stuart observed that 1970 DUELLMAN; HYLID FROGS 641 the frogs plugged the cavities in trees with their heads. Eggs are deposited in clumps in the wa- ter; in Yucatan eggs were found in shallow basins or solution pits. Tadpoles congregate in shaded areas and seek refuge in the decay- ing vegetation on the bottom of the basin or pit. Four reccnth- metamorphosed }oung ha\e snout- vent lengths of 15.5 to 16.1 (mean, 15.8) mm. These specimens have a protrud- ing snout and slightly flared lips. Remarks: The developmental and inter- nal cranial osteology of this species has been studied in detail (Trueb, 1970a). Etymology: The specific name petasatus is Latin meaning with a hat on and refers to the helmet-like casque. Distribution: Triprion petasatus occurs in the lowlands of the Y'ucatan Peninsula, southward in subhumid habitats to the sa- vannas of central El Peten, Guatemala (fig. 305). See Appendix 1 for the locality records of the 273 specimens examined. Fig. 305. Distribution of Triprion petasatus. Genus Pseudacris Fitzinger Pseudacris Fitzinger, 1843, p. 31 [type species, Rana nigrita LeConte, 1824, by nionotypy]. Ctwropliilus Baird, 1854, p. 59 [type species, Rana nigrita LeConte, 1825, by original designation]. Ilclocaetes Baird, 1854, p. .59 [type species, Htjla tiiscriata Wied, 1839, by subsequent designation (Schmidt, 1953)]. Generotype: The first usage of the name Pseudacris was in a subgencric position under Acris by Fitzinger ( 1843, p. 31 ) : "Pseuda- cris . . . Am. . . . Acr. nigrita Dum. Bibr." Dumeril and Bibron (1841, p. 509) used the combination Acris nigrita for the frog orig- inally named Rana nigrita by LeConte ( 182.'), p. 282). Since Fitzinger associated no other species with Pseudacris, Rana nigrita LeConte is the type species by monotypy. Etymology: The generic name is derived from the Greek pseudes, meaning false, and the Greek akris, in this case referring to the genus Acris. Definition: The frogs in this genus are small pond-breeding species; males attain snout-vent lengths of 41 mm. and females, 46 mm. The dorsum is tan, gray, or green with darker stripes or spots arranged in longi- tudinal series. All have a dark line from the nostril to the eye; the line is expanded pos- terior to the eye and in some species continues to the groin. In most species, a pale labial stripe is present. The webbing is vestigial on the hand, and the toes are less than one-third webbed. The discs are barely wider than the digits. A tarsal fold is absent, and dermal appendages on the limbs and an axillary membrane are lacking. The skin is smooth dorsally and not invoKed in co-ossification with the skull. Males have a single, median, subgular vocal sac but lack horny nuptial excrescences. The skull is weakly ossified and has a large frontoparietal fontanelle (fig. 306). The sphenethmoid is ossified anteriorly between the nasals to the end of the septum nasi. The nasal is moderately long and at least partially in bony contact with the sphen- ethmoid. The squamosal is not in bony con- tact with the crista parotica, and the anterior arm of the squamosal extends only about one-third of the distance to the maxillary. The columella is expanded distally. The quadra tojugal is present and articulates with 642 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 the maxillary. The prevomer is poorly ossified and the palatine is weak. The medial ramus of the pterygoid does not articulate \\'ith the prootic. Teeth are present on the premaxil- laries, maxillaries, and prevomers. The tad- poles have deep fins and small anteroxentral mouths with two upper and three lower rows of teeth. The mating calls consist of a series of quickly repeated notes, which in some species are so closely spaced that the call sounds like a trill. The chromosome numbers are )! = 12, 2/i=24 (known only in P. hracluj- phona and triseriata). Composition of Genus: Seven species are included in the genus; three of these are poly- typic. Only one species, Pseitdacris clarkii occurs in Middle America, and two Mexican specimens of that species have been exam- ined. Distribution: North America westward to the Rocky Mountains, northward to Hud- son Bay and northwestern Canada and south- ward to the Gulf of Mexico. In Middle Amer- ica, the genus occurs only in the lower Rio Grande Valley. Discussion: The frogs of the genus Pseu- daciis differ from most North and Middle American llyla by having small discs and greatly reduced webbing on the feet. No other external features will distinguish them from Hyla. If these frogs occurred in South America, they probably would not have been recognized gencrically. Pseitdacris seems to be more closely re- lated to the Hyla eximia group than to any other groups of Hyla or to Acris. Pscudacris differs from members of the Hyla eximia group by having a more extensively ossified sphenethmoid and better developed nasals which are in contact with the sphenethmoid. Thus, by comparison with Pseudacris, the skulls of Htjla eximia and its allies are re- duced, whereas the webbing of the feet and the sizes of the discs are reduced in Pseuda- cris as compared with Hyla eximia. The morphological similarities of the adults and tadpoles, the likeness of breeding habits, the general structural similarities of the mating calls, and the nearly complemen- tary geographic ranges of Pseudacris and the Hyla eximia group strongly suggest close phy- letic relationships between the groups. Possi- bly they both descended from a widespread Nearctic prototype, which gave rise to Pseu- dacris, in eastern North America and to the Hyla eximia group in western North America. A discussion of the intrageneric relation- ships of Pseudacris is inappropriate here. The \arious species have been reviewed by Schwartz (1957) and Smith and Smith (1952), and experimental evidence on repro- ductive isolating mechanisms was summa- rized by Mecham (1965). Pseudacris clarkii (Baird) Hclocaetcs clarkii Baird, 1854, p. 60 [svntypes, U.S.N.M. No. 3313 (fide Cochran, 1961, p 5b),is from Galveston, Galveston County, Texas; M. Dean collector]. Chorophilus triseriatus clarkii: Cope, 1875, p. 30. Pseudacris triseriata clarkii: Burt, 1932, p. 80. Pseudacris clarkii: Smith, 1934, p. 462. Diagnosis: This small, slender species with a subacuminate snout has a dorsal pat- tern of irregular dark green to reddish brown spots on a pale green, tan or gray ground color; a pale labial stripe is present; a dark interorbital triangular mark, not bordered by white usually is present. This color pattern, in combination with smooth skin, toes less than one-third webbed, and barely enlarged terminal discs on the digits distinguishes Pseudacris clarkii from other hylids. The only other small hylids in Mexico with a triangular interorbital mark are Acris crepitans, Hyla regilla, and Hyla sfaufferi. The former has the interorbital mark usually bordered by white, tubercular dorsal skin, and much more webbing on the feet. Hyla staufferi and Hyla regilla have a linear pattern on the dorsum, more webbing on the feet, and larger discs than Pscudacris clarkii. Desciuption: Males of this species attain a maximum snout-\ent length of 29 mm., and '■'Baird (1854, p. 60) did not designate type specimens liut stated that the habitat was "Galveston and Indianola, Texas." Yarrow (1882, p. 170) listed only U.S.N.M. No. 3313 under "Chlomphilus tri- .scriattis clarkii." Cope (1889, p. 347) listed the same specimen, plus U.S.N.M. No. 3317 from Indianola and U.S.N.M. No. 3315 from between Indianola and San Antonio, Texas; both were collected by John H. Clark and presumably along with U.S.N.M. No. 3313 formed the t\pe .series for Baird's description of Hclocaetcs clarkii. 1970 DUELLMAN: HYLID FROGS 643 Fig. 306. Dorsal (A) and ventral (B) views of the skull of Pscudacris clarkii, K.U. No. 60373. X 6. females reach 31 mm. The two known speci- mens from Mexico are juveniles having snout- vent lengths of 15 and 18 mm. The head is narrower than the body, and the top of the head is barely convex. In dorsal profile the snout is acuminate, in lateral pro- file, it is acuminate and projects beyond the margins of the lips. The snout is long, and slightly protuberant nostrils are situated at a point about two-thirds of the distance from the eyes to the tip of the snout. The canthus is round, and the loreal region is barely con- cave; the lips are moderately thick and not flared. A thin dermal fold extends posteriorly from the eye, above the tympanum, and downward to a point abo\e the insertion of the arm. The fold obscures the upper edge of the tympanum, which otherwise is distinct and separated from the eye by a distance equal to about two-thirds of the diameter of the tympanum. The arms are moderately long and robust; an axillary membrane is absent. There are no rows of tubercles on the ventrolateral edge of the forearm, but a distinct dermal fold is present on the wrist. The fingers are long and slender and bear discs that are only slightly wider than the fingers. The subartic- ular tubercles are moderately large and round; none is bifid. The supernumerar\' tu- bercles are moderately large and round. A large quadrangular palmar tubercle is present. The prepoUex is slightly enlarged and in breeding males does not bear a nuptial ex- crescence. The webbing on the hand is ves- tigial (fig. 307A). The legs are short and robust; the heels of the adpressed limbs barely overlap. The tibiotarsal articulation extends to the tympanum. A distinct transverse der- mal fold is present on the heel, and a well- developed, flap-like tarsal fold extends the full length of the tarsus. The inner metatarsal tubercle is small, elliptical, and elevated. A conical outer metatarsal tubercle is present. The toes are long and slender and bear very small discs; the subarticular tubercles are large and round, whereas the supernumerary tubercles are barely evident only on the proxi- mal segments of each digit. The toes are webbed only basally ( fig. 307B ) . The anal opening is directed posteriorly near the upper level of the thighs; a short, broad anal sheath is present. The skin on the dorsum is weakly granular, whereas that on the venter is strongly granular. The tongue is cordiform, shallowly notched posteriorly, and barely free behind. The dentigerous pro- cesses of the prevomers are small rounded elevations that are widely separated medially and lie between the ovoid choanae. Usually there are only two or three teeth on each ele- \ation. The vocal slits extend from the mid- lateral base of the tongue to the angles of the jaws. The vocal sac is single, median, sub- gular, and greatly distensible. The general coloration of Pseudacris clarkii is pale green or olive-green above with elongate brown spots usually forming three rows on the back (pi. 64, fig. 4). The dorsal surface \'aries from pale gray to green to dull olive-gray. The spots on the back and transverse bars on the limbs vary from brown to dark olive-green. There is a dark brown 644 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 stripe from the nostril, to the eye, and onto the anterior part of the flank. A narrow cream labial stripe is present. The venter is creamy white. The iris is pale bronze with black flecks. In preservative, the dorsum varies from pale tan to grayish brown; the dorsal markings are to dark brown, and the venter is creamy tan. Fig. 307. Hand (A) and foot (B) of Pseudacris clarkii, K.U. No. 110232. x 8. Tadpoles: No tadpoles of this species are available from Middle America; the following description is based on individuals from Ar- lington, Te.xas, provided by William F. Py- burn. A typical tadpole in developmental stage 33 has a body length of 8.8 mm. and a total length of 23.0 mm. The body is deeper than wide; in dorsal profile the snout is blunt- ly rounded, and in lateral profile it is round. The eyes are small, widely separated, and directed dorsolaterally. The nostrils are di- rected anterolaterally at a point about mid- way between the eyes and the tip of the snout. The spiracle is directed posteriorly at a point below the midline and about three-fifths of the distance from the snout to the posterior edge of the body. The anal tube is short and dextral. The caudal musculature is slender and extends to the tip of the pointed tail. The caudal fins are deep; at midlength of the tail the depth of either fin is half again the depth of the caudal musculature. The dorsal fin extends onto the body (fig. .308). In preservative the tadpoles are dark brown or nearly black above, and the venter is transparent. The caudal musculature is pale creamy tan below and dark brown above. The caudal fins are transparent and marked by a few small black flecks. The mouth is moderately small and situ- ated anteroventrally. The median part of the upper lip is bare; elsewhere the lips are bor- dered by one or two rows of small papillae. The beaks are slender and bear short, pointed serrations. The upper beak is very broad and has a short, blunt lateral processes; the lower beak is broadly V-shaped. There are two up- per and three lower rows of teeth. The upper rows are much longer than the lower ones, and the second upper row is broadly inter- rupted medially. The first and second lower rows are much longer than the third lower row, and the first lower row is narrowly inter- rupted medially in some specimens (fig. .309). Mating Call: The mating call of Pseuda- cris clarkii consists of a series of quickly re- peated, low-pitched notes. Analysis of the calls of four individuals from Montgomery County, Kansas, indicates the call rate is 130 to 160 (mean, 144) notes per minute. The duration of the note varies from 0.15 to 0.18 (mean, 0.17) of a second, and the notes have 1970 DUELLMAN: HYLID FROGS 645 Fic. 308, Tadpole of Pseudacris clarkii, K.U. No. 116932. x 5. 90 to 97 (mean, 93) pulses per second. The fundamental frequency varies from 74 to 83 (mean, 78) cycles per second, and the domi- nant frequency varies from 2508 to 2652 (mean, 2554) cvcles per second (pi. 37, fig. 1). Natural History: Pseudacris clarkii in- habits prairie and subhumid scrub land. The species breeds at the time of the spring rains between early March and late June. Males call from clumps of grass in shallow water. The tadpoles develop in shallow grassy ponds. Remarks: Pseudacris clarkii is known from Mexico on the basis of two specimens (S.U. Nos. 15449 and 15450) from 8 kilo- meters west of Matamoros, Tamaulipas (Lynch, 1965a, p. 31). Etymology: The specific name is a patro- nym for John H. Clark, the collector of the type specimen. DisTRLBUTioN : Pscudacris clarkii occurs in the central United States from south-cen- tral Kansas to the Gulf of Mexico; the species is known in Mexico only from the lower Rio Grande Valley in Tamaulipas (fig. 310). See Appendix 1 for the locality records of the two specimens examined. Genus Acris Dumeril and Bibron Acris Dumeril and Bibron, 1841, 1. 506 [t\pe species Rana grylhis LeConte, 1825, by fiat]. Generotype: Dumeril and Bibron (1841) included Raiw grylhis LeConte, 1825, and Rana nigrita LeConte, 1825. Neither was des- ignated as the type of the genus, although grylhis was listed first (page 507; nigrita was treated on page 509). Fitzinger (1843, p. 31) proposed the following arrangement: "1. Gen. Acris. Dum. Bibr. Pseudacris . . . Am. . . . Acr. nigrita. Dum. Bibr. Acris . . . Am. . . . Acr. gryllus. Dum. Bibr." hus, by selecting nigrita (one of the two 100° 98° V. 1 1 °\ \ > o 4. crepitans 28° V. • P. clarkii / ?R° k "y t III < \l s I ze" 0 100 "^ V.'l' z€ KILOMETERS 1 100° 98° Fig. 309. Mouth of tadpole of Pseudacris clarkii. K.U. No. 116932. x 30. Fig. 310. Distribution of Pseudacris clarkii and Acris crepitans in Mexico. 646 MOiNOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 species included in Acris by Dumeril and Bib- ron) as the type species of the genus Pseu- clacris Fitzinger, by fiat restricted grylhis to Acris. Etymology: Dumeril and Bibron (1841, p. 506) noted that the generic name was Greek, "Akpis, I'un des noms de la Sauter- elle." Thus, the generic name is a name for a grasshopper and is appropriately applied to these frogs capable of prodigious leaps. Definition: Members of this genus are small pond-breeding species; males attain snout-vent lengths of 29 mm. and females 34 mm. The dorsum is pale brown or gray usually with a dark interorbital triangular mark and with or without a green or rusty tan middorsal patch. A prominent longitudi- nal black bar is present on the posterior sur- faces of the thighs. The webbing is vestigial on the hand, and the toes are about three- fourths webbed; the terminal phalanges are not expanded. A tarsal fold is present and dermal appendages on the limbs and an ax- illary membrane is lacking. The skin on the dorsum is tubercular and not involved in in- tegumentary-cranial co-ossification. Males have a single, median, subgular vocal sac and lack horny nuptial excrescences. The skull is weakly ossified and has a large fronto- parietal fontanelle (fig. 311). The spheneth- moid is greatly reduced. The nasals are small and widely separated medially; they are not in contact with the sphenethmoid or maxillar- ies. The squamosal is not in bony contact with the crista parotica, and the anterior arm of the squamosal extends only about half of the distance to the maxillary. The quadrato- jugal is present and in contact with the maxil- lary. The prevoniers are greatly reduced and do not articulate with the maxillaries or pre- maxillaries. The palatine is slender and not in bony contact with either the maxillary or the sphenethmoid. The medial ramus of the pterygoid is not in bony contact with the prootic. Teeth are present on the premaxil- laries, maxillaries, and prevomers. The tad- poles have moderately deep fins and small anteroventral mouths with two upper and two lower tooth rows. The mating call con- sists of a long series of short clicking notes. The chromosome numbers are n=ll, 2n=22 (Cole, 1966). Fic. 311. Dorsal (A) and ventral (B) views of the skull of Acris crepitans, K.U. No. 59952. X <3- Composition of Genus: Two species (A. crepitans and A. gryUiis) are generally rec- ognized, although some previous workers have suggested that these two species intergrade. Two subspecies usually are recognized in gryllus, whereas the status of subspecies {hlanchardi and pahidicola) of crepitans is in question. Only crepitans occurs in Middle America, and 1 have examined 33 preserxed frogs from Mexico. Distribution: The genus occurs through- out eastern United States from New York and Michigan to South Dakota and eastern Colo- rado and southward in isolated populations through eastern New Mexico and western Texas to Coahuila, Mexico. Discussion: The frogs of the genus Acris are distinctive among the hylids in a number of morphological and behavioral features. They are non-arboreal, aquatic-margin spe- 1970 DUELLMAN: HYLID FROGS 647 cies and thus fill the ecological position of a small Rana. The frogs are active and call by day, as well as at night. The eggs are de- posited singly or in small groups adherent to aquatic \egetation. In the tadpoles, the en- tire upper lip is devoid of papillae, and the eyes are dorsal; in each of these characters, the tadpoles are like those of most North American Rana. The presence of long toes, no expanded digits, and a considerable amount of webbing are obvious adaptations for their semi-aquatic habits. The smooth skin on the throat and chest is unusual for a hylid and is more like the condition in Rana. The skull of Acris is so greatly reduced that the usefulness of cranial characters is re- stricted. Chantell (1968) noted the distinc- tiveness of Acris among North American hy- lids and suggested that it possibly was most closely related to Limnaoedus, which obvi- ously is a hylid with reduced cranial elements. Studies of chromosomes ( Duellman and Cole, 1965; Cole, 1966; Duellman, 1967b) have shown that Acris is unique among Holarctic and Neotropical hylids by having chromo- some numbers of 7i=ll, 2n=22; this number is common in Australian species of Hyla (Straughan, pers. comm.). Most hylids have n=12, 2(1=24, but some groups have haploid numbers of 13, 14, or 15 and diploid numbers of 26, 28, and 30. All ranids, for which chrom- osome data are available, have n = 13, 2n=26 chromosomes. Despite the divergent nature of Acris with respect to other hylids and the superficial similarity of Acris to ranids, the inescapable facts remain that Acris has procoelous verte- brae, an arciferal pectoral girdle, intercalary cartilages, and claw-shaped terminal pha- langes— a combination of characters that seemingly inextricably ally the genus with the hylids. Acris crepitans Baird Acris crepitans Baird, 1854, p. 59 [no types were designated; t\pe localitj': "Northern States generally; t>pe locality restricted to Albany, Albany County, New York by Smith and Taylor (1950, p. 359); Albany is approximately 100 miles north of the northeastern- most known locality for the species]. Smith and Tay- lor, 1948, p. 77. Diagnosis: This small species, with a gray or tan dorsum and an acutely rounded snout, is immediately distinguishable from all other Middle American hylids by the following combination of characters: tips of digits not expanded; skin on dorsum tuberculate, and that on throat and belly smooth; a black longitudinal bar, usually bordered above and below by creamy white bars, present on the posterior surface of the thigh. Description: Males of this small species attain a maximum snout-vent length of 29.0 mm., and females reach 34.0 mm. In a series of 10 males from the vicinity of Jimenez, Coa- huila, Mexico, the snout-\ent length is 20.3 to 23.6 (mean, 22.5) mm.; the ratio of tibia length to snout-vent length is 0.508 to 0.609 (mean, 0.563); the ratio of foot length to snout-vent length is 0.479 to 0.578 (mean, 0.541); the ratio of head length to snout-vent length is 0.328 to 0.389 (mean, 0.355); the ratio of head width to snout-vent length is 0.322 to 0.369 (mean, 0.348), and the ratio of the diameter of the tympanum to that of the eye is 0.346 to 0.654 (mean, 0.495). Five females from the same area have snout-vent lengths of 24.6 to 25.8 (mean, 25.3) mm. and do not differ significantly from the males in proportions. The head is narrower than the body, and the top of the head is barely convex. In dor- sal profile the snout is acutely rounded; in lateral profile it is round and slightly pro- truding beyond the margins of the lower jaw. The snout is long, and the nostrils are barely protuberant at a point about two-thirds of the distance from the eyes to the tip of the snout. The canthus is barely evident, and the loreal region is inclined to be moderately thick, around the lips. A thin dermal fold extends posteriorly from the eye and angles at a point above the tympanum downward to the insertion of the arm. The fold obscures the upper and posterior edges of the tympan- um, which is barely separated from the eye. The arms are moderately short and slen- der; an axillary membrane is absent. Two or three small tubercles are present on the ven- trolateral edge of the forearm, and a distinct transverse dermal fold is present on the wrists. The fingers are short and slender; the tips are not dilated into a disc. The subarticular tubercles are round; none is bifid. The super- numerary tubercles are absent. A large, ele- 648 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 vated palmar tubercle is present. The prepol- le.\ is barely enlarged, and in breeding males does not bear a horny nuptial excrescence. A vestige of a web is evident between the fingers (fig. 312A). The legs are long and robust; the heels of the adpressed limbs over- lap by about one-third of the length of the tarsus. The tibiotarsal articulation extends to the nostril or to the tip of the snout. A distinct transverse dermal fold is present on the heel, and an elevated, flap-like tarsal fold is present. Two or three tubercles are present on the outer edge of the tarsus. The inner metatarsal tubercle is elongately ovoid and rounded. The outer metatarsal tubercle is large and conical. The toes are long and slen- der and do not bear expanded discs. The subarticular tubercles are moderately small and subconical; the supernumerary tubercles are either absent or minute and few in num- ber on the proximal segments of the digits. The toes are about three-fourths webbed ( fig. 312B ) . The webbing extends from the base of the terminal phalanx of the first toe to the base of the terminal phalanx of the second and on to the base of the penultimate phalanx of the third, from the distal end of the penul- timate phalanx of the third to the base of the penultimate phalanx of the fourth toe and on to the base of the terminal phalanx of the fifth toe. The anal opening is directed posteroven- trally near the upper level of the thighs; a short, broad anal sheath is present. Two large and several small tubercles are present below the anal opening. The skin on the dorsum is tuberculate; that on the throat, chest, and ventral surfaces of the limbs is smooth, and the skin on the posterior part of the belly is weakly granular. The tongue is narrowly cordiform, shallowly notched behind, and barely free behind. The dentigerous pro- cesses of the prevomers are small, widely separated, posteromedially inclined processes between the small, ovoid choanae. Adults of both sexes have two, three, or four teeth on each process. The vocal slits lie along the median edge of the lower jaw. The vocal sac is single, median, and subgular. The general coloration of Acris crepitans is dull brown or dull gray with or without a differently colored middorsal stripe (pi. 64, Fic. 312. Hand (A) and foot (B) of Acris crepitans, K.U. No. 116930. X 8. 1970 DUELLMAN: HYLID FROGS 649 fig. 5 ) . A darker brown or dull green tri- angular shaped mark, with the apex directed posteriorly, usually is e\ident on top of the head. A pair of dorsolateral darker areas usu- ally are e\ident on the back, and dark brown trans\erse bands arc present on the dorsal surfaces of the limbs. The posterior surfaces of the thighs are marked by a longitudinal black or dark brown stripe, bordered above and below by broad creamy white stripes. Distinct creamy white or pale green spots or vertical bars are present on the upper lip, and a similarly colored stripe extends from the posteroventral edge of the eye to the angle of the jaw. The anterior part of the flank is dark brown or black, whereas pos- teriorly, the flanks are creamy white with brown flecks. The belly is pale creamy white, or stark white and the throat is suffused or flecked with gray or brown in breeding males. Some females also have flecks on the throat. The iris is pale bronze. I have not observed living frogs of this species from Mexico, and consequently I am unable to determine the nature of the dorsal stripe. Pyburn ( 1961 ) noted that there were four vertebral stripe colors in Acr(.s crepitans in Texas and Louisiana. He concluded that among the red, green, gray, and red-green stripes, that the green stripe is not permanent. He demonstrated that the presence of the green stripe at metamorphosis is determined by a single dominant gene and that the re- cessive homozygote is gray-striped. There is some evidence that green-striped frogs form a higher proportion in given populations in the eastern part of the range of the species than in the western part. Pyburn suggested that selection in relation to vegetation density might be the major cause for geographic differences in the frequency of the green stripe. In preservative, the dorsum is dull tan to dark gray; in many individuals, markings are barely discernible. In all individuals, the dark longitudinal stripe on the posterior surfaces of the thigh is evident; however, in some specimens from Mexico, there is no evidence of a pale stripe above the dark one. Tadpoles: No tadpoles of Acris are known from Mexico. Mating Call: The call of Acris crepitans consists of a prolonged series of short notes, sounding like "click-click-click-click." No re- cordings are available from Mexico. The analysis of a typical call from an individual in Douglas County, Kansas, reveals a note repetition rate of 128 notes per minute. The duration of the notes vary from 0.04 to 0.05 of a second, and there are approximately 70 pulses per second. The energy is spread throughout the frecjuency spectrum; the fundamental frequency is at about 175 cycles per second, and the dominant frequency is at about 3150 cycles per second (pi. 3.5, fig. 1). Natural History: Acris crepitans is an aquatic-margin inhabitant. The specimens from Mexico were obtained in riparian situa- tions along streams in otherwise arid regions. The males usually call from shallow water or floating vegetation. Remarks: Schmidt and Owens (1944, p. 100) provided the first definite record of this species from Mexico, based on one adult male and 10 recently transformed juveniles from La Lajita, on the Rio Sabinas, near Musquiz, Coahuila. Netting and Coin (1946, p. 253) discussed these specimens in relation to others from trans-Pecos Texas. In 1952, a field party from the University of Kansas obtained 21 specimens of Acris crepitans from the vicinity of Jimenez, Coahuila, Mexico. These two lo- calities are the only ones currently known for the species in Mexico. Milstead (1960) in- cluded Acris crepitans among the 14 relict species of the Chihuahuan Desert and sug- gested that Acris had invaded the Chihua- huan Desert during pluvial times. Etymology: The specific name is Latin, meaning rattling, and apparently refers to the call of this species. Distribution: Acris crepitans occurs prin- cipally at low elevations from New York and northwestern Florida westward to South Da- kota and eastern Colorado and New Mexico southward into Coahuila, Mexico (fig. 310). See Appendix 1 for the locality records of the 32 specimens examined. Nomina Dubita Two names based on specimens supposed- ly from Middle America cannot be assigned to known populations. In both cases holo- types are lost; thus, accurate determination 650 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 and comparisons are not possible. One other name obviously does not apply to a Middle American frog. The individual problems con- cerning each name are discussed below. Hyla cherrei Cope Hyla cherrei Cope, 1894, p. 195 [type unknown; type locality, Alajuela, Costa Rica; R. Alfaro collector]. Gunther, 1901 (1885-1902), p. 264. Taylor, 1952c, p. 846. ?Hyla inicroccphala microccphala, Duellman and Fouquette, 1968, p. .526. The holotype (the only specimen ever re- ferred to this name) is lost; consequently, it is necessary to rely entirely on Cope's (1894) description. On the assumption that Cope was correct when he stated "Manus almost without web; pes fully palmate" and gave the coloration as straw-colored and a narrow white stripe from the orbit to the sacrum, it is not possible with any degree of certainty to associate the name with any known popu- lation of hylid frog in Central America. The presence of a dorsolateral light stripe immedi- ately suggests Htjla microcephala and Hyla angiistilmeata; the latter differs from the de- scription of cherrei in other aspects of col- oration, size, and webbing. Duellman and Fouquette (1968, p. 527) tentatively, and perhaps correctly, placed cherrei in the sy- nonymy of microcephala. However, micro- cephala has the fingers about one-third webbed (more than cherrei) and the toes about three-fourths webbed (less than cher- rei). Obviously, the status of the name is open to question and probably can never be settled, unless the holotype is found. Hyla molitor O. Schmidt llyla molitor O. Schmidt, 1857, p. 11 [Psyntype, N.M.W. No. 16494; type locality, "Chiriqui-Flusse unvveit Bocca del toro," Panama; J. von Warszewicz collector]. Brocchi, 1882, p. 40. Gunther, 1901 ( 1885- 1902), p. 279. One faded specimen (No. 16494) in the collection of the Naturhistoriches Museum Wien purportedly is a syntype of this species. Dr. Josef Eisclt of that museum informed me (personal communication) that there is no documentation of the specimen other than a notation in Steindachncr's writing that the specimen is a syntype of Hyla molitor. The specimen agrees reasonably well with the detailed description given bv O. Schmidt (1858, p. 245). The snout-vent length is 36.5 mm. The fingers are slender, about one-fourth webbed, and bear small discs; the toes arc about two- thirds webbed. A strong tarsal fold is present, and a heav^ supratympanic fold obscures the upper part of the tympanum, which is less than one-half of the diameter of the eye. The anal region is slightly protruding, and a short anal sheath is present. There are four teeth on each of a pair of rounded elevations be- tween the smaller round choanae. The tongue is cordiform, flattened behind, and free pos- teriorly for about one-fourth of its length. No \ocal slits are evident; presumably the speci- men is a female. The dorsum is uniform pale brown, and the venter is creamy tan. If the specimen ac- tually is one of the three individuals on which Schmidt based his description, the distinctive colors ha\'e faded. The coloration was de- scribed by Schmidt (1858, p. 246): "Dorsum uniformly gray, more intensive on the back, fading away laterally and on extremities; in cvery-day life this blue color would be called Mueller's Blau. A delicately dotted black line runs on the canthiis rostralis from the opening of the nose to the corner of the eye. In the armpits, on the flanks and the thighs two of our three specimens have black marblings." ( Free translation from the German. ) The mention of blue color laterally and black marbling on the flanks and thighs caused Duellman and Trueb (1966, p. .322) to suspect that Hyla molitor might be the same as the species that they named Smilisca sila. However, details of the description and of the supposed syntype negate that possibil- ity. Cochran (1951, p. 58) listed, without qualification, Hyla puma Cope, 1885a, as a synonym of Hyla molitor. Schmidt (1857, p. 12) diagnosed -'Hyla molitor. Varict. marmorata. An nova spe- cies?" In 1858 (page 246) he described one individual having a snout-vent length of .38 mm. (5 mm. larger than the three specimens of molitor and slightly bolder dorsal colora- tion. Hyla marmorata O. Schmidt, 1857, is preoccupied by Bufo marmoratiis Laurenti, 1768 {—Hyla marmorata, Daudin, 1803). 1970 DUELLMAN: HYLID FROGS 651 Careful examination oi the supposed syn- type of IlyJa molitor and study of Schmidt's description by Charles F. Walker, Jay M. Savage, and me have resulted in our being unable to assign the name to any known population of Central American hylids. A possibilitN- exists that, except for the speci- mens obtained by Warszewicz, the species has not been discovered. A few years ago, I \\ould ha\e gi\cn credence to such a sugges- tion, but from 1964 through 1966, Charles W. Myers and I explored the lowlands and moun- tains of Bocas del Toro Pro\ince in Panama without finding frogs that were referable to the species of Htjla named by Schmidt. Grant- ed, this is only negative evidence, but when combined with the fact that molitor is unlike any Hijla known from Central America, we are advised to seek other possible explana- tions. Warszewicz obtained amphibians as a sideline to this plant collecting in Panama and Bolivia; apparently the amphibians were not indi\idually tagged. Consequently, the distinct possibility exists that some of the frogs reported by Schmidt as having originated in Panama actually came from Bolivia. Unfortu- nately, the herpetofauna of Bolivia is so poor- ly known that definite association of Schmidt's supposed Panamanian species cannot be made with known populations in Bolivia at this time. Hyla splendens O. Schmidt Hyla splendens O. Schmidt, 1857, p. 11 [holotype, Krakow No. 1008 (1340); type locality, "Chiriqui- Flusse unweit Bocca del toro," Panama; J. von War- szewicz collector]. Brocchi, 1882, p. 40. Giinther, 1901 (1885-1902), p. 266. Recent discovery of the holotype has pro- vided the opportunity to ascertain the status of this long unapplied name. The type is in rather poor condition; the color is greatly faded — no green mentioned by Schmidt ( 1S58, p. 244) is apparent. The specimen is a male having a snout-vent length of 51..3 mm. The skin is co-ossified with the fronto- parietals, nasals, and pars facialis of the max- illaries. The skin is smooth dorsally and granular ventrally. Apparently the frog is a member of the Andean complex of Gastro- theca, containing the species boliviana, mar- supiatum, and peruana. Obviously, the frog must have been obtained in South America by Warzsewicz and subsequently mislabeled. IJyIa splendens is not a member of the Mid- dle American fauna. Determination of the status of the specific name splendens in the genus Gastrotheca is beyond the scope of the present paper. Species Incjuirienda Hyla sp. Stuart (1948b, p. 38) in his description of two tadpoles collected by him on February 10, 1940, in Arroyo Las Palmas at Finca Los Alpes, Departamento Alta Verapaz, Guate- mala, stated: "The specimens arc of particular interest owing to the tremendous development of the mouth, to form a sucking disc. More- over, the lips are very broad and set with numerous, large papillae. These characters seem to indicate that the tadpole is specially adapted to life in swift waters, and the adults of so modified a tadpole undoubtedly live within the stream itself or in the vegetation abo\e it." On July 15, 1960, I obtained a single tad- pole of the same species in Arroyo Las Pal- mas, and on August 1, 1961, 1 obtained a large series from the same stream. The tadpoles were found in a quiet pool in a torrential stream, where the tadpoles adhered to stones on the bottom of the pool. Attempts to raise the tadpoles to metamorphosis were unsuc- cessful; one individual reached developmen- tal stage 41, at which time it had a body length of 17.8 mm. and a total length of 49.5 mm. Resorbtion of the tail had begun in this individual. The largest specimens available for study are in developmental stages .35 to 37. A typi- cal tadpole in developmental stage 35 has a body length of 17.5 mm. and a total length of 50.3 mm. The body is robust, depressed, and slightly wider than deep. In dorsal pro- file the snout is broadly rounded, and the posterior edge of the body is bluntly rounded. In lateral profile the snout gradually slopes anterovcntrally from the nostrils, which are about one-third of the distance from the eyes to the tip of the snout. The eyes are small, widely separated, and directed dorsolaterally. The spiracle is sinistral and directed postero- dorsally; the spiraeular opening is below the 652 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Fig. 313. Tadpole of Hyla sp., K.U. No. 68522. X 2.5. midline at a point about two-thirds of the distance from the snout to the posterior edge of the body. The anal tube is long and dex- tral. The tail is long and terminally rounded. The caudal musculature is robust and extends nearly to the tip of the tail. The depth of the tail is nearly constant throughout its length, and at midlength of the tail, the depth of the musculature is about equal to the depth of either fin. The dorsal fin does not extend onto the body (fig. 313). In life, the body is dark olive-brown above. The caudal musculature has alternating dark brown and yellowish tan blotches dorsally. The iris is pale bronze. In preservative, the body is dark above and pale gray laterally and ventrally. The caudal musculature is pale tan, except dorsally where elongate dark brown blotches, narrowly separated by tan, are evident in most specimens. In small tad- poles (stage 25) the tip of the tail is notice- ably darker than the rest of the tail. The dark pigment apparently disperses in larger indi- viduals. The tadpole in developmental stage 41 has large brown spots on the sides of the body. The mouth is very large, nearly as wide as the body, and directed \entrally. The lips are not invaginated laterally and form an en- tire labial disc, completely bordered by a row of small papillae. A single row of large pa- pillae are present medially to the anteroLateral part of the lip, and three or four rows of large papillae are present medially to the lower lip. The beaks are moderately robust and bear small serrations. The upper beak is broadly bell-shaped and lacks long lateral processes; the lower lip is broadly V-shaped. There are two upper and three lower rows of small teeth. All of the rows extend laterally to the Hps, and all are complete (fig. 314). Adults of three stream-breeding hylids are known from Finca Los Alpes — Plectrohyla giiatemalensis, Plectrohyla qtiecchi, and Pti/- cholnjla spinipoUex. The tadpoles of all three species are known, and none is like that de- scribed here. Tadpoles are known for all of the presently recognized species of hylids on the Atlantic slopes and highlands of Gua- temala. Consequently, we are forced to con- clude that the tadpoles from Finca Los Alpes belong to a species of frog not represented by adults in northern Central America. The tadpoles from Finca Los Alpes are unique among hylid tadpoles in nothern Cen- tral America and Mexico; stream hylids in those regions either have small mouths with two upper and two lower rows of teeth or large mouths with a proliferation of tooth rows; no other tadpole from northern Central .Amer- ica has an enlarged mouth and only two upper and three lower rows of teeth, a common condition in hylid tadpoles in the highlands of Costa Rica and Panama. Comparison of the tadpoles from Finca Los Alpes with those of several species from the Costa Rican high- FiG. 314. Mouth of tadpole of Hyla ,sp., K.U. No. 68522. X 8. 1970 DUELLMAN: HYLID FROGS 653 lands reveals that the tadpoles from Finca Los Alpes are very much like the tadpoles of Hyla pictipes (figs. 135 and 136). The body is broad and depressed in both, and the mouths are alike, except for two minor, but consistent, differences. In Hyla pictipes there are two complete rows of large papillae medi- ally to the small fringing row anteriorly, whereas one incomplete row is present in the Guatemalan tadpoles. Furthermore, the beaks are more robust in the Guatemalan tadpoles than in Hyla pictipes. The absence of other kinds of tadpoles in northern Central America having the morpho- logical characters of the tadpoles from Finca Los Alpes and the similarity of the tadpoles to those of Hyla pictipes and less so to mem- bers of the Hyla rividaris group, suggests that the unknown species of Hyla from Finca Los Alpes possibly is closely related to Hyla pic- tipes. Obviously, no conclusions can be reached until the adults are found, but be- cause of the discontinuity of montane hylids across the Nicaraguan Depression, I am skep- tical that the unknown frog in Guatemala is conspecific with Hyla pictipes. LIFE HISTORY Despite the extensive field work on hylid frogs in Middle America and the accumu- lated voluminous notes, far too little is known about the life histories of most of the species. The general aspects of life history are known for about two-thirds of the species, but de- tailed observations are available for only ten species. Pyburn (1963, 1966) reported on Agahjchnis caUidnjas and Smilisca cijanostic- ta, respectively. Zweifel (1964) and Pyburn (1967) reported on Phrynohtjas venulosa. Breder (1946) provided excellent observa- tions on Hyla rosenbergi, and Duellman ( 1963d ) gave a detailed account of Agahjch- nis annae. Duellman and Trueb (1966) gave notes on the life histories of the species of Smilisca and pro\idecl detailed data on S. phaeota. Duellman and Klaas ( 1964 ) pre- sented extensive notes on Triprion petasatus, and Trueb (1968a) included valuable life history information in her study of Hyla lan- casteri. Detailed notes on the life histories of Hyla miotyinpamiui and H. pseudoptima arc presented in this paper; furthermore, the tadpoles of 29 species arc described for the first time. However, the tadpoles of 28 spe- cies are unknown. Much still needs to be learned about the breeding habits and larvae of the great ma- jority of Middle American hylids; a nearly complete absence of knowledge exists con- cerning reproductive cycles, growth rates, and life spans. There is a great need for some basic autecological investigations and re- search of reproductive cycles. These kinds of investigations, by their nature, must be car- ried out over long periods of time by investi- gators residing in Middle America. BREEDING Because the males of most hylids have a voice, collectors are attracted to calling males. Consequently, information can be accumu- lated on the dates that males were calling either by the evidence presented in field notes or by the presence of distended vocal sacs in preserved specimens. With the full realiza- tion that males of some species may call when there is no breeding in the population, I have used the presence of calling males in order to determine breeding times in the Middle American hylids. Presence of gravid females would be a better indicator, but females of most species arc relatively scarce in collec- tions. The accumulated data are incomplete (no data available for 23 species) and are biased by two factors. The amount of field work in Middle America has been highly seasonal; most collectors have worked there in June, July, and August. My own field work has been less limited, but it has been concentrated in the same months with only about half as much time in February, March, April, and May, somewhat less in January, a meager amount in September and Decem- ber, and none in October and November. The only year-round field work carried out has been in Costa Rica and Panama. The season- al incidence of collectors doubtlessly is re- flected in the data on breeding activity. For example, in each of the ten months of the year that I have worked in eastern Mexico, I ha\e found Hyla miotympanitin breeding; I seriously doubt if breeding activity in this species ceases in October and November, but we have no data to prove otherwise. The data are biased further b\' the discrepancies in the amount of information axailable. Our knowledge of the breeding seasons of some species is based on scores of observations, whereas data on other species are available from only one or two obser\ations. For ex- ample, absence of records of breeding activity in Smihsca haudinii prior to early June on the Pacific lowlands of Mexico is accepted as \alid, because this is a common species in a well-known area. Howexer, the two dates for breeding activity in the poorly known montane Hyhi saJvadorensis onl\' indicate that the species does breed in June and July but do not provide any assurance that the breed- ing season is restricted to those months. De- spite the limitations of the data, some of the results are noteworthy. Tabulation of the number of species known to loreed in any gi\cn month shows that there is an increase from 44 in April, to 50 in May, to 70 in June, and then a decrease to 63 in July and 53 in August (based on data for 91 species). The average number of breeding 654 1970 DUELLMAN: HYLID FROGS 655 UJ o UJ Q. to cr UJ CO D 40 - Pond- breeders Stream -breeders - 30 - • / /* \ / '' '" \ / / "'^ \ // \ \ ^ - 20 / / / / / • / / / / / V / » / \ / \ / t / \ P \ / \ / \ 1 \ \ \ \ \ \ V 10 • / 1 1 1 1 1 1 \ \ / '•--■ A 1 1 1 1 1 I Jan. Feb. Mar. Apr. May June July Aug. Sept Oct. Nov. Fig. 315. Sea,sonal incidence of breeding activity' in Middle American hylid frogs. Dec. species in these five months is 56, whereas in the other seven months the average is 21; November with 12 breeding species is the lowest month. Although the peak of breeding activity is in June, species with different breeding sites have different seasonal preferences (fig. 315). For example, of the 29 species known to breed in February, 21 are stream-breeders, and only si.x are pond-breeders. This ratio declines to 21/11 in March, 24/16 in April, and 22/23 in May. In June, July, August, September, Oc- tober, and November, the pond-breeders out- number the stream-breeders, whereas in De- cember and January the reverse is true. Throughout much of Middle America May- November are the rainy months, and Decem- ber-April are the dry months. Thus, there seems to be a close positive correlation in time of breeding in most pond-breeders with the rainv season, whereas a less noticeable negative correlation exists with the stream - breeders. The available data suggest that the brome- liad-breeders have extensive breeding seasons, which in Hy/fl cletuJroscarta and Anotheca spinosa extend from January through August. Breeding activity is known for four brome- liad-breeders in June. Even less information is available on those species that carry their eggs. Gravid females and females carrying eggs or young are known from May, Septem- ber, and December in Hemiphractus pana- 7nensis. Calling males of Gastwtheca cerato- phnjs have been heard in March through July, and those of Gastrotheca nicefori have been heard in January. The seasonal activity in many stream- breeding species is understandable when one realizes that most of these species inhabit humid montane forests that are moist through- out the year. Secondly, and perhaps more 656 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 significantly, the streams are usually clear and more quiet in the drier seasons. At the height of the rainy season streams in many places become rushing torrents of murky wa- ter that roll boulders along the stream beds; such streams are poor habitats for fragile tadpoles. For example, Hyla miotympaiuim calls throughout the dry season along a (juiet stream, 3 kilometers southwest of Huatusco, Veracruz, VIexico; tadpoles of this species are abundant in the stream. In June and July the stream, swollen by heavy rains, roars through the ravine. Few, if any, adults of Htjia miotympamim are found on vegetation along the stream. It is doubtful if tadpoles can survive in the stream. The positive correlation of breeding acti\- ity with the rainy season in many pond- breeders is understandable, because so many of these species utilize temporary ponds that are formed by the heavy rains. Furthermore, many lowland areas are suitable for am- phibian activity only during the rainy season. In the dry season the frogs are secreted in bromeliads, elephant-ear plants under sheaths of banana plants, or in other moisture-holding hiding places unknown to collectors. Little conclusive information is available on the duration of the breeding season. On the bases of apparent year-round activity and the presence of tadpoles in many stages of development at widely scattered times during the year, it is reasonable to assume that many of the montane stream-breeders, such as Plec- irohijla, Ptycljoliyla. and the Hyla rivularis. and uranochroa groups, have extended breed- ing seasons. Several of the species apparently breed throughout the year. On the other hand, prolonged breeding seasons are un- usual in lowland pond-breeding species and seemingly exist onl)- in a few species living in rain forest. Thus far, the discussion of breeding has been concerned with entire species. Further insights into some of the situations, and some new problems are apparent when populations of one species are examined. Unfortunately, only incomplete data, at best, are available. Although it is highly probable that the wide ranging montane species Hyla miotympamim breeds throughout the year, I have just men- tioned that at one locality breeding activity ceases at the height of the rainy season. Populations of this species at high elevations in Hidalgo were inacti\e in [anuary and Feb- ruary, whereas populations at lower elevations were breeding. In the Caribbean lowlands, recei\ing abundant rain throughout the year, in lower Central America Hyla chraccata ap- parently breeds year-round, but to the north, in southern Mexico, where a definite dry sea- son occurs, the species breeds only in the rainy season. These are only two examples of species having wide altitudinal or latitudinal distributions and exhibiting altitudinal or geo- graphic variation in breeding seasons. The available records for Triprion spafii- latus and Phrynoliyas vemilosa indicate that these species have breeding seasons from June through August and April through August, respectively. However, these are the accumu- lated records of many years from throughout the range of the species. Both species, especialh' Triprion, emerge for breeding onh' after torrential rains. Experience has shown that the frogs emerge and breed on the night following a heavy rain and then disappear again, in many instances not to reappear until the next year. Consequently, where the data indicate a breeding season of three months for the species, the breeding activit}' in a given population may be limited to a period of a few hours in any gi\en year. Another nearly unkno\vn aspect of the breeding biology of h\'lid populations con- cerns the reproductive cycles of individuals in the population. In many places it is possi- ble to hear frogs of a particular species calling every night for many consecutive weeks or even months, but are the same individuals calling throughout that period of time? Also, we can ask but cannot answer: does one indi- \idual breed more than once a year? Some hints to the answers are provided by data on Hyla pscudopuma and Agalychnis calli- dryas. At Tapanti, Costa Rica, Hyla pseudo- puma was breeding from early April until mid-May, when breeding activity ceased until August. These data suggest that there are at least two breeding seasons in this population, but we do not know if the same indix iduals breed in both seasons. Some gravid females of Agalychnis callidryas taken early in the breeding season contained ovulated eggs plus 1970 DUELLMAN: HYLID FROGS 657 another complement of ovarian eggs. Most indi\ iduals taken later in the season contained only one complement. These meager data suggest that individual females of A^ahjcJini.s cdUidnjas breed twice in a given breeding season. EGGS Eggs are known of only 45 species of hy- lid frogs li\ing in Middle America. The fol- lowing discussion is based on my own obser- \ations and the scant information available in the literature. The majority of Middle American hylids deposit their eggs in water. Of those groups for which eggs and /or tadpoles are known, we can be reasonably sure that .35 species deposit their eggs in ponds and 52 species lay their eggs in streams. The following species are known to deposit their eggs in masses, either free or attached to vegetation, in ponds: Acris crepitans, Pseudacris clarkii, PternoJujIa fodiem. Triprion petasatus, and the following species of Hyla — boons, hou- lengeri, elaeochroa, eitphorbiacea, eximia, lo- quax, microcephala, phlebodes, plicata, pseti- dopuma, regilla, rosenbcrgi. staufferi, and icalkeri. Two of the latter species ( boam and rosenbergi) deposit their eggs in shallow basins constructed by the males in mud or gra\el at the edges of rivers or sluggish streams. Thus, although the o\iposition sites are adjacent to flowing water, the eggs are actually deposited in still water. Five species are known to spread their eggs in a film on the surface of the water in ponds; these are HyJa rufitela, Phrynohijas venulosa, Smilisca baudinii, SmiJisca cijano- sticta, and SmiJisca phacota. Of the 52 Middle American hylids that are known, or suspected, to deposit their eggs in streams, eggs of only five species are known. Hijla arenicolor deposits small clumps of eggs in quiet pools, and HijIa cadaverina deposits single eggs in the same situations. Eggs of Hyla cohjmba were found under a rock in a stream ( Dunn, 1924 ) , and those of Hyla sumichrasti were found attached to a dead leaf between stones in a stream (Star- rett, 1960a). The eggs of Hyla miotympamim are attached to the lee sides of rocks or to vegetation in streams. Empty egg cases at- taclied to rocks in streams known to be in- haljitcd only by Plectrolujia are a good indi- cation that at least some of the species in that genus deposit their eggs on rocks in streams. I suspect that eggs are deposited in streams by all of the species of Plectrohylo and Pty- chohyJa and by the members of the following species groups of Hyla: bistiticta, erythrom- ma. miotympamim, mixomaculata, pictipes, pinonun, rivtdaris, salvadorensis, sumichrasti, taeniopus, and uranochroa groups. Some hylids deposit their eggs on \ege- tation above water. Presumably all of the Middle American species of Agahjchnis and PIn/Uomedusa (eggs not known for A. hto- dryas and P. venusta), and Pachymedusa dac- nicoJor attach clumps of eggs on leaves or branches of bushes or trees overhanging ponds. Hyla ebraccata and at least some of its relatives comprising the South American Hyla leucophyllata group usually deposit their eggs in a single layer on leaves of emer- gent herbs in ponds. Two species of Hyla (lancasteri and thorectes) are known to lay their eggs on vegetation overhanging moun- tain streams. Four species are known to deposit their eggs in water abo\e the ground. Eggs of Anotheca spinosa have been found in water- filled ca\ ities in trees and in bromeliads; Hyla bromeliacia, dendroscarta, zeteki, and prob- ably picadoi lay their eggs in bromeliads. It is highly likely that at least some of the fringe- limbed tree frogs of the Hijla miliaria group utilize tree holes for egg deposition. In Hemiphractus panamensis, eggs are car- ried in depressions on the back of the fe- male; there the eggs develop directly into small frogs. The eggs are carried in a dorsal brood pouch in Gastrothcca ceratophrys; pre- sumably these eggs also undergo direct de- velopment. The same condition exists in Gastrothcca nicefori. The numbers of eggs produced by indi- vidual females of various species seems to vary directly with differences in size of the species. This correlation holds true in groups of related species having the same kinds of life history, such as the Smilisca baudinii group (Duellman and Trueb, 1966). Stream- breeding species tend to have far fewer eggs than do pond breeders. E.xamples of the lat- 658 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 ter category include: Sniiliscu baudinii (2620- 3320, mean 2960 eggs), Smilisca cyanosticta (910), Smilisca phaeota (1665-2010, mean, 1848), and Triprion petasatm (1750). A va- riety of stream-breeders have fewer, but larger eggs: Hyla miotijmpamim (120), Hyla pic- tipes (126), Hyla suiniclirasti (50), Hyla uranochwa (69), Ptyclwhyla euthy.sanota (155), and Ptyclwhyla schinidtorum (191). Specialized methods of egg deposition seem to be correlated with a decrease in the num- ber of eggs. The species of Ai^alychnis all of which suspend their eggs on vegetation over ponds, have fewer eggs than do those species that deposit their eggs in ponds; for example, Agalychnis annae has 47 to 162 (mean, 102) eggs, and Agalychnis callidryas has 39 to 108 (mean, 78) eggs. The two species that de- posit their eggs on vegetation over streams produce very few, large eggs. Three clutches from Hyla tJwrectcs contained 10 eggs each, and two clutches from Hyla lancasteri con- tained 20 to 23 eggs. The same reduction ap- parently holds for bromeliad breeders; one seemingly complete clutch from Hyla brome- liacia contained 14 eggs and a gravid female of Hyla zeteki contained 24 eggs. Parental care by means of carrying eggs and young also results in fewer and larger eggs. One female of Gastrotheca ceratophrys contained nine eggs in the brood pouch. Numbers of ovarian eggs, egg scars on dorsum, or at- tached young in Hemiphractus panamensis vary from 12 to 14. Primitive hylids probably deposited their eggs in clumps in ponds. From this original type the other modes of deposition probably were evolved independently. Each of the secondary oviposition habits possibly evolved several times. The two groups of frogs in Middle America that deposit their eggs on vegetation over ponds ( phyllomedusines and Hyla leucophyllata group) are distantly re- lated and certainly evolved their oviposition habits independently. The same certainly is true for the surface-film deposition habit and for the bromeliad deposition habits of Ano- theca and the species of Hyla (probably in- dependently in two groups of Hyla). The stream habit apparently evolved twice in Mid- dle American hylids, and independently a third time in the South American Hyla colyrn- ba group. TADPOLES The morphological adaptations of tadpoles have been discussed in detail in a preceding section (Taxonomic Characters and Criteria in Hylid Frogs) . Only a brief summary of the ecology of tadpoles is presented here. Tad- poles are known for 83 species of Middle American frogs. These include all of the gen- era that have an aquatic larval stage and all of the species groups of Hyla, except the Hyla miliaria group. The following have tadpoles that develop in ponds: Paclnjmedtisa. A \ - UJ m 20 > \ - 10 \ A \ / \ V ,■■' Direct-^ ^><— Brc imeliod N^.^ •■A ■ A '• ~-\ • • »...-.-■ .--.--•--»' ■"• • • • • 1 1 1 1 1 1 1 \ 1 0 500 1000 1500 2000 2500 3000 ELEVATION IN METERS Fig. 319. Altitudinal distribution of liylid frogs in Middle America. 3500 A distinct correlation exists between alti- tudinal range and mode of larval develop- ment in hylids; 31 of the 34 species that reach their upper altitudinal limits at an elevation of no more than 1000 meters are pond breed- ers. Forty-nine species have their lower alti- tudinal limits at an elevation of no more than 500 meters; 39 of these are pond-breeders; only seven stream-breeders occur at eleva- tions below 500 meters (fig. 320). Sixty per cent of the 74 species that occur at elevations below 1000 meters are pond-breeders, whereas only 32 per cent of the 74 species are stream- breeders. The number of pond-breeders usu- ally is inversely proportional to the number of stream-breeders; this is not indicative of any kind of competition between the two groups but rather an expression of the avail- ability of breeding sites (fig. 319). The foregoing comments on altitudinal distribution pertain to the Middle American hylid fauna as a whole. Some wide-ranging 670 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 IT) UJ O UJ Q- O q: UJ CD 3 I'^l _ - UPPER LIMITS 10 • % 5 • PI 1 ^ 'T- § — g - p-| — — -^C^J!^ 1 ^ ^ . — . ^ — _ [ ; 0 500 1000 1500 2000 2500 3000 I 3500 meters 10 15 S^' 2 U. W^ \ 0 40 LOWER LIMITS □ Pond -breeders EH Stream- breeders ^ Bromeliad-breeders |#] Direct Development Fig. 320. Upper and lower altitudinal limits of hylid frogs in Middle America. .species exhibit different altitudinal distribu- tions at different latitudes. For example, Aiia- lychnis callidryas, Hyla chraccatci, Ihjhi lo- qitax, and Hyla microcephala occur only on the lowlands ( elevations less than 300 meters ) in southern Mexico, whereas in Panama and Costa Rica the altitudinal range of each of these species extends from the lowlands to elevations of more than 800 meters. Hyki ebmccata, loqtiax, and microceplwla occur at elevations up to 1200 meters. Similar variation in altitudinal distribution is exhibited by certain montane stream-breed- ing species that occur on both Atlantic and Pacific slopes. In nearly every species in which the altitudinal limits are well known, the altitudinal range is greater on the Atlantic than on the Pacific slopes. For example, in Oaxaca, Mexico, Ptycholiyhi Jconliardschuh- zei occurs at elevations from 700 to 1S50 me- ters on the Atlantic slopes but only from 900 to 1700 meters on the Pacific slopes. Partial altitudinal displacement occurs in IlyUi ttnino- chroa in the Cordillera Talamanca in Costa Rica and western Panama where the species has an altitudinal range of 600 to 1500 meters on the Atlantic slopes and 1400 to 1720 meters on the Pacific slopes. Variation in altitudinal distribution on different slopes is primarily due to altitudinal differences in climatic zones. This is especially noticeable in the case of cloud forest, which is more restricted alti- tudinally on the Pacific slopes than on the Atlantic slopes. Certain aspects of altitudinal and ecologi- cal distribution can be shown by transects across the Middle American highlands (figs. .321 and 322). In both of the transects that 1970 DUELLMAN: HYLID FROGS 671 CO m UJ 3000 o 2000 I UJ _i UJ 1000 - PACIFIC ■5 "li to 1^ -5 :. g-c ^ S a "^ s^'^ 5 S ^,. .... . •s I"" 1. 1 -5-5 ■♦J ^ 3 » 31 ■6 § '^ -5 :^ "•% (b «■■ ^^ 5) -9-5 .^ §Z^ > J L _L J L Fig. 323. Distributional patterns of hylid frogs in the lowlands of Middle America. The Atlantic lowlands are abo\e the political units, and the Pacific lowlands are below. Each line represents one species, unless noted otherwise by a number. TABLE 62 Distributional Limits of Hylid Frogs in the Lowlands of Middle America ( N=The Total Number of Species Present) Region N Atlantic Lowlands Tamaulipas 4 San Luis Potosi .. 4 Veracruz 10 Guatemala 9 Honduras 8 Nicaragua 14 Costa Rica 12 Panama 9 Pacific Lowlands Sonora 3 Sinaloa 6 Jalisco 7 Michoacan 7 Guerrero 7 Oaxaca 8 El Salvador 4 Nicaragua 5 Costa Rica 10 Panama 11 Northern Limit Southern Limit 2(50% 1(25% 6(60% 0(00% 0(00% 7(50% 1(08% 1(11% 2(67% 3(50% 1(14% 0(00% 1(14% 1(13% 0(00% 2(40% 5(50% 6(55% 1(25% 0(00% 1(10% 1(11% 1(12% 3(21% 4(33% 2(22% 0(00% 0(00% 0(00% 1(14% 0(00% 4(50% 1(25% 0(00% 2(20% 1(09% The humid lowlands of the Golfo Dulce region in southeastern Costa Rica form an environmental barrier to the distribution of some species that inhabit the subhumid Pa- cific lowlands. Smilisca baudinii reaches the southern limits of its distribution in the sub- humid lowlands of Costa Rica, whereas Htjh stauffeh is present on either side of the Golfo Dulce region but is absent from the humid lowlands in the Golfo Dulce region. On the other hand, seven species {Agahjchnis calli- clryas, Agahjchnis spurreUi, Hyla botdengeri, Hijla ebraccata, Hijla elaeochroa, Hijla rufi- tela, and SnuRsca phaeota) that inhabit the Caribbean lowlands of lower Central Amer- ica and not the subhumid Pacific lowlands occur in the humid Golfo Dulce region. The presence of these species in the Golfo Dulce region indicates that in the not too distant past a continuous humid forested environ- ment must have existed between the Golfo Dulce and the Caribbean lowlands. Perhaps the connection was via the Nicaraguan low- lands or maybe via the Arenal depression in the Cordillera de Tilaran in Costa Rica ( Hey- er, 1967). The interesting aspects about the three lowland connections between the Atlantic and 674 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Pacific lowlands concern the occurrence of species on one side and/ or the other of each isthmus (table 63). Eleven species occur on the lowlands of the Isthmus of Tehuantepec; only two of these are found on both Atlantic and Pacific sides. However, one species (Phrijnohyas venidosa) occurs on the Atlantic side and not on the Pacific side of the isthmus proper, but it does occur elsewhere on the Pacific lowlands. Thirteen species occur in the Nicaraguan isthmus; none is restricted to the Pacific lowlands, but two species ( Hijla microcephala and Hyla staufferi) occur no farther south on the Atlantic side, and Hyla microcephala occurs no farther north on the Pacific side. None of the 16 species occurring in the restricted central part of the Isthmus of Panama occurs on both coasts of the isth- mus proper. However, four species that are members of the Atlantic side fauna occur in Pacific foothills to the east or west of the isthmus. The elevations separating the Atlantic from the Pacific drainages in the isthmuses are lower than the highest elevations commonly reached by any of the species occurring in the isthmuses. Apparently no geographical barriers exist, but definite environmental dif- ferences are present between the Atlantic and Pacific lowlands in the three isthmuses. At each isthmus, the Pacific lowlands receive much less rainfall, have a longer dry season, and support less luxuriant vegetation than the Atlantic lowlands. The distributions of most of the hylid frogs in Panama and northwestern South America follow the cross-over pattern point- ed out by Dunn (1940a), in which species that occur on the Pacific lowlands of Central America are found on the Caribbean lowlands of South America and species that occur on the Caribbean lowlands of Central America are found on the Pacific lowlands of South America. The distributional patterns are TABLE 63 Distribution of HyHd Frogs in Three Middle American Isthmuses Tehuantepec Nicaragua Panama Pacific Only Hyla smithii Pachymeclusa dacnicolor Triprion spatulatus Hyla crepitans Hyla rosirata Hyla microcephala Hyla roscnbergi Hyla rubra Hyla staufferi Phrynohyas vcniilosa Smilisca sila Both Sides Hyla staufferi Smilisca haudinii Hyla microcephala Hyla staufferi Phrynohyas venidosa Smilisca haudinii Atlantic Only Agalychnis callidryas Hyla ehraccata Hyla lo(fuax Hyla microcephala Hyla picta Phrynohyas venidosa Agalychnis callidryas Agalychnis saltator Hyla houlcngeri Hyla ehraccata Hyla elaeochroa Hyla locpiax Hyla phlehodes Hyla rufitela Smilisca phaeota Agalychnis calcarifer Agalychnis callidryas Agalychnis spurredi Hyla houlcngeri Hyla ehraccata Hyla phlehodes Hyla rufitela Smilisca phaeota 1970 DUELLMAN: HYLID FROGS 675 slightly more complicated than intimated by Dunn. Sixteen specii's of hylids occur in tlic lowlands of both Panama and northwestern South America ( table 64 ) . Six of these show the usual cross-o\er pattern from Caribbean Central America to Pacific South America; seven others are on the Pacific lowlands of Central America and the Caribbean lowlands and/or the Amazon Basin of South America. Two species are principally Caribbean in Central and South America, and one occurs on the Pacific lowlands of both. The generalities of the patterns hold true for most of northwestern South America and western Panama. However, in eastern Pana- ma we find a broad zone of interdigitation and overlap of Caribbean and Pacific species. The near absence of mountain ranges of e\en moderate elevations and the presence of hu- mid forests and open forest-savanna associa- tions results in a mosaic of distributions not encountered elsewhere in lower Central America. As examples, we need only to ex- amine the lists of species known from the lower Rio Chucunaque, Darien. Four of the nine species are primarily Caribbean in Cen- tral America, whereas the other five are Pa- cific. Six of the thirteen species known from the Rio Tuira Basin are primarily Caribbean; the other se\en are typical of the Pacific low- lands in Central America. The present distribution patterns of the hylids in eastern Panama and adjacent Co- lombia seems to be primarily dependent on the distribution of en\ironments in the region. Some species probably exist in peripheral en- vironments and survive under suboptimal con- ditions. Some of these same species and some others exist in relict populations in isolated subhumid areas. The mixture of lowland spe- cies of hylids in eastern Panama extends west- ward only to the Canal Zone. To the west and thence northward into Costa Rica the Caribbean and Pacific faunas are separated by high mountains. Haffer (1967a and 1967b) concluded that the distribution of birds in northwestern Colombia and adjacent Panama also was principally governed by "forest" and "non-forest" habitats. The highlands of Middle America are di- vided by lowlands into three major mountain masses, from north to south: the Mexican highlands, the Chiapan-Guatemalan high- lands, and the Costa Rican-western Pana- manian highlands. The Mexican highlands are the most extensive and the highest. The great elevated area consists of the Mexican Plateau bordered on the east by the Sierra Madre Oriental, on the west by the Sierra Madre Occidental, and on the south by the Cordillera Volcanica. In addition the Sierra de Coalcoman and Sierra Madre del Sur parallel the Pacific coast in southwestern Mex- ico. The highlands of Nuclear Central America are comprised by the Sierra Madre extending from eastern Oaxaca to Honduras, the central highlands of Chiapas, the Sierra de los Cuchu- matanes in western Guatemala, and a vast complex of small east-west small mountain ranges extending from Guatemala to northern Honduras. The highlands of lower Central America are comprised principally by the Cordillera Talamanca in Costa Rica and western Panama and the eastern extension, the Sierra de Ta- basara in Panama. The Cordillera Central and Cordillera de Guanacaste in Costa Rica complete the highland complex. The sizes of the hylid faunas in these three highlands correlates with the size of the high- land areas. Thirty-two species occur in the extensive Mexican highlands, and 21 occur in the Guatemalan highlands, whereas only 15 are found in the Costa Rican highlands. The hylid faunas of the three highlands are highly distinctive; only fi\'e species are shared between the Mexican and Guatemalan high- lands, whereas one is shared between the Guatemalan and Costa Rican highlands. The genus Plectrohyla is endemic to the Guate- malan highlands, whereas the genus Ptyclio- hijla is shared with the Mexican highlands. The Hijla salvadoren.sis group has one species in the Guatemalan highlands and one in the Costa Rican highlands. Except for a few species that occur only on the Caribbean or Pacific slopes, the hylid fauna is rather evenly distributed throughout the highlands of Costa Rica and western Panama. There are notable differences in the hylid faunas on the Atlantic and Pacific slopes of the Guatemalan highlands; only four spe- cies occur on both slopes. The highlands of 676 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 CD W pa < v ^ o hJ M-i o c ;-( « O 1 s < -C o St: bO « P « ■-4-. ™ o ii lu 3 -£2 <^ •o o o O 3 -cS(J ^ CO pa go, N -go- So XXX X X X X X X X X X X X X XX ; X X X X X X X X X X X X X X X X X X X X X X X :2 s 2 ^ tjO tii ^ (^ M •t •- CO -a 3 C to a r^ -S S p s 2^ «S J3 , <- "3 ~ c? i 2 -2 -2 .5 .S -2 ^^ '^^ S5 S5 21 :i) 'Bi Si a, Si's! "Si i: "S "H 1970 DUELLMAN: HYLID FROGS 677 most of Honduras are too poorly known to comment on the distribution of hylids throughout those ranges. Of the 32 species of h\lids in the Mexican highlands, three wide-ranging species occur in all of the four major Cordilleras, of which the Sierra Madre Occidental and Cordillera Volcanica each have four species of hylids. Fourteen species occur in the Sierra Madre del Sur; of these, six are endemic, and six are among the 23 species (14 endemic) in the Sierra Madre Oriental. Five isolated small highland areas must be mentioned. The Sierra de los Tuxtlas in southern \'eracruz has a rich hylid fauna composed mostly of species occurring either in the surrounding lowlands or on the slopes of the Sierra Madre Oriental. The Azuero highlands in the southern part of the Azuero Peninsula, Panama, are practically devoid of hylids. Only Smilisca sila is a member of the depauperate fauna there. The Serrania dc Darien, Serrania de Pirre, and Cerro Sapo are three separate and nearly parallel moun- tain ranges in eastern Panama. Gastrotheca nicefori occurs on Cerro Pirre and Cerro Sapo, plus several localities in Colombia; it is the only hylid species restricted to these ranges in Central America. The numerous continental islands along both coasts of Middle America are nearly de- void of hylid frogs. No hylids occur on the islands in the Golfo de California, Islas Tres Marias, Isla Coiba, Archipielago las Perlas, or Archipielago de San Bias. Two species occur on Isla Cozumel, three on Isla Grande del Maiz, one on Isla Cebaco, and four on the islands comprising the Archipielago de Bocas del Toro. EVOLUTION OF THE MIDDLE AMERICAN HYLID FAUNA An attempt at a synthesis of the taxonomic and distributional data is a fascinating chal- lenge. The absence of a fossil record of hylids or any group of \'ertebrates in Middle Amer- ica makes it necessary to rely on the charac- ters and distributions of the living species in combination with the evidence of the geologi- cal and climatic history of Middle America in order to draw zoogeographic conclusions. The historical zoogeography of Central America has been adequately discussed by Savage (1966) and Stuart (1966), and various regional studies in Mexico have contributed equally substantial information (Duellman, 1960b, 1965c; Savage, 1960). Consequently, I will not elaborate on the geological history of Middle America and instead refer my read- ers to the above papers and the many publi- cations cited therein. Previous workers deaHng with the history of the Middle American fauna religiously fol- lowed the Matthewsian concepts of a northern origin and southward dispersal of the families of amphibians and reptiles. Dunn (1931c) considered Central American groups that had relatives in South America to be members of his Old Northern Fauna. Schmidt (1943) called the same groups hanging relicts. Stuart ( 1950 ) recognized these Central American groups that had differentiated from their South American relatives as the Autochtho- nous Middle American Fauna. Savage (1966) recognized these groups as the Mesoamerican Fauna, a zoogeographical element equal in its distinctness to the Nearctic and Neotropical faunas. Dunn and Schmidt were devoted disciples of Matthews; Stuart and Savage, although they recognized the distinctive nature of the Middle American fauna, formulated their ideas at a time when the permanence of the present continental land masses was held sa- cred. However, in recent years the earlier unpopular concept of continental drift has received tremendous support from the geolo- gists who have amassed a wealth of data on paleomagnetic crustal movements, midoceanic ridges, and intercontinental continuity of beds (see Wilson, 1963; Takeuchi, Uyeda, and Kanamori, 1967; and Hurley, 1968). Sudden- ly Gondwanaland is in vogue. It seems that the question is not if, but when, continental drift occurred. Excellent evidence is avail- able in support of continental connections in the Southern Hemisphere in the Paleozoic, and the early Mesozoic, and suggestive bits of evidence are present for the Jurassic and pos- sibly early Cretaceous. Some substantial bio- logical evidence supports the concept of Hol- antarctic distributions (Darlington, 1965; Brundin, 1965; and Hallam, 1967). The recent work of Brundin ( 1967 ) on chironomid 678 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 midges presents a masterful synthesis of a Holantarctic center of origin and dispersal. The recent find of a Triassic labyrinthodont in Antarctica (Barrett, Baillie, and Colbert, 196S) is the most significant vertebrate evi- dence demanding the consideration of Ant- arctica in the paleogeography of vertebrates. Coin and Coin recently suggested Antarctica as the center of origin of frogs.'*' The time is ripe to re-examine the zoo- geographic concepts that have been applied to the Middle American herpetofauna; we must now inquire about the probabilities of a northern origin and a southward dispersal versus a southern origin and a northward dis- persal. We should not make the mistake that has been so prevalent in the past of assuming that the entire fauna has been derived from one direction. Instead, each group must be examined independently. What I have to say about the origin and the dispersal of the hylid fauna may or may not be applicable to other groups of organisms; that is something for other workers familiar with other groups to determine. I recognize four principal hylid faunas in the world: 1) Neotropical — the largest and most diverse. 2) Australo-Papuan — rather large in numbers of species and moderately diverse. 3) Mesoamerican — moderately large and very diverse. 4) Holarctic — small and lacking diversity. The large number of species and incredible diversity of the South Ameri- can hylids is a strong indication that the group differentiated and dispersed from that region. Only the Neotropical fauna contains all four subfamilies of hylids. The Mesoamerican hylid fauna is distinc- tive but obviously related to the Neotropical fauna by virtue of the presence of three Meso- american autochthonous genera belonging to two subfamilies that do not occur in the Hol- arctic fauna. I can find no evidence to ally the Holarctic fauna with the Mesoamerican fauna. Furthermore, due to my lack of knowl- edge of the few Old World members of the Holarctic fauna, I have no suggestions con- '" This suggestion was made in a paper entitled "Antarctica as the center of origin of frogs" presented at the annual meeting of the American Society of Ichthyologists and Herpetologists in Ann Arbor, Mich- igan, in June, 1968. ccrning the possible relationships of the Hol- arctic hylids with those in the Australo- Papuan region. The Holarctic hylids seem to be separate from the others. This separa- tion is more than just spatial. Perhaps the Holarctic hylids are relatively recent forms that have yet to develop a variety of char- acteristics. Conversely they might be archaic forms, but due to the broad distributions of many species and the near absence of diver- sity the fauna probably does not possess great antiquity. My concept of tlic origin of the Middle American hylid fauna centers on two in\a- sions from Southern America and one minor invasion from North America ( fig. 324 ) . These are summarized below. 1. An invasion from South America, prob- ably in the Cretaceous, of at least three stocks of hylids representing bylines, amphignatho- dontines, and phyllomedusines. This group evolved in isolation through most of the Ceno- zoic into the diverse Mesoamerican hylid fauna. 2. A second invasion from South America after the reformation of a land connection with Central America in the Pliocene. All species are members of the Neotropical hylid fauna. 3. A dispersal of two species groups of the Nearctic component of the Holarctic h\'- lid fauna from North America into the high- lands of northern Middle America. This prob- ably did not occur before mid-Pliocene. Thus, in Middle America we ha\c three historical groups — the Mesoamerican and rep- resentatives of the Neotropical and Nearctic hylid faunas. The following discussions of the evolution of these faunas contain, I fear, as much conjecture as fact, but nothing like the tree frog witnessed by Giles ( Woodhouse, 1966). The Mesoameric.\n Hylids The largest historical element in the Mid- dle American hylid fauna is the Mesoameri- can element, which comprises 73 per cent of the hylid fauna in Middle America. In this element we have all of those phyletic lines that cvohed in Middle America while South America was isolated by seaways during most of the Cenozoic. Included in the Meso- 1970 DUELLMAN: HYLID FROGS 679 Fig. 324. Proposed phylogenetic history of the major hylid faunas in Middle America. Numbers in cones are the numbers of species. I=NucIear Central American and Mexican Highland Component. II=Lo\ver Cen- tral American Highland Component. HIn^Lowland Component. anierican hylids are three components, which I refer to as: 1) the Mesoamerican Lowland Component, 2) the Lower Central American Highland Component (the Talamancan Her- petofauna of Savage, 1966), and 3) the Nu- clear Central American and Mexican High- land Component (in part the Guatemalan Highland Herpetofauna of Savage, 1966). The second and third have been deri\'ed from the first, perhaps entirely independently but possibly through an intermediate form that has passed into oblivion. The all too meager evidence suggests that a basic Mesoamerican hylid fauna was composed of one phyllome- dusine stock, one amphignathodontine stock and one or two hyline stocks. The two Meso- american highland components are quite dis- tinct in their evolutionary histories and are discussed separately. The stream-adapted tadpoles of the spe- cies in the two highland components are spe- cialized derivatives of the pond-type of tad- pole characteristic of the lowland component. Certain structural features of the adults also suggest that the montane species are derived from lowland types. The conclusions based on morphology are supported by the physio- logical work by Brattstrom (1968, p. 110), who stated: "High altitude forms [anurans] in the recent mountains of Central America are physiologically essentially lowland tropi- cal forms that have been carried or forced into a variety of restrictive physiological plas- ticities." 680 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Mesoamefican Lowland Component: Prior to the separation of Central America from South America in the Eocene certain groups of hyhds inhabited what is now Cen- tral America; each of these groups had rela- tives in South America. Two of these groups ( phyllomedusines and amphignathodontines ) can be dispensed with readily. The primitive phyllomedusine stock in Mid- dle America probably was a generalized phyl- lomedusine, perhaps not much difFerent from Pachijmedusa dacnicolor. Certainly the Meso- american stock and the Neotropical stock which developed into PhijUotnediisa had a walking gait and arboreal eggs. Probably a Pachijmedusa stock was isolated on the Pa- cific slopes of Mexico by the early Miocene, by which time uplift of the Mexican and Nu- clear Central American highlands created in- creased diversity of environments. The Pachij- medusa stock evolved towards increasing aridity, whereas the Agalijchnis stock re- mained in humid forests and differentiated into six species in Middle America. The primi- tive Agahjchnis stock probably was much like Agahjchnis saltator, which evolved in the Cen- tral America peninsula south of the Nica- raguan Embayment, whereas its close ally, callidnjas apparently developed north of the embayment. The evolution of annae and moreletii apparently are correlated with the elevation of the Talamanca range in lower Central America and the Nuclear Central American highlands, respectively. Agahjchnis spurreUi is a specialized lowland species which probably e\olved in the humid forests of lower Central America in the late Tertiary and spread into Chocoan South America and in eastern Panama differentiated into lito- drijas. The Agahjchnis calcarifier-craspedoptis stock migrated into South America after the Pliocene continental connection. East of the Andes the stock differentiated into craspedo- pus, whereas on the Pacific lowlands it evolved into calcarifer, which subsequently extended its range northward into Central America. The Amphignathodontinae, as rather loosely defined in the present work, is repre- sented in Middle America by one stock that was i.solated there throughout the Cenozoic. This stock had certain amphignathodontine morphological features, but lacked the brood pouch, which was developed in females in the Neotropical representatives. Thus, the only Mesoamerican amphignathodontine, Anotheca, remained relatively primitive in its reproductive modifications. Perhaps all primi- tive amphignathodontines were like Anothe- ca and deposited their eggs in bromeliads and/ or water-filled cavities in trees. The remaining members of the Mesoameri- can lowland component are bylines currently recognized in three genera plus four species groups of Hijh. It is reasonable to assume one ancestral stock for all of these, except the enigmatic Hijla miliaria group comprising the so-called fringe-limbed tree frogs. The ab- sence of data on life histories, mating calls, and cranial osteology for the five species placed in this group precludes any meaning- ful phylogenetic conclusions. I am uncertain about the five species being placed in one group. The northernmost species, vaJancifer and echinata, seem to be more closely related than either is to the three southern species. Whatever their relationships might be with one another, the relationships with other hy- lids, either Neotropical or Mesoamerican, are even more obscure. On the basis of our pres- ent knowledge of these frogs it is not possible to determine if they originated from the com- mon Mesoamerican stock or are representa- tives of a separate stock that was isolated in Middle America. Returning now to the main hyline stock in Middle America, we haxe a generalized lowland pond-breeder with unspecialized tad- poles having 2/3 tooth rows. I assume that this stock had a generalized skull (quadrato- jugal, pterygoid-prootic articulation, and squamosal-crista parotica articulation pres- ent), teeth on the prevomers, and a fronto- parietal fontanelle. Probably the members of the Hijhi godmani group are most like this early stock. At the present time this group occurs on the Atlantic lowlands and foothills, godmani to the northwest of the Isthmus of Tehuantepec and loc/uax in Central America. The Ilijhi picta group seems to be closely re- lated to the godmani group and to have dif- ferentiated by reduction in size, reduction of cranial elements, and modification of colora- tion. The two species in the picta group in- 1970 DUELLMAN: HYLID FROGS 681 habit peripheral subhumid lowlands in north- ern Middle America. Hyla smitliii occurs on the Pacific lowlands of Mexico, and Hyla picta inhabits the Atlantic lowlands of Mex- ico southward to northern Honduras. Their present ranges are narrowly separated by an apparent barrier — the xeric Plains of Tchuan- tepec. Structuralh' the adults of the Hyla bromeliacia group are hke those of the picta group. The only major differences are in life history. The members of the bromeliacia group deposit their eggs in bromeliads and have tadpoles with long muscular tails and ventral mouths containing two upper and four or five lower rows of teeth. This group, containing bromeliacia in northern Central America and dendroscarta in southeastern Mexico, apparently diverged from the low- land pond-breeding picta-stock by adapting to arboreal breeding habits in a successful at- tempt to in\-ade the foothills and low moun- tains, where ponds are scarce. An early derivative from the basic Meso- american hylid stock is represented now by the frogs of the genus Smilisca, which accord- ing to Duellman and Trueb (1966) evolved in the niesic tropics of the southeastern part of the Central American paleopeninsula prob- ably in the early Miocene, but possibly earlier. The Smifoca-stock difl^erentiated into two groups — the baudinii group on the Caribbean lowlands and the sordida group on the Pacific slopes of lower Central America. Probably before the elevation of the Talamancan Range in Costa Rica and western Panama the sor- dida-stock invaded the Caribbean of Costa Rica. One species, puma, evolved on the Caribbean lowlands, whereas two others, sor- dida and sila on the Caribbean and Pacific slopes, respectively, adapted to life in streams as the Talamanca Range uplifted in the Plio- cene. The baudinii group of Smilisca re- mained in the lowlands of Middle America and differentiated into two species (cyano- sticta and phaeota) in the humid en\'iron- ments, whereas baudinii became widely dis- tributed in the subhumid lowlands. Increasing aridity in the Pliocene and Pleistocene were met with some striking adap- tations for survival in arid environments. Ac- cording to Trueb (1970a), an apparent early divergent stock from the Smilisca progenitor developed a casque head characterized by broad labial flanges and a prenasal bone. This stock e\'olved into Triprion, one species in the Yucatan Peninsula {peicusatua) and anoth- er on the Pacific lowlands of Mexico (spatu- latus). Trueb (1970a) showed a progression of cranial dermal proliferation from Smilisca baudinii to Pfernohyla dentata and finally Pternohyla fodiem, thereby demonstrating the highly probable course of evolution of Pternohyla from a Smilisca battdinii-hke an- cestor. Pternohyla dentata occurs in the Rio Santiago Rasin on the Mexican Plateau, and P. fodiens, which inhabits the lowlands of western Mexico, extends northward into Arizona, the northernmost occurrence of any Mesoamerican hylid. Lower Central American Highland Component: The basic Mesoamerican hyline stock pro\ided a progenitor to the hylids of the mountains of lower Central America. This fauna consists of 12 species in five groups of Hyla. The primitive generalized Hyla in the lower Central American highlands is repre- sented by the present-day members of the Hyla pseudopuma group. This group contains two species (angustilineata and pseudopuma) having generalized skulls and tadpoles that develop in montane ponds. The two species differ principally in coloration and mating call. All other species of Hyla in this com- ponent seem to have e\'ol\'ed from a pseudo- puma-like ancestor in response to montane habitats lacking ponds for breeding. With the exception of one group of bromeliad-breeders, all of these derived species are stream-breed- ers and differ from one another principally in the kinds and degrees of stream adaptations of the tadpoles and progressive reduction of the skulls of the adults. The buccal adapta- tions of the tadpoles include enlargement of the mouth or the development of a funnel- shaped mouth but no increase in the number of tooth rows; tadpoles of all members of the lower Central American highland Com- ponent have two upper and three lower rows of teeth, except the arboreal tadpoles of the zeteki group, which lack definitive rows of teeth. Although in external appearance the adults of the monotypic Hyla lancasteri group differ strikingly from the members of the Hyla 682 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 pseudopuma group, these differences are su- perficial. The only significant cranial modifi- cations in lanca.steri are the shortening of the snout region and reduction of the nasals. The tadpoles are moderately elongate and have onh' slightly enlarged mouths. The lancasieri group occurs at moderate elevations on the Caribbean slopes; some populations are ap- parently unique in the Lower Central Ameri- can Highland Component by depositing their eggs on vegetation over streams. The frogs in the llyla uranochroa group {uranoclnoa and riifioculis) have only slightly reduced skulls but have extremely modified stream-tadpoles, which have long muscular tails with shallow fins and funnel-shaped mouths. The frogs in this group have red eyes, a unique character in the Lower Central American Highland Component; they could have evolved from either a pseiidopuma-\ikc ancestor or an early lancasteri-\ike stock. Both the adults and tadpoles of uranochroa and ntfioculis are very much alike structurally; the two species probably differentiated as a result of altitudinal separation, although they now occur sympatrically at intermediate elevations on both Pacific and Caribbean slopes. A separate adaptive line includes the stream-breeding Hyla pictipes and rivularis groups. These montane stream inhabitants are characterized by reduced cranial ossifica- tion (loss or reduction of quadratojugal, no bony articulation of the scjuamosal and crista parotica, and no bony connection of the me- dial ramus of the pterygoid with the prootic) and highly modified tadpoles ha\ing greatly enlarged ventral mouths and long muscular tails. These groups obviously are descended from a stock having a more fully developed skull and ha\ing more generalized tadpoles; thus, they probably e\olved from an ancestor much like Hyla pseudopuma. Certainly, the pictipes and rivtdaris groups represent an en- tirely separate ph\letic line from that whicli gave rise to the urarwchroa group. On the basis of its somewhat more generalized .skull (shorter sphenethmoid and broader fronto- parietals) the monotypic Uijla pictipes group seems to be an early divergent line from the stock that gave rise to the rivularis group. Hyla pictipes is now restricted to high ele\ a- tions in the Cordillera Central and Cordillera Talamanca. The four species in the Hyla rivularis group are very similar in the struc- ture of the adults and tadpoles. Hyla iica has the shortest sphenethmoid and most gen- eralized mating call. The sphenethmoid is progressi\ely longer in rivularis, xanthosticta, and debilis. The four species are partially segregated altitudinally, and no member of the group is completely geographically allo- patric to all other members of the group. The four species probably differentiated through geographic and altitudinal isolation in the constantly changing Cordilleras in the late Tertiary and subsequently established their present partially sympatric distributions. Aside from the generalized Hyla psetido- piima group, the only members of the Lower Central American Highland Component that are not stream-breeders are the members of the Hyla zetcki group. The two species (ze- teki and picadoi) in this group adapted to the montane forests by developing the habit of depositing their eggs in bromeliads. Be- cause of this divergent reproductive behavior and only moderately reduced cranial ele- ments, the progenitor of the zeteki group probabh' was a generalized pond-breeder, possibly much like the members of the pseu- dopuma group. The two species in the zeteki group are broadly sympatric, but picadoi oc- curs at higher elevations than zeteki; their differentiation may ha\e been the result of either altitudinal isolation or geographic sepa- ration in the Cordillera Central and Cordillera Talamanca with subsequent migration of each species into the other cordillera. It is necessar\- to note here that the brome- iiad breeding habit apparently e\olved inde- pendently in the Hyla zeteki and bromeliacia groups. The tadpoles of the former group have anterodorsal mouths and reduced tooth ro\\'s, whereas those of the bromeliacia group ha\(> \entral mouths and no reduction of tootli rows. The Nuclear Central American and .Mexican Highland Component: In the high- lands of northern Central America and in Mexico there exists a hylid element containing 40 species currently recognized in nine spe- cies groups of Hyla and the genera Plectro- liyla and Ptychohyla. One additional species occurs in lower Central America. All of these 1970 DUELLMAN: HYLID FROGS 683 groups are stream-breeders. Many members of this component have tadpoles with only slightly enlarged mouths and the basic tooth row formula of 2/3. In those tadpoles ha\ing enlarged mouths the number of tooth rows is \ariously increased to as man\' as 7/11. This is in marked contrast to the members of the Lower Central American Highland Com- ponent, in which even in those species having greatly enlarged mouths the tooth row for- mula is always 2/3. One species in the foothills of lower Cen- tral America, Htjla legleri is closely related to the northern Hyla saloadorensis; by virtue of both ha\ing a tooth row formula of 2/. 5 these species are placed in the Nuclear Cen- tral American Component. We can assume only one other transgression of the Nicara- guan lowland gap by a member of a highland assemblage. The Guatemalan tadpole de- scribed under Species Inquirienda obviously belongs with the Lower Central American Highland Component. Like the tadpoles of the Hyla pictipes and rivularis groups, the Guatemalan tadpole has an immense ventral mouth with 2/3 tooth rows. The combination in the same component of frogs in the highlands of Nuclear Central America with those in the highlands of Mex- ico is contrary to the faunal dissimilarities of the two highland areas presented in recent summaries of the herpetofauna ( Duellman, 1960b, and 1966c; Savage, 1966). Ne\-erthe- less, the Mesoamerican hylids in the two highland areas separated by the narrow low- lands of the Isthmus of Tehuantepec definite- ly seem to be members of one faunal element. Four species occur in both areas; members of two other species groups are found in both areas, and two closely related groups are separated geographically by the isthmus. A realistic phylogenetic history of the northern Mesoamerican highland hylids can be con- structed only by taking into account the spe- cies and groups in both highland areas. Cer- tainly the frogs in the two areas did not have entirely separate evolutionary histories. The origin of Central American and Mexi- can Highland Component is obscured in the absence of any seemingly primitive type that could be intermediate between the highland component and the Mesoamerican Lowland Component. This position is filled by the Hyla pseudopuma group in Lower Central American Highland Component. Considera- tion should be given to the one known group that spans the Nicaraguan gap and occurs in both highland areas, namely the Hyla salva- dorcn-sis group. The adults in this group are sufficiently generalized that they might be relatively unchanged from a progenitor of the northern highland component. However, their tadpoles are more specialized than many of those in other groups in this component, although the tadpoles in the salvadorensis group could ha\'e become modified after the differentiation of other groups. We can arrive at a basic cranial type if we consider the presence of a quadratojugal, the bony articulation of the squamosal with the crista parotica, and the bony connection of the medial ramus of the pterygoid to the prootic as generalized and primitive cranial conditions. Furthermore, tadpoles that have relatively small anteroventral mouths with two upper and three lower tooth rows and rather deep caudal fins arc obviously the least specialized of the stream tadpoles and are thus considered to be the generalized stream tadpoles in this component. Therefore, it seems an easy task to find the primitixe group in this northern highland component; we need only to find a group having the generalized cranial and larval characters. But no such group exists, probably because the e\olution of cranial features is entirely separate from the evolution of larval characters. Conse- r[uently, we have no extant group that can be considered as an idealized progenitor of the Nuclear Central American and Mexican High- land Component. Nevertheless, the frogs in this component seem to belong to a single historical group; furthermore, certain evolu- tionary trends and phyletic lines are evident within the group. The five most important evolutionary trends are correlated with increased adapta- tion for life in and along montane streams. These trends are: 1) reduction of certain cranial elements, especially the loss of the cjuadratojugals ( this reduction is not confined to frogs that are adapted to the stream habi- tat), 2) reduction and loss of vocal sacs and voice, 3) increase size of hands and length- 684 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 ening of digits, apparently as an adaptation for grasping rocks in streams, 4) depression of the body, lengthening of the tail, and re- duction of the caudal fins in tadpoles, and 5) enlargement of the mouth to form a ven- tral oral sucker and a corresponding increase in the number of tooth rows. No one group exists that has e\ol\cd all of the above char- acters to their most advanced state. Several instances of parallelism are evident, such as the loss of \'oice in distantly related groups and the loss of the quadratojugals in two sepa- rate phyletic lines. For the sake of simplicity the frogs in this highland component can be divided into three subcomponents. Each subcomponent repre- sents a major phyletic line which is charac- terized by a combination of traits or trends not present in the other subcomponents. The first of these contains the taeniopus, salva- dorer^is, enjthromma, and pinonim groups, a total of eight species. Most of these frogs have a well-developed quadratojugal. The Hi/la taeniopus group, comprising three spe- cies (clianeque, taeniopus, and altipoteits) , is probably one of the most primitive groups in the subcomponent. The pterygoid is in bony contact with the prootic in all three species, and the squamosal articulates with the crista parotica in clianeque and taeniopus. A voice is present in some populations of chaneqiw; small vocal slits are present in taeniopus but absent in altipotens. The tadpoles have long muscular tails and small mouths with 2/3 {altipotens and taeniopus) or 2/4 (chaneque) tooth TOWS. Hi/Ia altipotens and taeniopus ha\e greatly enlarged testes; both species ap- parently evolved from a chaneque-Mke ances- tor. The differentiation of these species seems to have been the result of geographic isola- tion; thus, chaneque evolved in the Guate- malan highlands, taeniopus in the Sierra Madre Oriental, and altipotens in the Sierra Madre del Sur. This differentiation must have occurred prior to the Wisconsin, the most recent time when cloud forest might have existed on the low ridges of the Isthmus of Tehuantepec and thereby allowed Hyla chaneque to cross into the Mexican highlands. The Hyh salvadoremis group consists of salvadorensis and legleri; both have general- ized skulls and tadpoles with 2/5 tooth rows. Probably the ancestral stock of the salva- dorensis group extended along the Pacific slopes of the moderately uplifted highlands from El Salvador to Costa Rica in the Plio- cene; subsequently two populations were iso- lated by the intervening Nicaraguan lowlands in which subhumid conditions developed. The northern population evoKed into salvadoren- sis, and the southern population became leg- leri. Probably in the Pliocene a stock of small stream breeding hylids that was derived from the salvadorensis stock occurred on the slopes of the highlands of southern Mexico; this stock subsequently differentiated into the erythrom- ma and pinorum groups — the former in the Sierra Madre Oriental, and the latter in the Sierra Madre del Sur. Both groups are char- acterized by the loss of the pterygoid-prootic articulation and by a reduction of the quad- ratojugal. The tadpoles of the monotypic erythromma group developed 4/6 tooth rows and dispersed around the edge of the Mexi- can highlands; thus, it came to occur sym- patrically with members of the pinorum group in the Sierra Madre del Sur. Prior to the Pleistocene the pinorum group stock invaded the Chiapan highlands to the east of the Isth- mus of Tehuantepec and differentiated into Hyla melanomma, while the stock in the Si- erra Madre del Sur evolved into Hyla pinor- um; both species retained the 2/5 tooth for- mula of the ancestral salvadorensis group. No later than Wisconsin time, melanomma in- vaded the Sierra Madre del Sur, where it oc- curs sympatrically with pinorum. Perhaps melanomma and pinorum differentiated in different areas in the Sierra Madre del Sur, and subsequent to their differentiation mela- nomma crossed the Isthmus of Tehuantepec to the Chiapan highlands. The second subcomponent contains four groups of Hyla and the genus Ptychohyla, a total of 15 species. All of these frogs lack a quadratojugal and a bony articulation of the pterygoid with the prootic. Only in the most primitive Hyla miotympanum group does the squamosal ha\e a bony articulation with the crista parotica. The differentiation in this group e\idently was correlated with, or oc- curred subsequent to, the first major uplift of the highlands in the Miocene. The Hyla 1970 DUELLMAN: HYLID FROGS 685 miotympanum group consists of two species (iiiiotympanum and arborescandcns); the for- mer occurs at lower ele\ations in the Sierra Madre Oriental and has tadpoles with 2/3 tooth rows, whereas the latter lives at higher ele\ations in the same mountains and has tad- poles with 2/4 tooth rows. The species prob- ably differentiated at different ele\'ations; the tadpoles of arhorescandens with their longer tails and larger mouths having more tooth rows reflect adaptation to the more swift streams typical of higher elevations. The two species in the Hyla hazelae group {hazelae and thorectes) retained the generalized stream tadpoles of the miotympanum group, but lost the bony articulation between the squamosal and crista parotica. The hazelae group probably is a relatively recent diver- gent line from the miotympanum group. Pos- sibly the members of the hazelae group are relicts of a former more widespread miotym- panum group that were isolated in separate highland areas due to increasing aridity in post-Wisconsin time. The Hyla mixomaculata and sumichrasti groups are the most specialized members of the second subcomponent. Both have reduced cranial elements; tadpoles of the former group have 7/11 tooth rows, and tadpoles of the sumichrasti group have 3/7 tooth rows. Mem- bers of the mixomaculata group apparently lack a voice. Present distributional evidence suggests that the mixomaculata group orig- inated in the Sierra Madre Oriental and that the sumichrasti group originated on the Pa- cific slopes of Me.xico. Each group probably evolved independently from a generalized an- cestral stock, possibly the progenitor of the miotympanum group. A mixomaculataAike stock apparently spread southward into the Sierra Madre del Sur and there gave rise to pellita, whereas the population that remained in the Sierra Madre Oriental evolved into mixomaculata. Hyla mixe and nuhicola seem to be closely related deri\atives of the mixo- maculataAike stock. They presumably arose as isolates in the Sierra Madre Oriental, per- haps during climatic fluctuation in the Pleisto- cene. The last group in the second subcompo- nent is the genus Ptychohyla, an assemblage of five species differing from all other Middle American hylids by ha\ing large xentrolateral glands in the breeding males. The genus con- tains two species groups differing in larval, adult, and ethological characters. Duellman (1963c) suggested that Ptychohyla was re- lated to the Hyla uranochroa group in lower Central America. Now that the stream hy- lids of Middle America are better known such an arrangement does not seem to be so plausi- ble, although tadpoles with funnel-shaped mouths occur in Ptychohyla and in the Hyla uranochroa group. Perhaps Ptychohyla evolved from a generalized Hyla miotympan- umAike stock in the Guatemalan highlands. There the stock differentiated into two groups, probably by means of selection for lar\al differences. This differentiation must ha\e occurred by mid-Pliocene, after which time the continued uplift of the Guatemalan highlands separated the Ptychohyla eiithy- sanota stock into euthysanota on the Pacific slopes and a spinipoUex-leonhardschultzei stock on the Atlantic slopes. The latter stock crossed the Isthmus of Tehuantepec in a glacial period of the Pleistocene and differ- entiated into leonhardschultzei in the Mexican highlands, whereas the residual stock in Gua- temala developed into spinipoUex. The schmidtorum group also crossed the isthmus and developed into ignicolor in the Mexican highlands; at the same time the Guatemalan population e\olved into schmidtorum. The third subcomponent contains the Hyla bistincta group (nine species) and Plectro- lujla (10 species). These two groups exhibit parallel progressive adaptations to the moun- tain stream habitat; on the basis of their similar morphology, they must be closely re- lated. In both groups the skulls are well ossi- fied, but the quadratojugal is greatly reduced or absent. Species in both groups have blunt heads and large hands with long fingers. Al- though the tadpoles have long muscular tails they have only slightly enlarged ventral mouths with two upper and three lower tooth rows. The advanced species in both groups lack a voice. Probably the ancestral stock to these two groups was widespread in the moderately uplifted highlands of southern Mexico and Guatemala in the Miocene. Sub- sequently, great uplift of the highlands in the Pliocene resulted in the isolation of the an- 686 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 cestral Plectrohyla in the Chiapan-Guatemal- an highlands and the Hijla histincta stock in the Mexican highlands. The bistincta group is nicely di\ided into three subgroups — two generalized species (histincta and pentJieter) having vocal slits and unspecialized hands, two related diver- gent species (cliryses and charadricola) , and five related allopatric specialized species ( roh- ertsorum, pachyderma, siopela, crassa, and hogertae) . The primitive member of the group, Hyla histincta, is widespread in the Mexican highlands. The closely related Hyla pentlieter probably evolved in the Sierra Madre del Sur from the widespread histincta stock, which also differentiated into a more specialized form in the high mountains of the Sierra Madre Oriental. The latter stock probably was continuously distributed through those mountains in plu\'ial periods in the Pleistocene, but now is represented only by relict populations that have differen- tiated sufficiently to be considered as five allo- patric species. From north to south these species are robertsorum, pachyderma, siopela, crassa, and hogertae. Hyla chnjses and char- adricola also probably represent relicts of a former widespread derivative of the Hyla histincta-Mke stock, but their relationships with bistincta and pentheter are not clear. Concommitantly with the diversification of the Hyla histincta group in the Mexican highlands, Plectrohyla was differentiating in the Chiapan-Guatemalan highlands. Conceiv- ably, in the course of the uplift in the Plio- cene the Plectrohyla stock was separated into a highland component and another compo- nent at moderate elevations on the slopes. The latter component retained vocal slits and evolved into a group of small species, whereas the highland component evolved into a group of larger species lacking vocal sHts. The former group, which for conve- nience can be called the sagoriim group, even- tually established populations on the Atlantic and Pacific slopes of the highlands. Possibly, the species now known as quecchi was the original inhabitant on the Atlantic slopes, whereas matudai was endemic to the Pacific slopes. Through isolation, differences in voice, shape of the snout, and mouthparts of the tad- poles developed. Subsecjucnt climatic fluctu- ation, probably in the Pleistocene, permitted migration southward of the f/t/ecc/i /-stock and northward of the matudai-stock. Depression of climatic zones and uplift through volcanism again resulted in isolation of populations on Atlantic and Pacific slopes, but this time two species were present on each slope. The matudai-stock on the Atlantic slope differen- tiated into /.v;7, and the f/uecc/n'-stock on the Pacific slope evolved into sagoriim. The relationships of the species in the highland component (guatemalensis group) are more obscure. Plectrohyla glandulosa and pycnochila seem to be the least specialized species; the condition of the prcpollical pro- cess in these .species probably is relati\ely un- changed from that of the Plectrohyla proto- type and is much like that in the Hyla bistinc- ta group. It is possible that glandidosa de- veloped in the Sierra de los Cuchumantanes in Guatemala, while pycnochila was isolated in the highlands of central Chiapas. Appar- ently Plectrohyla avia represents an evolu- tionary intermediary between the generalized glandulosa-pycnochila stock and guatemalen- sis and harticegi. The prcpollical spine is long and pointed in avia (independently e\ol\ed in the sagorum group) and is bifid in guate- malensis and harticegi. Plectrohyla avia is endemic to moderately high elevations on the Pacific slopes; harticegi occurs at the same elevations but farther west. Plectrohyla gua- temalensis occurs nearly throughout the geo- graphical (but not the altitudinal) range of the genus. It occurs sympatrically with avia and possibly with harticegi. The Neotropical Hylids All four subfamilies of hylid frogs have their greatest di\ersity in South America. With the exception of the Hylinae, only two genera (Anotheca and Pachymedusa) are en- tirely extra-Neotropical. As can be expected in any large fauna such as the Neotropical hylids, there are many diverse types of morphological, developmental, and behavioral adaptations. Some of the adaptations that are characteristically Neotropical and not present in Mesoamerican groups are summarized be- low. 1970 DUELLMAN: HYLID FROGS 687 1. Ph\lIoniedusincs ha\ing grasping feet (PhyUomcdusa). 2. Triangular dermal helmet (hemiphrac- tines ) . 3. Aniphignathodontines carrying eggs on the back or in a dorsal pouch { Amphipmiho- don, Cnjptohatrachus, Flectonottis, Frilziana, Gastrotheca, Nyctimantis, and Stcfania). 4. Paired lateral \ocal sacs behind the angles of the jaws {Osteoceplwhis, Phnjno- hijas, Argenteohyla, and Trachycephahis) . 5. Odontoids on mandible, palatine, and parasphenoid (Phyllodytes). 6. Single, median, subgular vocal sac formed by longitudinal dermal folds on throat ( Sphaenorhychiis ) . 7. Hyla ha\'ing angular prevomerine den- tigerous processes, such as are characteristic of members of the albomarginata, boons, geo- graphica, and lanciformis groups. 8. Hyla having projecting snouts and re- duced webbing between the first and second toes (rubra group). 9. Hyla having pelagic tadpoles with 2/4 tooth rows {albomarginata and boons groups ) . 10. Hyla having tadpoles with xiphicercal tails and terminal mouths lacking teeth, such as are characteristic of members of the leuco- phyllata, microcephala, and parviceps groups. 11. Haploid number of 15 chromosomes (leucophyllata. microcephala, and parviceps groups); the same number occurs in the Papuan Hijla angiana. The above features, taken together or in- di\'idually, characterize hylid frogs that un- derwent their diversification in South Amer- ica. Much, if not all, of this diversification probably occurred during the period of the Cenozoic ( Paleocene-Pliocene ) when South America was isolated from Central America by seaways. Subsequent to the connection of Central America with South America in the late Plio- cene, members of several different phyletic lines of the diverse Neotropical hylid fauna invaded Central America. It is possible that some of these immigrants arrived somewhat earher by means of island-hopping through the archipelago existing in the Cenozoic Pana- manian Portal. However, hylids seem to be notoriously poor at this means of dispersal. as witnessed by the poverty of the West In- dian species of hylids in comparison with the exceedingly rich Eleiitherodactylus fauna there. It is not necessary to assume immigra- tion of the Neotropical hyHds into Central America prior to the continental connection, because the distribution and minor differentia- tion of this fauna in Central America can be explained adequately on the basis of the later arrival. The Neotropical hylid fauna in Central America consists of 22 species. Four of these species occur only in lower Central America and at present are not known from South America, although in each case a closely re- lated, and possibly conspecific, taxon is known in South America. Phyllomediisa lemur in Costa Rica and Panama is closely related to P. buckleyi and medinae in South America. Phyllomedusa veniisto, which is known from only one locality in eastern Panama, is re- markably similar to P. edentula in Amazonian South America. Hemiphractus panameiisis is very much like the Ecuadorian H. fasciatus; acquisition of material from Colombia should show that both nominal species belong in the same taxon. The relationships of the Pana- manian Gastrotheca cerotophrys are with the Chocoan G. cornutum, which might be con- specific. Most of the other species are more widely distributed in Middle America (table 6.5). Species in four groups have undergone differ- entiation in Middle America. Hyla ebraccata, the only Middle American species in the large Amazonian Hyla leucophyllata group, barely enters South America in Pacific Colombia. Hyla rufitela is a Middle American endemic and northernmost member of the South Amer- ican Hyla albomarginata group. Five of the 25 species of the Neotropical Hyla rubra group occur in Middle America; two of these species (elaeochroa, staufferi) apparently dif- ferentiated in Central America and are re- stricted to Middle America. The Neotropical Hyla microcephala group is composed of about a dozen species, four of which occur in Middle America. Evidently the Middle America members of this group differentiated from a single stock. Two of the resulting species (robertmertensi and sartori) are re- stricted to Middle America; phlebodes barely MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 TABLE 65 Distribution of Species of Neotropical Hylicl Groups in Middle America Species °3 a a CS CO P-M Ji' S .^ o Phijllomednsa lemur .._ PhijUomediisa vemista Hemiphractus patuimensis Gastrotheca ceratophnjs _.-. Gastrotheca nicefori Phrtjnohijas venulosa — Hyla rubra Hyla elacochroa Hyla staufferi Hijla boulengeri Hyla rostrata Htjla microcephala Hyla robertmertensi Hyla phlebodes Hyla sartori Hyla ebraccata Hyla subocularis Hyla rafitela Hyla crepitans Hyla rosenbergi Hyla boans Hyla colymba ? ? ? X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X " ? indicates the presence of closely related, possibly conspecific species in South America. enters South America, and microcephala is widespread in South and Middle America. The relationships of the montane, stream- breeding Hyla colymba are with three Andean stream-breeding species comprising the Hyla bogotensis group. Apparently Hyla colymba immigrated into Central America from South America after the closure of the Panamanian portal in the late Pliocene. The Neotropical hylid fauna in Central America apparently became established there after 16 separate phyletic lines entered Cen- tral America from South America. The evolu- tion of these separate phyletic lines took place in South America prior to the immigration of members of these groups into Central Amer- ica. If my earlier suppositions about the rela- tionships of the Central American Phyllome- dusa, Hemiphractus, and Gastrotheca are cor- rect, differentiation in Middle America has taken place in only five of the 16 groups. The immigrations are summarized below: 1. Phyllomedusa buckleyi-medinae-lemur series from Amazonian South America to foot- hills of lower Central America. 2. Phyllomedusa edentida-venusta from Amazonian South America into Panama. .3. Hemiphractus fasciatus - panamensis probably from Amazonian South America in- to Chocoan region and Panama. 4. Gastrotheca cormitum-ceratophrys from Chocoan South America into Panama. 5. Gastrotheca nicefori from the Andes to mountains of eastern Panama. 6. Phrynohyas venulosa from non-forested lowlands east of the Andes into lowlands of Middle America. 7. Hyla rubra group {boulengeri-\ike stock) from Amazonian lowlands into Cen- tral America; subsequent differentiation of 1970 DUELLMAN: HYLID FROGS 689 rostrata and migration of rostrata into non- forested lowlands of northern South America. 8. Hijla rubra group {ruhra-elaeochroa- staufferi stock) from Amazonian South Amer- ica into Central America; subsequent differ- entiation of elaeochroa and staiifferi from rubra. 9. Hyla tnicrocephaluAike stock from Am- azonian South America; subsequent differen- tiation of phlebodes, sartori, and robertmer- tensi from microcephaJa, of which phlebodes extended its range into Chocoan South Amer- ica. 10. Hijla leucophijUataAike stock from Amazonian South America; subsequent dif- ferentiation of Middle American populations into ebraccata, which extended its range into Chocoan Colombia. 11. Hyla subocidaris from Amazonian South America into eastern Panama. 12. Hyla crepitans from non-forested areas of northern South America into eastern Pana- ma and subsequently to northern Honduras. 13. Hyla rosenbergi from Chocoan South America into lower Central America, or from a boans-\ike stock from Amazonian South America with subsequent differentiation into rosenbergi in lower Central America followed by migration into Chocoan South America. 14. Hyla boons from Amazonian South America into eastern Panama. 15. Hyla albomarginata-MVe stock from Amazonian South America into lower Central America; subsequent differentiation of Central American populations into rufitela. 16. Hyla colymba from Andean foothills into mountains of lower Central America. The temporal sequence of these invasions is shrouded by our lack of knowledge of the climatic history of the isthmian link. But even so, some temporal arrangements seem to be rather obvious. A relatively early invasion can be postulated for those groups that have undergone differentiation in Central America and/ or have migrated into northern Middle America. Those species that have not differ- entiated from South American populations and have restricted ranges in lower Central America could have entered Central America at a later time. I conceive of five temporal invasions; these are not thought to be com- pletely distinct "faunal waves" but rather as the approximate temporal sequence of in- vasion. Probably the earliest members of the Neo- tropical hylid fauna to enter Middle America were the Hyla microcephala, rubra, and bou- lengeri stocks, all of which subsequently dif- ferentiated into several species in Middle America. Perhaps at about the same time, Phrynohyas vemdosa entered Middle Amer- ica. Some members of all these groups in- habit the subhumid lowlands of Middle Amer- ica, all of these groups have dispersed north- ward into Mexico. I consider that this first invasion of Neotropical frogs took place soon after the closure of the Panamanian Portal in the late Pliocene. If any of the Neotropical hylids reached Central America by island- hopping prior to the closure of the portal, they certainly must have been members of these groups. The Hyla albomarginata and leucophyllata groups and Hyla crepitans could ha\e entered Central America somewhat later, possibly at the end of the Pliocene or in early Pleistocene time. The first two groups differentiated from their parental Neotropical stocks and dis- persed through humid lowland forests of low- er Central America; both reached the Golfo Dulce region of the Pacific lowlands. Hyla ebraccata (the Middle American derivative of the leucophyllata group) subsequently ex- tended its range to southern Mexico and northwestern South America. The Middle American Hyla crepitans is undifferentiated from the populations in northern South Amer- ica, but it has existed in Central America for a sufficient length of time to extend its range from the subhumid savannas of Panama to the subhumid lowlands of northern Honduras without leaving any relict populations in the intervening lowlands now covered mostly with humid forest. This feat also was accom- plished by Cnemidophorus lenmiscatus. A third group of Neotropical species ap- parently entered Central America during a Pleistocene glacial period when temperatures were depressed and probably some lowland areas had more rainfall than they do now. All of the species in this group (Phyllome- dusa lemur, Hemiphractus panamensis. Gas- trotheca ceratophrys, G. nicefori, and Hyla colymba) live in humid foothill forests, and 690 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 none has greatly differentiated in Central America. Two species {PlnjUomedusa lemur and Hyhi colymba) extend to Costa Rica; the others are restricted to Panama and Colombia. PlnjUomedusa vetmsta, Hijla boans, and H. subocidaris barely enter Central America in eastern Panama; each species is a member of a widespread group in Amazonian South America. They are the most recent immi- grants. In Middle America the Neotropical hylids are principally lowland in their distribution. Neotropical species comprise 70 per cent of the 16 species of hylids in the Canal Zone, but only 40 per cent of the 13 species on the Caribbean lowlands of Costa Rica and only 37 per cent of the eight species in the low- lands of southern Veracruz, Mexico. Five of the Neotropical hylids live in foothills or low mountains in Central America. Three of these are only in Panama; two also occur in Costa Rica, and none is a part of the rich highland hylid fauna in Nuclear Central America and Mexico. The relative paucity of Neotropical hylids in the lowlands of northern Middle America and their absence in most of the highland regions can be explained on the bases of time, lack of adaptations to environmental conditions, and competitive factors. Some of the earlier invaders, such as Hyki staufferi and Phrynohtjas vemdosa, which arc adapted to subhumid environments, have migrated northward to the northern limits of the tropics in Mexico. Subhumid conditions seem to be a controlling factor to the dispersal of some lowland species, such as Hyla ebraccata, clae- ochroa, and nifitela, although elaeochroa and rufitela do not seem to have reached their potential northern limits of distribution, pos- sibly due to lack of time. There are no ob- vious ecological or physical barriers at the presently known limits of distribution of Gas- trotheca ceratopJirys and Hemiphractus pana- mensis in Central America. Again, perhaps only more time is required for them to extend their ranges along the foothills of the Cordil- lera Talamanca and Cordillera Central of Costa Rica, unless, of course, they already exist there and have not been found by the many collectors who have swarmed over Costa Rica in the last decade. The extensive Middle American highlands are devoid of Neotropical hylids save Hi/la colymba, the only Neotropical stream-breeder in Central America. The absence of suitable breeding sites for the pond-breeding Neo- tropical hylids in the mountains of Middle America is an important limiting factor to those species. I can find no e\'idenee that any of the many groups of montane stream-breed- ing hylids in Middle America descended from a Neotropical stock. This absence of stream descendants from Neotropical lines is strik- ing in comparison with the multitudinous stream-inhabitants that seemingly descended from Mesoamerican lowland pond-breeders. Although lack of time and presence of com- petitors may be of some importance, I think that the absence of evolutionary potential in the Neotropical pond-breeders precludes their diversification into montane habitats in Cen- tral America. With the exception of the mem- bers of the Ilyki nd)ra, and albomarginata groups, all of the Neotropical lowland pond- breeders in Middle America have either spe- cialized breeding behavior for ponds {boans group ) or specialized pelagic tadpoles ( leuco- phyUata, microceplmla, and parviceps groups). In fact, the tadpoles of none of the Neotropical groups is sufficiently generalized to adapt to stream conditions. Competition may be an important factor in the distribution and relative abundance of Neotropical versus Mesoamerican species in the Middle American lowlands, especiallv the extensive subhumid areas, characterized by prolonged dry seasons. Adaptations by Meso- american hylids for survival under these se- vere environmental conditions include sur- face-film eggs (Smilisca), integumentary- cranial co-ossification (Pternohyla and Tri- prion), and rapid development of tadpoles (all three genera mentioned). Among the Neotropical species in Middle America, only Phrynohyas vemdosa has corresponding adap- tations (surface-film eggs, rapid tadpole de- velopment, and thick glandular .skin). Even though the Neotropical species com- prise only 19 per cent of the total Middle American hylid fauna, these groups form a significant part of the fauna in lower Central America. Some Neotropical species have spread throughout the lowlands of Middle 1970 DUELLMAN: HYLID FROGS 691 America, but the Neotropical hylids ha\e had only moderate success in the highlands of lower Central America and arc absent from the highlands north of Costa Rica. The Nearctic Hylids In comparison with the Neotropical and Mesoamerican elements, the Nearctic hylid fauna is characterized by a paucity of species and little di\crsit)'. In the present systematic arrangement 26 species are grouped in four genera, the largest of which is Hijla with 16 species in four groups. Pseudacris (seven species) and Limnaoechis (one species) are weakly differentiated from Hijla. However, Chantell (1968) suggested that Limnaoechis might be more closely related to Acris. The two species in the latter genus are notably dis- tinct from other Nearctic hylids. Although there are fragmentary fossil re- mains from various parts of North America from the Lower Miocene through the Pleisto- cene (see Auffenberg, 1956; Chantell, 1964; Holman, 19.59, 1961, 1962, 1963, "1966" 1 1968], 1967; Lynch, 1964, 1965b, 1966c; and Tihen, 1960), none of these fossils contiubutes sig- nificantly in unraveling the systematic and zoogeographic relationships of the Nearctic hylids, neither among the groups recognized in North America nor with the Mesoamerican hylids. A possible minor exception is the Upper Miocene-Lower Pliocene Pseudacris nordensis from Nebraska, Chantell (1964) suggested that this species might be inter- mediate between Uijla and Pseudacris, al- though he found material referable to Pseuda- cris clarkii in the same fauna. No workers have successfully related Ne- arctic species to members of the Mesoameri- can hylid fauna. Blair ("1958" [1959] and 1960) placed Sniilisca haudinii in the Hyla versicolor group and Hijla staufferi in the Hyla eximia group. These erroneous group- ings were based solely on similarities of the mating calls of haudinii, staufferi, and Nearc- tic species without consideration of the mating calls of the Mesoamerican relatives of hau- dinii and staufferi: furthermore, morphologi- cal characters were not considered. Several authors have suggested that frogs in the Hyla eximia group are closely related to the Hyla arhorea complex in Eurasia; the most recent statement is by Taylor (1962, p. 346): "The arhorea group of Hyla also occurs in America. A species group in Mexico (including eu- phorhiacea, cardenasi, eximia, arhoricola, la- frentzi, and wrightorum) must be regarded as members of the arhorea group. Some pop- ulations of arhorea are so similar to lafrentzi that they can be separated only with con- siderable difficulty, if at all." The relationships of the Nearctic hylids presimiabh' are with the Palearctic species. Anthony Gaudin is currently investigating the osteological characters of the Holarctic hylids. When his work is completed, and the results are compared with osteological data on the Mesoamerican hylids, a convincing argument might be put forth for the distant relation- ships of the Nearctic and Mesoamerican hy- lids. On the basis of the present evidence I can only assume such a relationship. Acris and Pseudacris barely enter northern Mexico and would not be included in an ac- count of Middle American hylids were it not for the fact that the Mexican-United States boundary is the arbitrary northern limit for this study. The Nearctic species of Hyla can be placed in four groups. The monotypic Hyla crucifer group and the Hyla cinerea group {cinerea and gratiosa) are confined to eastern North America. The Hyla versicolor group (areni- color, versicolor, chrysoscelis, avivoca, and femoralis) is widespread east of the Sierra Nevada in the United States. Hyla arenicolor is the westernmost member of the Hyla versi- color group; it dispersed southward on the Mexican Plateau probably in plu\ial periods in the late Pleistocene and post-Wisconsin. The only other Nearctic group in Middle America is the Hyla eximia group, represent- ed by the wide-ranging, variable Hyla regilla in western North America, probably Hyla squirella in southeastern North America, pos- sibly Hyla andcrsonii in eastern United States, and fi\e species (cadaverina, plicata, eximia, eupliorhiacea, and icalkeri) that inhabit Mex- ico. I agree with Jameson, Mackey, and Rich- mond (1966) that the eximia group (their Hijla regilla stock) was more widespread in pluvial (glacial) periods of the Pleistocene. However, those authors were working under the erroneous assumption that plicata (their 692 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 lafrentzi) and the northern populations of eximia (their icrightontm) were conspecific with regilla. Moreover, they did not consider the two southern species {euphorbiacea and tcalkeri ) . The dispersal and subsequent differentia- tion of the eximia group in western North America and in Mexico is correlated with the Madro-Tertiary Geoflora (Peabody and Sav- age, 1958). The historical components of southwestern North America include a Ma- drean Complex of the Young Northern Ele- ment (Savage, 1960). The Hyla eximia group, in Mexico at least, is part of the Madrean Complex. Apparently an early eximia group- stock was present in the Mexican highlands in the Pliocene. Uplift of the Cordillera Vol- canica in the Pliocene probably tended to isolate montane populations of a former more widespread stock; these montane isolates are known today as Ihjla plicata. This same up- lift also isolated populations to the north on the Mexican Plateau and to the southeast in the highlands of Oaxaca. The populations on the Mexican Plateau were subjected to con- siderable climatic fluctuations in the Pleisto- cene. At glacial or pluvial times the frogs dispersed over the plateaus and extended northward into Arizona and New Mexico, whereas during interglacial times their ranges were constricted to higher, more mesic areas. A variety of minor morphological, color, and ethological differentiation took place in the populations, which were alternatively isolated and confluent. The result of this history is the mosaic of varieties of Hyla eximia. The southeastern Mexican populations of the eximia group differentiated from the northern populations, and dispersed through the elevated region of Oaxaca and across the Isthmus of Tehuantepec into the Chiapan highlands. The dispersal across the isthmus must have occurred in late Pliocene or during an early Pleistocene glacial period. Subse- quent differentiation on either side of the isthmus resulted in the evolution of euphor- biacea in the Oaxacan highlands and tcalkeri in the Chiapan-Guatcmalan highlands. Hyla cadaverina apparently is an early divergent line from the eximia-regilla stock and became adapted for existence in sub- humid areas with the onset of increasing aridity in the Pliocene, whereas the regilla stock remained in more mesic montane en- vironments. The dispersal of regilla south- ward into southern Baja California probably occurred in a Pleistocene pluvial period. Sub- sequent isolation resulted in minor differen- tiation of the southern population into Hyla regilla ctirfa. The West Indian Hylids There is no evidence that any Middle American hylids were derived from the de- pauperate West Indian hylid fauna, but it is possible that some of the West Indian hy- lids were derived from Middle America. Dunn (1926) considered the four Jamaican species to have resulted from a single invasion of that island from Hispaniola, which also contains four species. The only other true West Indian hylid is Hyla septentrionalis on Cuba, Isle of Pines, the Bahamas, and south- ern peninsular Florida. Discounting the con- tinental islands of Trinidad and Tobago, the only other Hyla on a West Indian Island is the South American Hyla rubra on St. Lucia. Thus, we can view the West Indian hylid fauna as being comprised of nine endemic species — four on Hispaniola, four on Jamaica, and one centered on Cuba and the Bahamas. On evidence provided by a study of the cranial osteology, Trueb (1970a) concluded that a Hyla septentrionalis group containing septentrionalis, dominicensis, vasta, brunnea, and lichenata possibly evolved from a Hyla ])oans-]ike progenitor that waifed to the West Indies from South America. She considered that two phyletic lines are evident in the group. One of these contains as the primitixe form Hyla vasta on Hispaniola; domincensis on Hispaniola and septentrionalis on Cuba are treated as derived species. Trueb placed the Jamaican Hyla brunnea and lichenata in a second phyletic line in the septentrionalis group and concluded that they probably orig- inated /)! situ from a common ancestor that migrated from Hispaniola. Dunn's (1926) supposition that all of the Hispaniolan hylids are closely related can be disproved. Certainly Hyla heilprini with its green peritoneum, external pigmentation, and projecting prepollex is strongly suggestive of a South American Hyla albomarginata group 1970 DUELLMAN: HYLID FROGS 693 progenitor, despite Noble's (1927) contention tli;it Iwilpiini is a montane derivative of vasta. The relationships of the small Ilyla pulchri- lincata are not known. CertainK- it represents a separate stock from heilprini and the septen- trionalis group. The two small Jamaican species, Hijla niari(2nac and uilderi, were placed by Dunn ( 1926) with the larger species on the island — brunnea and lichenata. The bromeliad breed- ing behavior and similar adaptive types of tadpoles in all four species were his principal criteria for placing all of the species in one group. Dunn concluded that the speciation in the Jamaican hylids was the result of "fratricidal competition" in the tadpoles, which resulted in the metamorphosis of frogs at greatly varying sizes. Granting that Dunn's conclusions represent one solution to the problem of the Jamaican hylids, I question the validity of his argument and suggest that new evidence be sought. Trueb's suggested relationships of the Jamaican Hijla brunnea and lichenata with the septentrionalis group are based on the supposition that the ancestral stock that reached Jamaica was a casque- headed form. The development of a casque head is specialized. Casque-headed hylids are considered to be at the ends of various phyletic lines and not to be ancestors of more generalized forms. However, Dunn's theory of the paedomorphic status of uilderi and marianae offers an intriguing possibility that might be substantiated by developmental studies of the Jamaican species. Although HijUi uilderi and marianae have highly specialized arboreal tadpoles, it is con- ceivable that they evolved the larval charac- teristics independently of brunnea and lich- enata. I find no apparent close relationship of wilderi and marianae with Hispaniolan species and suggest the possibility that these two species might be derivatives of a gen- eralized Mesoamerican hylid stock. The two Jamaican species do not possess any morpho- logical characters that rule out this possibility. The foregoing comments on West Indian hylids are not intended to be conclusive but rather, I hope, inducive to stimulate research on this group of hylids. E.xcept for Trueb's (1970a) comments on the cranial osteology of some of the species, no new information has come forth after Dunn's ( 1926 ) work on the Jamaican species. SUMMARY AND CONCLUSIONS One hundred and fifteen species of liylid frogs arc known from Middle America (Mex- ica and Central America). On the bases of morphological characters of the adults and tadpoles, and features of their life histories, these species are placed in 15 genera. Si.x of these genera arc endemic to Middle America, and two others have their greatest diversity in Middle America. The 73 species of Hyh in Middle America are arranged into 28 groups, 18 of which are restricted to Middle America. The following taxonomic changes are pro- posed in this paper: 1) Hyla arboricola Tay- lor, 1941=Hijla eximia Baird, 1854. 2) Hijla bocourti Mocquard, 1899=Hy/a euphorhia- cea Gimthcr, 1859. 3) Ihjla cardenasi Taylor, 1939=Wy/rt eximia Baird, 1854. 4) HijJa dar- lingi Smith, Smith, and Werler, 1952=H(//« miotympamim Cope, 1863. 5) Hyla regilla deserticola Jameson, Mackey, and Richmond, 1966=Hyla regilla hypochondriaca Hallowell, 1854. 6J Hyla duellmani Lynch and Smith, 1966=:H ylachaneque Duellman, 1961. 7) Hyla immema Taylor, 1952=Hyla miliaria (Cope, 1886). 8) Hyla lythrodes Savage, ]968=Hyla rufioculis Taylor, 1952. 9) Hyla phantasmargoria Dunn, l943=Hyla miliaria (Cope, 1886). 10) Hyla pugnax Schmidt, l857=Hyla crepitans Wied, 1824. 11) Hyla richardtaylori Taylor, l954=Hyla fimbrimem- bra Taylor, 1948. 12) Hyla wrightorum Tay- lor, 1939=W(//« eximia Baird, 1854. 13) Ce- rathyla Jimenez de la Espada, lS7l=Hemi- phractm Wagler, 1828. No new taxa are pro- posed. The present study represents the first at- tempt to work out the systcmatics of a large, diverse group of frogs by utilizing characters such as cranial osteology, mating calls, and larval morphology, in addition to the conven- tional external morpliological characters of the adults. The utilization of a wide spectrum of characters has provided a wealth of evi- dence concerning the relationships of the species. A \ariet\' of modes of life history is ex- hibited by the Middle American hylids. The evolution of stream adaptations in tadpoles apparently has occurred at least twice in Middle America. Probably the habit of de- positing eggs in bromeliads has evolved inde- pendently in three groups. Although there is considerable continuit\- in the hylid fauna of the lowlands, the rela- tively depauperate fauna on the Pacific low- lands is distinct from that on the Caribbean lowlands. Significant faunal breaks occur at the Isthmus of Tehuantepec and the Nica- raguan Depression. The hylids in the three major highland areas are quite distinct; the highest percentage of endemism occurs in the highlands of Costa Rica and Western Panama. No species is shared between these highlands and those in Nuclear Central America, which has fi\e species in common with the Mexican highlands. The hylid fauna of Middle America con- tains three historical elements. The major element is the Mesoamerican fauna, which evolved in tropical Middle America from early South American stocks that were isolated in Middle America during most of the Cenozoic. A significant part of the present Middle Amer- ican hylid fauna is composed of species be- longing to the Neotropical fauna. These are late Cenozoic immigrants into Central Amer- ica. A third, relatively insignificant group is the Nearctic fauna, a part of the Holarctic hylid fauna that reaches its southern limits of distribution in the New World in northern Middle America. The lengthy presentation of my researches on Middle American hylid frogs answers many questions and raises several others. The relationships of some species are unknown. Although I have been tempted to in\oke the doctrine of special creation, I have followed the precedent established by Lucretius (58 B.C. ) : "Nothing from nothing ever yet was born." 694 APPENDIX 1 All of the specimens of hylids from Middle America that have been examined during the course of this study are listed be- low. The species are arranged alphabetically within the genera, which in turn are in alpha- betical order. Localities and specimens are given in the following order: country (ar- ranged from north to south — Mexico, British Honduras, Guatemala, El SaKador, Honduras, Nicaragua, Costa f-lica, Panama); states (de- partments, provinces) in alphabetical order in each country; localities in alphabetical order in each state; museum abbre\'iations arc given in alphabetical order as listed in Materials and Methods, and the number of specimens in each museum collection are given in pa- rentheses. Unless otherwise indicated, speci- mens are preserved frogs. Skeletons, lots of tadpoles, and clutches of eggs are so indi- cated. No distinction is made between cleared and stained specimens and those that are dried skeletons. For example, K.U. ( 16, 2 skeletons, 1 tadpoles) denotes that from a given locality there are in the collections at the University of Kansas, 16 preserved frogs, two skeletons, and one lot of tadpoles. Lo- calities that ha\'e not been located to state or equivalent political unit are listed immedi- ately after the name of the country. Speci- mens with data giving only the country or state are listed first in that political unit under "No specific locality." Acris crepitans blanchardi MEXICO: Coahuila: 19 kilometers north of Jimenez, 19 kilometers west of Jimenez, K.U. (10); 1.6 kilometers west of Jimenez, K.U. (1); 3.2 kilo- meters west of Jimenez, K.U. (10); Rio Sabinas, near Miisquiz, F.M.N.H. (11). Agalychnis annae COSTA RICA: Alajuela: Cinchona, K.U. (2 tad- poles). Cartago: Cartago, A.N.S.P. (4), F.M.N.H. (13), K.U. (73, 1 skeleton); Moravia, K.U. (4, 1 tadpoles, 1 eggs), U.S.C. (1); 2 kilometers south of Paraiso, U.S.C. (3); Tapanti, K.U. (39, 5 skeletons, 10 tadpoles, 4 eggs), M.C.Z. (2), M.V.Z. (3), U.I.M.N.H. (1), U.S.C. (1). Quanacaxte: El Silen- cio, La Laguna, U.S.C. (3). Limon: Jimenez, A.M.N.H. (1). San Jose: Guadalupe, K.U. (1); U.S.C. (1); La Hondura, A.N.S.P. (1); La Palma, K.U. (1, 7 tadpoles, 2 eggs), M.C.Z. (1), U.S.C. (1); San Jose, A.M.N.H. (3), A.N.S.P. (16), K.U. (2), M.C.Z. (2), S.U. (1), U.M.M.Z. (9), U.S.C. (12); San Pedro, A.M.N.H. (3), U.S.C. (12); Santo Domingo, M.V.Z. (4). Agalychnis calcarifcr COSTA RICA: HerecUa: Finca La Selva, U.S.C. (1). Limon: Siquirres, N.H.R.M. ( 1 ). PANAMA: Canal Zone: Barro Colorado Island, A.N.S.P. (1), F.M.N.H. (1), M.C.Z. (1). Darien: Laguna, K.Lf. (3, 1 eggs). Agalychnis callidryas MEXICO: Campeche: 7.5 kilometers west of Escarcega, K.U. (5); Matamoros, F.M.N.H. (1); Pacaitun, F.M.N.H. (1); Tuxpeiia, U.M.M.Z. (1). Oaxaca: 3 kilometers north of Donaji. U.M.M.Z. (10); 22 kilometers south of Jesiis Carranza (Vera- cruz), U.I.M.N.H. (6); 3.7 kilometers nortli of Sara- bia, U.M.M.Z. (8); 3 kilometers south of Tolocita, K.U. (6); Tuxtepec, K.U. (1. 1 tadpoles); 1 kilo- meter south of Ubero, U.M.M.Z. (27); 1 kilometer north of Valle Nacional, U.I.M.N.H. (10); 1.6 kilo- meters south of Valle Nacional, K.U. (23, 4 skeletons, 1 tadpoles), U.I.M.N.H. (18). Tabasco: 10 kilo- meters south of Cardenas, K.U. (1); La Venta, U.S.N. M. (2); Santo Tomas, U.S.N.M. (1); Teapa, U.M.M.Z. (9, 2 tadpoles); 10 kilometers north of Teapa, U.M.M.Z. (2); 13 kilometers north of Teapa, U.M.M.Z. (2); 21 kilometers north of Teapa, U.M.M.Z. (1). Veracruz: 7 kilometers south of Acayucan, U.I.M.N.H. (1); 33 kilometers south of Acayucan, U.I.M.N.H. (2); 1.6 kilometers east soutli- east of Alvarado, U.M.M.Z. (.5); 2.4 kilometers east- .southeast of Alvarado, U.M.M.Z. (2); 4.5 kilometers south of Aquilera, U.M.M.Z. (1); Berta, U.S.N.M. (1); 10 kilometers south of Catemaco, U.M.M.Z. (1); 8 kilometers southwest of Coatzacoalos, U.M.M.Z. (9); Cuatotolapam, U.M.M.Z. (33); Encinal, LT.M.M.Z. (20); 10 kilometers soutlieast Hueyapan, U.M.M.Z. (5); 21.6 kilometers south of Las Choapas, T.C.VV.C. (1); 24.8 kilometers soudi of Las Choapas, T.C.W.C. (2); 3.5 kilometers west of Lerdo de Tejada, U.M.M.Z. (4); 2 kilometers south of Naranja, K.U. (1 skeleton), U.M.M.Z. (17); Rodriguez Clara, U.I.M.N.H. (1); San Andres Tu.xtla, U.I.M.N.H. (6); 2 kilometers south of San Andres Tuxtla, U.M.M.Z. (2); Tierra Colorada, C.A.S. (1), S.U. (2), U.I.M.N.H. (12); 8 kilometers south of Veracruz, U.M.M.Z. (4); 8 kilometers soutli of Veracruz, U.M.M.Z. (4); 8 kilometers east of Zapoapan, T.O.W.C. (1). yucatan: Chichen-Itza, F.M.N.H, (29), U.M.M.Z. (1) 2.4 kilometers east of Chichen- Itza, U.M.M.Z. (1 tadpoles); 10 kilometers south of Chichen-Itza, U.M.M.Z. (4, 1 tadpoles); Culuba, 28 kilometers east of Sucopo, F.M.N.H. (13). BRITISH HONDURAS: Cayo: Cohune Ridge, U.NLM.Z. (6); Pine Ridge Road, U.M.M.Z. (4). 695 696 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 GUATEMALA: Alta Verapaz: Finca Chama, U.M.M.Z. (66, 1 eggs); Finca Samanzana, U.M.M.Z. (2 tadpoles). El Peten: 3 kilometers southeast of La Libertad, K.U. (7); Tikal, U.M.M.Z. (8); Sacrificio, .\.M.N.H. (1); Toocog, 15 kilometers southeast of La Libertad, K.U. (13, 12 skeletons, 1 tadpoles, 2 eggs). Izabal: 8 kilometers .south of Puerto Barrios, K.U. (8). HONDURAS: Atlantidad: Lancetilla, M.C.Z. ( 1); Toloa Junction, U.S.N. M. (1). Colon: Balfate, A.M.N.H. (2). Cortes: Agua Azul, A.M.N.H. (4); Lago Yojoa, A.M.N.H. (6); Rio Lindo, A.M.N.H. (2). NICARAGUA: Boaco: 14 kilometers north of, 13 kilometers east of Boaco, K.U. (1). Jinotega: Jino- tega, FM.N.H. (1). Managua: Casa Colorada, 22 kilo- meters south of Managua, K.U. (5, 1 skeleton, 2 tad- poles, 3 eggs). Matagalpa: Finca Tepeyac, K.LI. (9, 1 tadpoles); Hacienda La Cumplida, K.U. (2, 1 skele- ton, 1 eggs), U.M.M.Z. (18, 3 tadpoles). Zclaya: Bluefields, F.M.N.H. (1); Cukra, A.M.N.H. (1); El Recreo, K.U. (1); Isla Pequena del Maiz, M.C.Z. (2); Masahuas, Rio Huaspuc, A.M.N.H. (1); Rio Grande, M.C.Z. (1). COSTA RICA: Alajucla: Laguna Monte Alegre, K.U. (1); 3 kilometers northeast of Muelle de Arenal, U.S.C. (2). Cartago: Peralta, K.U. (1); Turrialba, K.U. (8), U.M.M.Z. (5), U.S.C. (10). Guanacaste: Finca San Bosco, K.U. (65, 5 skeletons, 1 tadpoles, 3 eggs), U.S.C. (35); Silencio, U.S.C. (16); Tilaran, K.U. (2). Heredia: Finca La Selva, U.M.M.Z. (1); Puerto Viejo. K.U. (2). Limon: Batan, K.U. (2); Colorado Bar, A.M.N.H. (1); El Tigre, 9 kilometers southwest of Siguirres, U.S.C. (1). Guapiles, A.N.S.P. (1); La Ca.stilla, A.N.S.P. (3); La Lola, U.F. (1), U.S.C. (4); 1.6 kilometers south of Limon, A.M.N.H. (1); Los Diamantes, U.M.M.Z. (2); Pandora, U.S.C. (7); Rio Lari at Rio Dipnari, U.S.C. (2); Rio Toro Amarillo, 7 kilometers west of Guapiles, K.U. ( 1 tad- poles); Suretka, K.U. (3); Tortugero, M.C.Z. (2), U.F. (3). Puntarenas: 3 kilometers northwest of Buenos Aires, K.U. (1); 6 kilometers northwest of Buenos Aires, K.U. (2); 10 kilometers east of Esparta, K.U. (1, 1 tadpoles); Golfito, U.M.M.Z. (3), U.S.C. (8); 4 kilometers east southeast of Palmar Sur, K.L'. (1); Parrita, U.S.C. (9), 10 kilometers northwest of Piedras Blancas, K.U. (4); 8 kilometers northeast of Potrero Grande, U.S.C. (1); 4.5 kilometers west of Rincon de Osa, K.U. (2, 1 tadpoles); Rio Ferruviosa, 7.2 kilometers south of Rincon de Osa, U.S.C. (1); 21.7 kilometers west of San Ramon, U.S.C. (14); 1.6 kilometers northwest of Villa Neily, U.S.C. (3). San Jo.se: San Lsidro el General, K.U. (4); 14 kilometers southwest of San lsidro el General, U.S.C. (1); 19 kilometers southwest of San lsidro el General, K.U. (2). PANAMA: Bocas del Toro: 3.2 kilometers north- west of Almirante, K.U. (5); 9.6 kilometers west of Almirante, K.U. (1); 12.8 kilometers west of Almir- ante, K.U. (1); Cayo de Agua, K.U. (3); Cayo Zapatilla Grande, K.U. (5); Isla de Colon, K.U. (4); Isla Popa, K.U. ( 1 ); mouth of Rio Cahuita, K.U. ( 1 ). Canal Zone: Barro Colorado Island, A.M.N.H. (3), A.N.S.P. (1), F.M.N.H. (2) K.U. (5, 4 .skeletons, 5 tadpoles. 2 eggs), T.N.H.C. (2); U.M.M.Z. (1); Camp Chagras, K.U. (4, 1 tadpoles); Gatiin, C.A.S. (1), M.C.Z. (1); Juan Mina, A.N.S.P. (2); Madden Forest, A.M.N.H. (7); 3.5 kilometers north of Mira- flores bridge, T.N.H.C. (19); San Pablo, M.C.Z. (1). Chiriqui: 13 kilometers west-northwest of Concepcion, K.U. (1); Progreso, U.M.M.Z. (1); Puerto Armuelles, C.A.S. (1). Colon: Achiote, K.U. (4). Darien: Camp Creek, below Yavisa, A.M.N.H. (1); Chalichi- man's Creek, Rio Sucubti, A.M.N.H. (2); Laguna, K.U. (13, 3 tadpoles); Rio Chucunaque at Rio Can- clon, U.M.M.Z. (1); Rio Tuira at Rio Mono, K.U.( (18); Rio Ucurganti, 7 kilometers above mouth, K.U. (3); Tacarcuna, K.U. (19. 2 .skeletons, 2 tadpoles, 2 eggs). Panama: Cerro La Campana, F.M.N.H. (25), K.U. (33, 2 tadpoles, 7 eggs), U.U. (4); 3 kilometers west-southwest of Chepo, K.U. (l);Tapia, A.M.N.H. (2). PANAMA: K.U. (1). Agalychnis litodryas Darien: Rio Tuira at Rio Mono, Agalychnis moreletii MEXICO: Chiapas: Acacoyagua, U.M.M.Z. (1 eggs); 6 kilometers northeast of Escuintla, U.M.M.Z. (6, 1 tadpo'es); Finca San Jeronimo, U.I.M.N.H. (14); Finca Juarez, S.U. (2), U.I.M.N.H. (22), U.M.M.Z. (4), Region Soconusco, U.I.M.N.H., U.I. (3). Oaxaca: Campamento Vista Hermosa, K.U. (2, 1 tadpoles); Mirador, A.M.N.H. (10); Nuevo Raza Sacatepec, U.I.M.N.H. (1); 28.2 kilometers north of Pochuda. U.M.M.Z. (1). Veracruz: Cuautlapan, K.U. (3), U.I.M.N.H. (6), U.M.M.Z. (1), U.S.N.M. (11); Escamilla, U.M.N.H. (5); 6.4 kilometers east of Fortin de las Flores, C.A.S. ( 1 ); 8 kilometers east of Orizaba, C.A.S. ( 1 ); San Andres Tu.xtla, U.S.N.M. ( 1 ); Volcan San Martin, K.U. (1). BRITISH HONDURAS: No specific locality, F.M.N.H. (2). Cai/o: Pine Ridge Road, U.M.M.Z. (22); Valentin, U.M.M.Z. (14). GUATEMALA: No specific locality. U.S.N.M. (5). Alta Verapaz: Finca Chichen, U.M.M.Z. (5); Finca Chicoyou, K.U. ( 66, 2 skeletons, 5 tadpoles, 4 eggs); Finca La Primavera, U.M.M.Z. (1); Finca Samac, U.M.M.Z. (3, 1 tadpoles); Finca Volcan, U.M.M.Z. (3); Senahii, U.S.N.M. (1). Huehuetenan- go: Barillas, U.M.M.Z. (4); Finca San Rafael, 16 kilometers southeast of Barillas, F.M.N.H. (4); Max- hal, north of Barillas, F.M.N.H. (1). Santa Rosa: Finca El Progreso, U.M.M.Z. (9); Finca La Gloria, U.M.M.Z. (2); Suchitepequez, Patutol, F.M.N.H. (1). Agalychnis saltator NICARAGUA: Zclai/a: Eden Mine, A.N.S.P. (2). COSTA RICA: Guanacaste: Finca San Bosco, K.U. (27, 1 skeleton), S.U. (2), U.S.C. (1). Heredia: Finca La Selva, U.S.C. (1); Puerto Viejo, K.U. (15, 2 skeletons, 1 tadpoles). Limon: La Castilla, .'\.N.S.P. (1). 1970 DUELLMAN: HYLID FROGS 697 Agalychnis spurrelli COSTA RICA: Funtarenas: Rincon de Osa, K.U. (1), U.S.C. (1); 4.5 kilometers west of Rincon de Osa, U.S.C. (1), K.U. (3 tadpoles, 1 eggs); Rio Ferroviosa, 7 kilometers south of Rincon de Osa, U.S.C. (4). San Jose: 16 kilometers southwest of San Isidro el General, U.S.C. (12). PANAMA: Bocas del Tom: Rio Urri, R.H. (1). Carial Zone: Barro Colorado Island, A.M.N.H. (1), A.N.S.P. (1), M.C.Z. (1), K.U. (8, 1 skeleton, 1 tadpoles). Darien: Tacarcuna, K.U. (9). Panama: Cabima, U.S.N.M. (1). Anotheca spinosa MEXICO: Oaxaca: Vista Hermosa, K.U. (13, 4 skeletons, 2 tadpoles), U.M.M.Z. (2); Yelagago, A.M.N.H. (1); 8 kilometers south of Yetla, K.U. (1). Veracruz: Barranca Metlac, U.M.M.Z. (1); Cuautla- pan, F.M.N.H. (48), K.U. (3), M.C.Z. (11), U.I.M.N.H. (62), U.M.M.Z. (18, 1 skeleton), U.S.N.M. (14); 9 kilometers southwest of Fortin de las Flores, U.M.Z. (9); 13 kilometers west northwest of Potrero, K.U. (2); Volcan San Martin, K.U. (3, 1 tadpoles), U.M.M.Z. (5, 1 tadpoles). COSTA RICA: Alajuela: 3 kilometers west of La Fortuna, U.S.C. (1). Cartago: Moravia, K.U, (2); Paloma, Valle de Orosi, U.S.N.M. (1). PANAMA: Bocas del Toro: Rio Changena, 830 meters, K.U. (2). Code: El Valle, U.M.M.Z. (1). Gastrotheca ceratophrys PANAMA: Bocas del Toro: 5 kilometers west of Almirante, K.U. (1); Rio Changena, 830 meters, K.U. (1); Rio Claro near junction with Rio Changena, K.U. (3, 1 skeleton). Darien: Laguna, K.U. (1); Tacarcuna, U.S.N.M. (1). Panama: Upper Rio Pequeni, U.S.N.M. (1). San Bias: Camp Summit (3). Gastrotheca nicefori PANAMA: Darien: South slope of Cerro Citurio, Serrania de Pirre, K.U. (2); Ridge between Rio Jaque and Rio Imamado, Serrania del Sapo, K.U. ( 1 ). Hemiphractus panamensis PANAMA: Bocas del Toro: north slope Cerro Pando, 1450 meters, K.U. (1); Rio Changena, B.Y.U. (1 + young), R.H. (1). Colon: Signal Loma, 5 kilo- meters south of Santa Isabel, U.S.N.M. (2). Darien: Cerro Citurio, K.U. (18), Cerro Pirre, G.M.L. (1), K.U. (5, 2 skeletons). Panama: Altos de Pacora, K.U. (2). San Bias: Camp Summit, K.U. (3). Hyla altipotens MEXICO: Oaxaca: 33 kilometers north of San Gabriel Mixtepec, K.U. (1); 37 kilometers north of San Gabriel Mixtepec, K.U. (25, 2 skeletons); 3 kilo- meters east of San Sebastian (Los Fustes), T.C.W.C. (1). Hyla angustilineata COSTA RICA: Heredia: Rama Sur Rio Las Vueltas, south slope of Volcan Barba, K.U. (21, 2 skeletons, 4 tadpoles), M.C.Z. (2), U.S.C. (2). Funtarenas: 3 kilometers east-northeast of Monteverde, U.S.C. (2). Sanjose: La Palma, U.S.N.M. (3). Hyla arborescandens MEXICO; Hidalgo: 10 kilometers south of Zacual- tipan, I.P.N. (2). Oaxaca: 4.2 kilometers south of Campamento Vista Hermosa, K.U. (14, 1 skeleton, 1 tadpoles); 6 kilometers south of Campamento Vista Hermosa, K.U. (9, 1 skeleton); 7.5 kilometers south of Campamento Vista Hermosa, K.U. (1); 10 kilo- meters south of Campamento Vista Hermosa, K.U. (4); 15 kilometers south of Campamento Vista Her- mosa, K.U. (2), U.M.M.Z. (1); Cerro Machin, U.I.M.N.H. (3); Cerro San Felipe, F.M.N.H. (3), U.I.M.N.H. (4), U.S.N.M. (1); 13 kilometers north- west of Ixtlan de Juarez, T.N.H.C. (3); 25.5 kilo- meters north of Nochixtlan, U.I.M.N.H. (2); 52 kilo- meters north of Oaxaca, U.I.M.N.H. (2). 68 kilome- ters south of Valle Nacional, U.M.M.Z. (4). Fuebla: 14.4 kilometers west of Huachinango, Paraje Verde, U.I.M.N.H. (2), U.S.N.M. (7); Puente Colorado, U.I.M.N.H. (2), U.M.M.Z. (1); Rio Octapa, 3.7 kilo- meters north-northeast of Tezuitlan, K.U. (9, 1 skele- ton). Tlaxcala: Apizaco, U.S.N.M. (1). Veracruz: Acultzingo, F.M.N.H. (2) I.P.N. (1), U.I.M.N.H. (4); 5 kilometers southwest of Acultzingo, U.I.M.N.H. (1); Cumbres de Acultzingo, F.M.N.H. (3), U.I.M.N.H. (5), U.M.M.Z. (16), U.S.N.M. (5). 1.9 kilometers southwest of La Joya, F.M.N.H. (30); La Perla, M.C.Z. (1); Pan de Olla, M.C.Z. (2), U.I.M.N.H. (17); U.S.N.M. (25); Tegueyutepec, M.C.Z. (3) Hyla arenicolor MEXICO: Aguascalientes: 29 kilometers west, 3 kilometers south of Aguascalientes, K.U. ( 1 ) ; 14.4 kilometers north of Rincon de Romos, U.I.M.N.H. (1). Chihuahua: Balleza, U.S.N.M. (1); north rim of Barranca del Cobre, 37 kilometers south, 2.4 kilo- meters east of Creel, K.U. (8); Batopilas, U.S.N.M. (1); Chihuahua, U.M.M.Z. (1); 24 kilometers south, 10 kilometers east of Creel, K.U. (3); Divisadero, 25.6 kilometers south, 21 kilometers west of Creel, K.U. (13); 21 kilometers west of Guatemoc, M.C.Z. (1); 13 kilometers southwest of Hidalgo del Parral, T.C.W.C. (1); 60 kilometers southwest of Juanito, U.S.N.M. (1); La Pulvosa, U.M.M.Z. (2); Maguaric- hic, U.M.M.Z. (2); 3 kilometers west of Minaca, K.U. (2); Mojarachic, F.M.N.H. (2), U.I.M.N.H. (1), U.M.M.Z. (1); Rio del Sauz, west of Sauz, A.M.N.H. ( 1 ) ; 3 kilometers south, 8 kilometers west of San Francisco, K.U. (3); San Rafael, between Santa Barbara and Parral, A.M.N.H. (1); Sauz, F.M.N.H. (1); 5 kilometers northeast of Temoris, K.U. (8); Wasichichic, U.M.M.Z. (1). Coahuila: No specific locality, F.M.N.H. (3). Distrito Federal: Pedregal de San Angel, I.P.N. (1); 2.5 kilometers west of 698 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Santa Fe, A.M.N. H. ( 1). Durango: No specific local- ity, U.S.N. M. (1); Cerro de Mercado, A.N.S.P. (1); Coyotes, U.M.M.Z. (1); El Salto, U.S.N. M. (1); 42,7 kilometers northeast of El Salto, U.I.M.N.H. (6); 1.6 kilometers west of E! Salto, L.B.S.C. (2); 5 kilometers west of El Espinosa, L.B.S.C. (1); Laguna del Pro- greso, U..M.M.Z. (3); 50 kilometers southwest of V'ictoria de Durango, U.I.M.N.H. (2). Guanajuato: No specific localit>', U.S.N.M. (4); 6 kilometers west of Acambaro, F.M.N.H. (1); 6 kilometers north of, 8 kilometers west of Leon, K.U. (1); 11 kilometers northwest of Leon, U.LM.N.H. (5); 7 kilometers south of Valle de Santiago, U.I.M.N.H. (1). Guer- rero: No specific locality, U.M.M.Z. ( 1 ); Acahuitzotla, K.U. (1), T.C.W.C. (9); 3 kilometers north of Aca- huitzotla, F.M.N.H. (5), U.I.M.N.H. (1); Agua del Obispo, F.M.N.H. (4), T.C.W.C. (1), U.I.M.N.H. (3), U.M.M.Z. (1). Chilapa, U.S.N.M. (1); east of Chilapa, K.U. ( 1 ); 5 kilometers south of Chilpancingo, U.F. (2); 19 kilometers south of Chilpancingo, F.M.N.H. (22), U.I.M.N.H. (15); Palo Blanco, F.M.N.H. (1), U.I.M.N.H. (1); 18 kilometers south of Puente de L\tla (Morelos), F.M.N.H. (4), U.I.M.N.H. (8); San Juan, U.S.N.M. (1); 8 kilo- meters north of Ta.xco, T.C.W.C. (2). Hildalgo: 18 kilometers southeast of Actopan, K.U. (2), T.C.W.C. (13); 8 kilometers west of Actopan, T.C.W.C. (4); 30 kilometers east of Huichapan, T.C.W.C. (5); 11 kilometers southwest of Huichapan, K.U. (1); 9.4 kilometers north of Metzquititliin, K.U. (1); Miguel, M.C.Z. (1); Tianguistengo, F.M.N.H. (1). Jalisco: No specific locality, U.S.N.M. (2); 11 kilometers west of Ameca, U.M.M.Z. (2); Atemaje, A.M.N.H, (2); 3 kilometers west of Ayutla, K.U. (5), Cerro del Col, A.M.N.H. (1); Cerro Pelon, Rio Blanca, north of Zapopan, A.M.N.H. (2); Chapala, U.S.N.M. (1); 4 kilometers northeast of Ciudad Guzman, F.M.N.H. (1); 8 kilometers northwest of Cuautla, K.U. (1); 5 kilometers northwest of Degollado, K.U. (1); Guada- lajara, K.U. (10); 5 kilometers nordi of Guadalajara, K.U. (2); 33 kilometers .southwest of Guadalajara, K.U. (2); Hostolipaquillo, A.M.N.H. (4); 3 kilome- ters east of I.xdahuacan del Rio A.M.N.H. (1); 10.4 kilometers north-northwest of I.xtlahuacan del Rio, K.U. (1); La Mesa Maria de Leon, K.U. (11); 5 kilo- meters northeast of Magdalena, K.U. (4); Rancho Primavera, near Guadalajara, U.I.M.N.H. (2); Rio Blanco, near Guadalajara, K.U. (2); San Gabriel, U.M.M.Z. (1); 3 kilometers northeast of Talpa, K.U. (2); 7 kilometers west of Tenchitlan, K.U. (1); be- tween Tecjuesquite and Hostolipaquillo, A.M.N.H. (5); Tlaquepaque, A.M.N.H. (7); Tonolii, A.M.N.H. (2); between Tonold and Tlaquepaque, A.M.N.H. (1); 14.4 kilometers northeast of Union Tula, K.U. (1); 2.5 east of Villa Guerrero, K.U. (1); 6.4 kilo- meters west of Villa Guerrero, K.U. (5); 1.2 kilome- ters north of, 1 1 kilometers west of Yahualica, K.U. Yahualica, K.U. (2). Mexico: 11 kilometers south of Yahualica, K.U. (2). Mexico: II kilometers south of, Acambay, K.U. (1); San Juan Teotihuaciin, K.U. (I), M.C.Z. (2), U.M.M.Z. (1); Tonatico, I.P.N. (1). Michoacdn: Agua Cerca, U.M.M.Z. (1); Cascada Tzararacua, U.M.M.Z. (5); Chinapa, U.M.M.Z. (1); 6 kilometers east of Cojumatlan, F.M.N.H. (1), U.I.M.N.H. (1); Dos Aguas, U.M.M.Z. (1); El Espinal, U.M.M.Z. (1); El Sabino, F.M.N.H. (22), U.I.M.N.H. (6); Lago de Camecuaro, U.M.M.Z. (1); Lombardia, U.M.M.Z. (2); Tupataro, U.S.N.M. (1); 1.2 kilometers northwest of Zinapecuaro, K.U. (1). Morelos: 19 kilometers north of Cuautla, T.C.W.C. (2); 18 kilometers southeast of Cuautla, T.N.H.C. (4); Cuernavaca, U.I.M.N.H. (3), U.M.M.Z. (2), U.S.N.M. (1); 3 kilometers north of Cuernavaca, F.M.N.H. (4); 11 kilometers east of Cuernavaca, U.I.M.N.H. (2); Huajintlan, F.M.N.H. (6); 2 kilo- meters south of Jonacatepec, T.C.W.C. (3). Nayarit: 1.6 kilometers east of I.xtlan del Rio, U.M.M.Z. (1); La Mesa de Nayarit, A.M.N.H. (4), Sierra de Nayar, A.M.N.H. (1); 37 kilometers south of Tepic, L.B.S.C. (6). Oaxaca: 3 kilometers east of Huajapan de Leon, K.U. (1); 32 kilometers southeast of Huajapan de Leon, U.I.M.N.H. (1). Puebla: 13 kilometers south- east of Izucar de Matamoros, C.A.S. (1). Queretaro: Cadereyta, U.M.M.Z. (3); 11 kilometers west-south- west of San Juan del Rio, K.U. (1); Tequisquiapan, A.M.N.H. (3). San Luis Potosi: Ahualulco, U.S.N.M. (1); Alvarez, M.C.Z. (2), Cerro de Alvarez, A.N.S.P. (2); Cerro de MigueHto, A.N.S.P. (3); Morales, M.C.Z. (1); U.M.M.Z. (1); San Luis Potosi. M.C.Z. (5), U.I.M.N.H. (4); 36 kilometers north of San Luis Potosi, M.C.Z. (4); 43 kilometers south of San Luis Potosi, M.C.Z. (6); 18 kilometers southwest of San Luis Potosi, U.M..M.Z. (3); 5 kilometers west of San Luis Potosi, U.I.M.N.H. (8); San Pedro, A.N.S.P. (1); 3 kilometers north of Santa Maria del Rio, A.M.N.H. (1). Sinaloa: No specific localitv-, U.S.N.M. (2); Plumosas, U.S.N.M. (I); 70 kilometers north- east of Villa Union, L.B.S.C. (6), 75 kilometers northeast of Villa Union, L.B.S.C. (1). Soiiora; 14.4 kilometers north of Imuris, K.U. (1); 3 kilometers from La Poza, 10 kilometers north of Guavmas, F.M.N.H. (6); Nogales, U.S.N.M. (1); Pi'lares, U.M.M.Z. (5); San Jose de Guayamas, M.C.Z. (1); northern Sonora, U.S.N.M. (1). Veracruz: 10 kilo- meters southwest of Jacales, K.U. (2); 6 kilometers west-southwest of Zacualpilla, K.U. (73). Zacatccas: 3 kilometers .southeast of Laguna Valderana, U.M.M.Z. (3); 17.6 kilometers northwest of Jalpa, K.U. (3); 13 kilometers south of Moyahua, C.A.S. ( 1 ); Plateado, U.S.N.M. (1); 5 kilometers northwest of Teul, U.M.M.Z. (40). Hyla bistincta MEXICO: Durango: 5 kilometers west of El Espinosa, L.B.S.C. (1). Guerrero: Omiltemi, U.I.M.N.H. (3); between Puerto Chico and Aso- leadero, U.M.M.Z. (1); 22 kilometers southwest of Ye.vtla, I.P.N. (8). Hidalgo: Zacualtipan, A.N.S.P. ( I ) . Jalisco: 25 kilometers southeast of Autlan, U.M.M.Z. (1). Mexico: 19 kilometers west of Villa Victoria, U.I.M.N.H. (1). U.S.N.M. (1). Morelos: Cuernavaca, U.S.N.M. (1); 3 kilometers north of Cuernavaca, U.I.M.N.H. (3). Michoacdn: Cerro San Andres, U.M.M.Z. (1); Dos Aguas, U.M.M.Z. (1); 12.5 kilometers east-nordieast of Dos Aguas, U.M.M.Z. 1970 DUELLMAN: HYLID FROGS 699 (1); Los Conejos. U.M.M.Z, (3); Uruapan, K.U. (2, 1 skeleton), U.I.M.N.H. (2); U.M.M.Z. (33, 2 skele- tons, 1 tadpoles), U.S.N. M. (10). Nayarit: Santa Teresa, U.S.N.M. (1). 10 kilometers northeast of Avutla, .\.M.N.H. (1); Cerro San Felipe, U.I.M.N.H. (22); Huautla, A.M.N.H. (1); San Lucas Camotlan, U.S.N.M. (1). Sinaloa: 1.6 kilometers east of Santa Lucia, K.U. (1). Veracruz: No specific locality, U.S.N.M. (1); Cumbres de Acultzingo, F.M.N.H. (2),U.LM.N.H. (5), U.S.N.M. (1). Hyla boans PANAMA: Colon: Rio Candelaria, A.M.N.H. (2). Darien: Paya, U.U. (5); Rio Tuira at Rio Mono, K.U. (2). San Bias: Camp Sasardi, K.U. (15, 4 skele- tons, 3 tadpoles, 1 eggs). Hyla bogertae MEXICO: Oaxaca: tributary of Rio Atoyac, be- low V'ivero El Tapanal, 1.6 kilometers south of La Cofradia, Distrito de Sola de Vega, L.A.C.M. (13, 1 tadpoles, 1 young). Hyla boulengeri NICARAGUA: No specific locality, U.S.N.M. (1). Chontales: 1 kilometer north, 25 kilometers west of Villa Somoza, K.U. (1). Zelaija: El Recreo, K.U. (2), U.M.M.Z. (1); Kanawa, A.M.N.H. (1); Sioux Plantation, A.M.N.H. (1), M.C.Z. (1); Tuli Creek, A.M.N.H. (1). COSTA RICA: Alajuela: 9 kilometers north of Ciudad Quesada, U.S.C. (4); 18 kilometers north of La Florencia, U.S.C. ( 1 ); Laguna Monte Alegre, K.U. ( 1 ) ; Las Playuelas, 1 1 kilometers south of Los Chiles, U.S.C. (4); 3 kilometers northeast of Muelle de Arenal, U.S.C. (5). Cartago: Turrialba, K.U. (1). Guanacaste: Finca Taboga, 20 kilometers southeast of Las Caiias, K.U. (1), U.S.C. (1); 7 kilometers north of Liberia, U.S.C. (8); 13.6 kilometers north of Liberia, U.S.C. (3); 20.5 kilometers south of Liberia, U.S.C. (1); 4 kilometers northeast of Tilaran, L'.S.C. ( 1 ); 6 kilometers northeast of Tilaran, U.M.M.Z. ( 1 ), U.S.C. (7). Heredia: Puerto Viejo, K.U. (44, 6 skeletons), M.C.Z. (5); 1 kilometer northeast of Puerto Viejo, U.M.M.Z. (1); 4.2 kilometers west of Puerto Viejo, K.U. (1, 1 skeleton). Limon: Mountain Cow Creek, near Banano, K.U. (1); Suretka, K.U. (8, 1 skeleton); Tortugero, A.M.N.H. (1). Funtare- nas: Parrita, U.S.C. (1); 6 kilometers southwest of Rincon de Osa, K.U. (6); 4.5 kilometers southwest of Rincon de Osa, K.U. (3, 2 tadpoles) Rio Rincon, 4.8 kilometers south of Bahia Rincon, U.S.C. (1); 4.4 kilometers northwest of Villa Neily, U.S.C. (1); 10.5 kilometers west-northwest of Villa Neily, K.U. (1). San Jose: 21 kilometers west-southwest of San Isidro el General, K.U. (3). PANAMA: Bocas del Toro: 3.2 kilometers west of Almirante, K.U. (1). Canal Zone: Barro Colorado Island, A.N.S.P. (4), F.M.N.H. (1); Fort Clayton, U.I.M.N.H. (1); 2.8 kilometers southwest of Fort Kobbe, K.U. (1); between Gatuncillo and Guaya- balito, A.M.N.H. (8); Juan Mina, A.N.S.P. (1); K.U. (1), U.U. (2); 10 kilometers northwest of Miraflores Locks, A.M.N.H. (2); Road K2, T.N.H.C. (10); Road K9, T.N.H.C. (1); Summit Gardens, A.M.N.H. (1), A.N.S.P. (2), K.U. (1, 1 skeleton). Colon: Ciricito, C.A.S. (1); Rio Gatuncillo, near Nuevo San Juan, K.U. (1). Darien: El Real, K.U. (3). Hyla bromeliacia GUATEMALA: Aha Verapaz: Finca Chicoyou, near Coban, K.U. (1, 1 skeleton, 3 tadpoles, 1 eggs); Finca Samac, F.M.N.H. (1), U.M.M.Z. (6, 1 tad- poles). El Quiche: Finca San Francisco, U.M.M.Z. ( 2, 1 tadpoles ) . HONDURAS: Atlantidad: Mountains behind La Ceiba, M.C.Z. (1). Cortes: Mountains west of San Pedro Sula, F.M.N.H. (6), M.C.Z. (3). Hyla cadaverina MEXICO: Baja California Norte: Caiion Guada- lupe, Sierra de Juarez, L.B.S.C. (2), U.M.M.Z. (11); Caiion de las Palmas, Sierra de Juarez, L.B.S.C. (2); Cafion de Llanos, 14.4 kilometers soutli-southwest of La Rumorosa, M.V.Z. (14); Cafion de Tajo, west side of Laguna Salada, U.M.M.Z. (3), Caiion La Provi- dencia, east base of Sierra San Pedro Martir, U.S.N.M. (3); Ensenada U.S.N.M. (1); Isla Navidad, U.S.N.M. (1); La Laguna, Sierra La Laguna, U.S.N.M. (1). Osos Negros, U.S.N.M. (1); Playa Estero 14.4 kilo- meters south of Ensenada, A.M.N.H. (1); 32 kilo- meters east of Rosario, U.M.M.Z. (2). Hyla chaneque MEXICO: Chiapas: 5.6 kilometers south of Rayon Mescalapa, K.U. (1); 6.2 kilometers south of Rayon Mescalapa, K.U. (4, 1 skeleton), U.M.M.Z. (1). Oaxaca: 42.6 kilometers south of Valle Nacional, U.M.M.Z. (9); 43.5 kilometers south of Valle Nacio- nal, U.M.M.Z. (2), K.U. (12, 2 skeletons, 3 tadpoles), M.C.Z. (5); 6 kilometers south of Campamento Vista Hermosa, K.U. (5, 2 tadpoles), U.M.M.Z. (2 tad- poles ) ; 8 kilometers south of Campamento Vista Her- mosa, K.U. (3); 11 kilometers south of Campamento Vista Hermosa, K.U. (1, 2 tadpoles); 13 kilometers south of Campamento Vista Hermosa, K.U. (1); 16 kilometers south of Campamento Vista Hermosa, K.U. (1), U.I.M.N.H. (2); Sierra Madre above Zanatepec, K.U. (1), U.I.M.N.H. (1). 8 kilometers south of Yetla, K.U. (2). Hyla charadricola MEXICO: Hidalgo: Lago de Tejocotal, 11 kilo- meters east of Acaxochitliin, K.U. (1), U.M.M.Z. (2); 4 kilometers southwest of Tianguistengo, K.U. (2). Pucbla: 11.7 kilometers west of Huachinango, U.M.M.Z. (5); Rio Totolapa, 14.4 kilometers west of Huachinango, K.U. (38, 3 skeletons), M.C.Z. (2), U.I.M.N.H. (1), U.M.M.Z. (5, 1 skeleton). 700 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Hyla chryses MEXICO: Guerrero: between Puerto Chico and Asoleadero, 45 kilometers airline west-northwest of Chilpancingo, K.U. (1), U.M.M.Z. (3). Hyla colymba COSTA RICA: Cartago: Moravia, K.U. (2, 1 skeleton ) . PANAMA: Bocas del Toro: La Loma, M.C.Z. (2); Rio Changena, 650 meters, K.U. (1); Rio Changena, 830 meters, K.U. (1 tadpoles). Code: El Valle, A.M.N. H. (1). Daricn: Cerro Citurio, Serrania de Pirre, K.U. (1); Cerro Mali, G.M.L. (1); Laguna, K.U. (1, 1 tadpoles); Ridge liet\veen Rio Jaque and Rio Imamado, K.U. (2, 1 tadpoles). Panama: Altos de Pacora, K.U. (1). Hyla crassa MEXICO: No specific locality, M.N. H.N. (1). Oaxaca: Cerro San Felipe, U.I.M.N.H. (1). Hyla crepitans HONDURAS; Cortes: Tela, A.M.N. H. (1 ). PANAMA: Canal Zone: Alhajuela, U.M.M.Z. (7); Camp Chagres, K.U. (16, 1 skeleton); Fort Kobbe (1); Juan Mina, F.M.N.H. (1); junction roads K2 and K9, T.N.H.C. ( 19); San Pablo, A.M.N. H. (I), M.C.Z. (I), U.M.M.Z. (3); Summit Gardens, A.N.S.P. (3), F.M.N.H. (2), M.C.Z. (I). Code: El Valle, C.A.S. (1), K.U. (1). Los Santos: Guanico Arriba, K.U. (I); Tonosi, K.U. (1). Panama: 3 kilo- meters west-southwest of Chepo, K.U. (3); Chilibre, U.S.N.M. (I); Finca La Sumbadora. K.U. (3); Las Cumbres, A.M.N. H. (1); Panama, A.M.N. H. (1), K.U. (I), M.C.Z. (2); Rio Abajo, K.U. (1), U.U. (1); Rio Mamoni, 5 kilometers east of Chepo, K.U. ( 1 ); 17 kilometers east of Tocumen, M.V.Z ( 1 ). Hyla debilis COSTA RICA; Cartago: Tapanti, U.S.C. (15, 1 skeleton). Hcredia: Isla Bonita, F.M.N.H. (1),K.U. (1). PANAMA: Bocas del Toro: north slope of Cerro Pando, 1450 meters, K.U. (14, 2 skeletons, 4 tad- poles); Rio Claro near Rio Changena, K.U. (I). Chiruitii: Boquete, U.M.M.Z. (I). Hyla dendroscarta MEXICO: Oaxaca: Campamento Vista Hermosa, K.U. (4); 7.5 kilometers south of Campamento Vista Hermosa, K.U. (2). Veracruz: Barranca Metlac, M.C.Z. (I), U.I.M.N.H. (2), U.M.M.Z. (9); Cuaut- lapan, F.M.N.H. (I), K.U. (I), M.C.Z. (1), U.I.M.N.H. (136), U.S.N.M. (36); 12.4 kilometers .southwest of Fortin de las Flores, U.M.M.Z. (4); 3 kilometers southwest of Huatusco, U.M.M.Z. (10, 1 tadpoles); Mirador, K.U. (1 tadpoles), U.M.M.Z. (3, 1 skeleton, 1 tadpoles); 3 kilometers east of San Andres Tu.vtla, K.U. (25); Sumidero, M.C.Z. (I); 15 kilometers east-northeast Tlacotepec, K.U. ( I ). Hyla ebraccata MEXICO: Chiapas: 25 kilometers south of Chon- talpa, A.M.N.H. (1); 17 kilometers south of Teapa (Tabasco), U.I.M.N.H. (4). Oaxaca: 3 kilometers north of Donaji, U.M.M.Z. (17); 21.7 kilometers south of Jesus Carranza (Veracruz), U.I.M.N.H. (2); 78 kilometers north of La Venta, T.N.H.C. (24); 43 kilo- meters north of Matias Romero, U.I.M.N.H (27); 3 kilometers south of Papaloapan, A.M.N.H. (I); 3.7 kilometers north of Sarabia, U.M.M.Z. (6); 1.6 kilo- meters south of Tolocita, U.M.M.Z. (2); 3 kilometers south of Tolocita, K.U. (I); Ubero, U.M.M.Z. (14); 5 kilometers south of Ubero, U.M.M.Z. (I); 1.6 kilo- meters south of Valle Nacional, K.U. (21 ), U.I.M.N.H. (2). Tabasco: Teapa, U.M.M.Z. ( 15); 10 kilometers north of Teapa, U.M.M.Z. (1); 15 kilometers north of Teapa, U.M.M.Z. (7). Veracruz: 4.5 kilometers north of Aguilera, U.M.M.Z. (1); 10 kilometers south of Catemaco, U.M.M.Z. (3); Coyame, U.I.M.N.H. (6), U.M.M.Z. (7); Encinal, U.M.M.Z. (28); north side of Lago de Catemaco, K.U. (2); 25 kilometers south of Las Choapas, T.C.W.C. (19). BRITISH HONDURAS; No specific locality, F.M.N.H. (2). Cayo: 16 kilometers from Belize-Cayo Road on Hummingbird Highway. U.M.M.Z. (1); Cohune Ridge, M.C.Z. (I), U.M.M.Z. (18); 5 kilome- ters from Millionaro Camp on Pine Ridge Road, U.M.M.Z. (6). Stann Creek: Hummingbird Highway between Roaring Creek and Stann Creek, U.M.M.Z. (2). GUATEMALA: £/ Peten: 8 kilometers south of La Libertad, U.S.N.M. (5); Posa de Jicotea, 8 kilo- meters south of Piedras Negras, U.I.M.N.H. (2); Toocog, 15 kilometers southeast of La Libertad, K.U. (56, 10 skeletons, 2 eggs, 2 tadpoles); Yaxha, U.M.M.Z. (1). NICARAGUA: Matagalpa: 2 kilometers north, 6 kilometers east of E.squipulas, K.U. (1). Zelaya: Machuca. A.N.S.P. (1). COSTA RICA; Alajuda: Las Playuelas, 11 kilo- meters south of Los Chiles, U.S.C. (1). Cartago: Lago Bonilla, K.U. (6, 2 skeletons); Moravia de Tur- rialba, K.U. (22, 4 skeletons, 1 eggs), U.S.C. (12); 11 kilometers southwest of Moravia de Turrialba, K.U. ( 1 ); 1 kilometer east-northeast of Pacuare, K.U. (48); 2 kilometers south of Paraiso, U.S.C. (17); Turrialba, F.M.N.H. (56), K.U. (91, 5 skeletons), M.C.Z. (I), U.M.M.Z. (14), U.S.C. (45), U.S.N.M. (8). Guana- caste: Finca San Bosco, LI.S.C. (37); Laguna Arenal, U.S.C. (1); Silencio, U.S.C. (42); Tilaran, K.U. (38); 2 kilometers east of Tilaran, K.U. (20, 1 eggs, I tadpoles), U.S.C. (9). Heredia: Finca La Selva, U.S.C. (3); Puerto Viejo, K.U. (13, 3 tadpoles). Union: Bambu, U.S.C. (2); Guacimo, U.S.C. (I); Linion, A.M.N.H. (4); Los Diamantes, F.M.N.H. (2); Suretka, K.U. (1, 2 skeletons), M.C.Z. (2). Puntarenas: 1.6 kilometers south of Agua Buena, 1970 DUELLMAN: HYLID FROGS 701 U.S.C. (2); 5 kilometers west-northwest of Barranca, U.M.M.Z. (5); 3 kilometers northeast of Boca de Barranca, U.S.C. (1); 12 kilometers west-northwest of Esparta, K.U. (1); Esquinas Forest Preserve, be- tween Pahnar and Golfito, K.U. (1); 3 kilometers east of Golfito, K.U. (2); Palmar, K.U. (1); 3 kilometers southeast of Palmar Norte, K.U. ( 1 ); 4 kilometers east- southeast of Palmar Norte, K.U. (2); 3 kilometers northwest of Piedras Blancas, K.U. (1); Rincon de Osa, U.M.M.Z. (6), U.S.C. (7); 4.5 kilometers west of Rincon de Osa, K.U. (26); 6 kilometers southwest of Rincon de Osa, K.U. (10); Rio Ferruviosa, 7 kilo- meters south of Rincon de Osa, U.S.C. (12); 7 kilo- meters west of Villa Neily, U.S.C. (1); 10.5 kilome- ters west-northwest of Villa Neilly, K.U. (5, 2 skele- tons ) . PANAMA: Bocas del Tow: 2.5 kilometers west of Almirante, U.U. (2); 3.2 kilometers northwest of Almirante, K.U. (4); Isla Bastimentos, K.U. (1). Caual Zone: No specific locality, T.N.H.C. (90); be- tween Catuncillo and Guayabilito, A.M.N. H. (17); Juan Mina, U.U. (4); Rio Chagres, near Ganiboa, U.M.M.Z. (1); Summit, K.U. (5), M.C.Z. (1). Chiriqui: Progreso, U.M.M.Z. (2). Code: El Valle, A.M.N.H. (13), F.M.N. H. (1), K.U. (1), M.V.Z. (3). Colon: Acliiote K.U. (67, 4 skeletons); Ciricito, C.A.S. (4). Darien: Camp Townsend, Rio Chucu- naque, A.M.N.H. (1); Laguna, K.U. (2); Rio Tuira at Rio Mono, K.U. (23, 4 skeletons); Rio Ucurganti, 7 kilometers above mouth, K.U. (2); Tacarcuna, K.U. ( 40, 1 tadpoles ) . Panama: south slope of Cerro La Campana, F.M.N.H. (17), K.U. (17); 3 kilometers west-southwest of Chepo, K.U. (3); Tapia, A.M.N.H. (1); 33 kilometers east of Tocumen, M.V.Z. (3). San Bias: Sasardi, K.U. ( 1 ) . Hyla echinata MEXICO: Oa.xaca; Vista Hermosa, U.I.M.N.H. (1), U.M.M.Z. (1). Hyla elaeochroa NICARAGUA: No specific locality, U.S.N.M. (1). Boaco: 14 kilometers north, 13 kilometers east of Boaco, K.U. (1). Choniales: 1 kilometer north, 2.5 kilometers west of Villa Somoza, K.U. (13). Nueva Segovia: 5 kilometers north, 2.5 kilometers east of Jalapa, K.U. (2). Zelatja: Cukra, A.M.N.H. (1); El Recreo, U.M.M.Z. (1); Tuli Creek, A.M.N.H. (2). COSTA RICA: Alaiuela: 9 kilometers north of Ciudad Quesada, U.S.C. (5); Laguna Monte Alegre, K.U. (1); Las Vegas, 8 kilometers south of Tanque, U.S.C. (5); Los Chiles, A.M.N.H. (1). Cartago: Cervantes, U.S.C. (1); Chitaria, A.N.S.P. (1), 2 kilo- meters east of Chitaria, K.U. (1); El Silencio, 14.4 kilonifters northeast of Turrialba, K.U. (2); Juan \inas, U.S.C. ( 1 ); 9 kilometers from La Suiza, U.S.C. (18); 4.6 kilometers east-northeast of Pacuare, K.U. (35); 4 kilometers south of Pavones, K.U. (1); Peral- ta, K.U. (6); Turrialba, F.M.N.H. (33), K.U. (146, 19 skeletons, 2 tadpoles, 1 eggs), M.C.Z. (1), T.N.H.C. (1), U.M.M.Z. (12), U.S.C. (174); 5 kilo- meters south of Turrialba, U.I.M.N.H. (1). Guana- caste: 2 kilometers east of Tilaran, K.U. (22, 2 tad- poles). Hereclia: Finca La Selva, U.M.M.Z. (1); Puerto Viejo, K.U. (27, 11 skeletons, 7 tadpoles), M.C.Z. (2), U.M.M.Z. (1). Limdn: Bambu, U.S.C. (2); Batan, K.U. (10); La Castilla, Rio Reventazon, A.N.S.P. (7); La Lola, K.U. (21, 2 skeletons), U.M.M.Z. (3, 1 tadpoles); Limon, K.U. (8); Los Diamantes, K.U. (2), U.M.M.Z. (11); 2.5 kilometers east of Los Diamantes, U.S.C. (8); Pandora, U.M.M.Z. (4), U.S.C. (1); Portete, U.M.M.Z. (5); Sipurio, U.S.N.M. (1); Suretka, K.U. (16, 1 skeleton); Tor- tuguero, M.C.Z. (1), U.F. (10); Zent, M.C.Z. (4). Piintarenas: 5 kilometers northwest of Buenos Aires, K.U. (1); 10 kilometers east of Esparta, K.U. (1 tadpoles); Golfito, K.U. (3), U.M.M.Z. (2); 3 kilo- meters southeast of Golfito, U.S.C. (4); Gromaco, U.M.M.Z. (1); 2.5 kilometers southeast of Palmar Sur, K.U. (9); 4 kilometers .southeast of Palmar Sur, K.U. (2 tadpoles); 10.7 kilometers southeast of Palmar Sur, K.U. (1 skeleton); 13 kilometers southeast of Palmar Sur, K.U. (3, 1 eggs, 4 tadpoles); Parrita, U.S.C. (9); 3 kilometers northwest of Piedras Blancas, K.U. (14); 10 kilometers northwest of Piedras Blancas, K.\J. (14); 11 kilometers north, 3 kilometers west of Puntarenas, T.C.W.C. (7); Quebrada Boruca, 22 kilometers east of Palmar Norte, K.U. (1); Rincon de Osa, K.U. (15); 4.5 kilometers west of Rincon de Osa, K.U. (19, 1 tadpoles); Rio Zapote, 8 kilometers east of Palmar Norte, K.U. ( 1 ). PANAMA: Bocas del Tow: Almirante, K.U. ( 1 ), U.S.N.M. (1); 3 kilometers west of Almirante, K.U. (1), U.U. (1); Isla Bastimentos, K.U. (4); Rio Cricamola, 3.7 kilometers from mouth, K.U. (1); Rio San San, M.C.Z. (2). Chiriqui: Puerto Armuelles, A.N.S.P. (1); Progreso, U.M.M.Z. (6); Rio Garichine, 8.3 kilometers east-southeast of Paso de Ganoas, K.U. (2). Hyla erythromma MEXICO: Guerrew: Acahuitzotla, T.C.W.C. (1); Agua del Obispo, F.M.N.H. (1), K.U. (1, 2 tadpoles), T.C.W.C. (I), U.M.M.Z. (5); 1.6 kilome- ters east of San Andreas de la Cruz, K.U. (2); 3.3 kilometers north of San Vincente, K.U. (5). Oaxaca: 5 kilometers S. Yetla, K.U. (6); 6.9 kilometers south of Yetla, K.U. (1); 8 kilometers south of Yeda, K.U. (51, 2 skeletons, 5 tadpoles, 1 eggs); U.I.M.N.H. (4). Hyla euphorbiacea MEXICO: Oaxaca: Camotlan, A.M.N.H. (2); Canon Tlatixtac, 6.4 kilometers E. Oaxaca, A.M.N.H. (1); Cerro San Felipe, U.I.M.N.H. (26); South slope of Cerro Machin, K.U. (1 tadpoles); El Punto, A.M.N.H. (1 tadpoles), 1.6 kilometers north of El Punto, A.M.N.H. (3, 1 tadpoles); 3 kilometers north of El Punto, A.M.N.H. (2); Guelatao, U.M.M.Z. (1); 25 kilometers north of Guelatao, K.U. (1); Ixtlan de Juarez, U.M.M.Z. (91); 16 kilometers .south of I.xdan de Juarez, U.I.M.N.H. (5); 17 kilometers south of 702 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 Ixtlan de Juarez, U.M.M.Z. (9); 20 kilometers south of Ixtlan de Juarez, U.I.M.N.H. (26); 27 kilometers south of Ixtlan de Juarez, U.M.M.Z. (5); 29 kilometers .south of Ixtlan de Juarez, U.I.M.N.H. (62); Lachigola, A.M.N.H. (27); Llano de las Flores, A.M.N. H. (4, 1 skeleton), K.U. (89, 10 skeletons), U.I.M.N.H. (19), U.M.M.Z. (87); Oa-xaca, A.M.N.H. (1); U.I.M.N.H. (3), U.M.M.Z. (18), U.S.N.M. (1); 5 kilometers northeast of Oaxaca, U.F. (5); 1.6 kilometers east of Oaxaca, U.I.M.N.H. (42); 2.7 kilometers southeast of Oaxaca, U.M.M.Z. (16); 4 kilometers southeast of Oaxaca, K.U. (12, 1 tadpoles, 1 eggs); 5 kilometers southeast of Oa.xaca, U.M.M.Z. (11); 6 kilometers southeast of Oaxaca, K.U. (12); 8 kilometers southeast of Oaxaca, K.U. (49, 3 skeletons), U.M.M.Z. (7); 15 kilometers southeast Oaxaca, U.M.M.Z. (7); 4 kilo- meters west of Oaxaca, U.M.M.Z. (7); 14.4 kilometers northwest of Oaxaca, U.M.M.Z. (6); between Oaxaca and Tlacolula, U.I.M.N.H. (4); San Andres Chica- huasda, U.I.M.N.H. (1); 3 kilometers northeast of San Andres Chicahuastla, U.M.M.Z. (27); San Felipe, A.M.N.H. (1); 1.6 kilometers south of San Felipe Ixtapa, U.I.M.N.H. (3); 3 kilometers east of Santa Lucia, A.M.N.H. (10); Santo Domingo Ocotepec, U.I.M.N.H. (4); San Vincente Laehixio, K.U. (1); 4 kilometers southeast of Tlacolula, T.N.H.C. (8); 15 kilometers north of Tlaxiaco, K.U. (1 tadpoles). Puchla: Paraje Verde, U.I.M.N.H. (3); U.M.M.Z. (1); Puente Colorado, U.M.M.Z. (2). Veracruz: Cumbres de Acultzingo, I.P.N. (1), K.U. (5), M.C.Z. (1), U.I.M.N.H. (17), U.M.M.Z. (19), U.S.N.M. (11). GUATEMALA: Aha Vcrapaz: Coban, F.M.N.H, ( 3 ) ; Finca Samac, U.M.M.Z. ( 1 ) . Hyla eximia MEXICO: Aguascalientes: 13 kilometers east of Aguascalientes, F.M.N.H. (10), U.I.M.N.H. (15); 16 kilometers east of Aguascalientes, F.M.N.H. (20), U.I.M.N.H. ( 18); 4 kilometers ea.st of Caiiada Honda, S.U. (1); 8 kilometers north of Rincon de Romos, U.M.M.Z. (1); 30 kilometers .south of Rincon de Romos, U.I.M.N.H. (2). Chihuahua: No specific locality, M.C.Z. (1); Colonia Garcia, U.S.N.M. (3); 26 kilometers south, 21 kilometers west of Creel, K.U. (2); 37 kilometers south, 2.5 kilometers east of Creel, K.U. (16); Meadow Valley, U.S.N.M. (6); Mojara- chic, F.M.N.H. (1), U.I.M.N.H. (2), U.M.M.Z. (1); 3 kilometers west of Samachique, K.U. (1); 3 kilo- meters southwest of San Jose de Babicora, K.U. (22); 5 kilometers northeast of Temoris, K.U. (1). Distrito Federal: Atzacualco, U.S.N.M. (1); Ciudad Me.xico, I.P.N. (1), U.S.N.M. (2); Contreras, U.M.M.Z. (1); Guadalupe, A.M.N.H. (15); Lago Texcoco, U.M.M.Z. (7); 1.6 kilometers west of Nalpani, A.M.N.H. (37); Parque Cliapultepec, A.M.N.H. (1); Pedregal de San Angel, I.P.N. (16); San Juanico, A.M.N.H. (1); San Luis, 10 kilometers east of Xochimilco, U.M.M.Z. (4); San Mateo Chalpa, A.M.N.H. (1); 1.6 kilometers .southwest of Ticoman, A.M.N.H. (4); Tlalpam, F.M.N.H. (2); Valle de Me.xico, U.S.N.M. (1); Xochimilco, A.M.N.H. (15), A.N.S.P. (3), I.P.N. (7), U.M.M.Z. (1); 1.6 kilometers southwest of Xochimil- co, U.M.M.Z. (I). Durango: Buenos Aires, 6 kilome- ters southwest of La Ciudad, A.M.N.H. (29), K.U. (1 tadpoles); Coyotes, F.M.N.H. (1), U.M.M.Z. (4); Cerro Huehuento, 40 kilometers north-northwest of El Salto, U.M.M.Z. (1); Durango, M.C.Z. (1), U.M.M.Z. (12); Rio Mezquital, 16 kilometers north- of Durango, U.I.M.N.H. (21); 24 kilometers north of Durango, U.I.M.N.H. (3); 20 kilometers east of Durango, LI.M.M.Z. (5); 96 kilometers west of Dur- ango, M.C.Z. (2); 14 kilometers east of El Espinosa, L.B.S.C. (1); El Salto, A.N.S.P. (2), K.U. (2); 4 kilometers northeast of El Salto, U.I.M.N.H. (33); 16 kilometers northeast of El Salto, U.I.M.N.H. (3); 2 kilometers southwest of El Salto, I.P.N. (8); 11.3 kilometers southwest of El Salto, U.I.M.N.H. (1); 14 kilometers southwest of El Salto, K.U. (8), U.I.M.N.H. (3); 16 kilometers southwest of El Salto, C.A.S. (1), K.U. (17); 23 kilometers southwest of El Salto, U.M.M.Z. (7); 44.3 kilometers southwest of El Salto, K.U. (3); 49 kilometers southwest of El Salto, U.I.M.N.H. (1); 53 kilometers southwest of El Salto, L.B.S.C. (46); Laguna del Progreso, U.M.M.Z. (20); 7 kilometers northeast of Las Adjun- tas, U.I.M.N.H. (1); 4 kilometers southwest of Las Adjuntas, U.I.M.N.H. (4); 19 kilometers southwest of Las Adjuntas, K.U. (13); 8 kilometers north of Morcillo, U.M.M.Z. (3); Pueblo Nuevo, A.N.S.P. (2), U.M.M.Z. (12); Rio Mezquital. 16 kilometers north- east of Durango, U.I.M.N.H. (1). Guanajuato: No specific locality, U.S.N.M. (3); Acambaro, F.M.N.H. (4); Celaya, F.M.N.H. (1); 11 kilometers northwest of Leon, U.I.M.N.H. (18); 10 kilometers west of Panjamo, U.I.M.N.H. (2); Silao, U.S.N.M. (1); 22 kilometers south of Valle de Santiago, U.I.M.N.H. (63). Guerrero: 4 kilometers southwest of Almolon- ga, K.U. (3). T.C.W.C. (20), U.M.M.Z. (6); East of Chilapa, K.U. (1). Chilpancingo, M.C.Z. (1); 22 kilometers south of Chilpancingo, C.A.S. (1); 22 kilo- meters south of Ixtapan de la Sal (Mexico), K.U. (17); Omiltemi, M.C.Z. (1), T.C.W.C. (5); U.I.M.N.H. (10), U.S.N.M. (3); 3 kilometers north of Omiltemi, T.C.W.C. (1); 6 kilometers nortli of Omiltemi, U.S.N.M. (2); 10 kilometers east of Omil- temi, F.M.N.H. (3); 3 kilometers west of Omiltemi, T.C.W.C. (9); Tixtla, F.M.N.H. (1), U.I.M.N.H. (1). Hidalgo: 10 kilometers east of Aca.xochitlan, .\.M.N.H. (1); 5.6 kilometers west Acaxochitlan, U.M.M.Z. (6); Agua Blanca, near Apulca, U.M.M.Z. (1); 16 kilometers north of Agua Blanca, U.M.M.Z. (6); 16 kilometers south of Apulca, U.M.M.Z. (15); Atotonilco Grande, F.M.N.H. (32), U.I.M.N.H. (24); El Chico Parque Nacional, A.M.N.H. (8); Guerrero, M.C.Z. (3); 20 kilometers west-southwest of Huachin- ango (Puebla), U.M.M.Z. (2); Jacala, U.M.M.Z. (2); Lago Tejocotal, I.P.N. (1), U.M.M.Z. (1); 2.5 kilo- meters southwest of Tianguistengo, K.U. (2); 6 kilo- meters Tianguistengo, F.M.N.H. (4); 5 kilometers northwest of Tianguistengo, U.I.M.N.H. (1); 13 kilometers east-northeast of Tulancingo, U.M.M.Z. ( 1 ); 10 kilometers southwest of Tulancingo, U.M.M.Z. (2); 15 kilometers southwest of Tulancingo, U.M.M.Z. 1970 DUELLMAN: HYLID FROGS 703 (1); Velasco, A.M.N.H. (2); 6 kilometeis south of Zacualtipan, F.M.N. H. (11), U.I.M.N.H, (12). Jalisco: Agiia Delgada, 6 kilometers north of Guada- lajara, A.M.N.H. (14); 3 kilometers east of Ajijic, A.M.N.H. (1); 5 kilometers west of Arandes, K.U. (45); Atemajae, A.M.N.H. (2); Atotonilco del Alto, K.U. (4); Autlan Road, F.M.N.H. (2); 5 kilometers northeast of Autlan, F.M.N.H. (1), U.I.M.N.H. (2); 4 kilometers west of Ayo el Chico, U.I.M.N.H. (11); Cerro de la Venta, 22 kilometers west-northwest of Guadalajara, K.U. (1); Chapala, A.M.N.H. (1 tad- poles), F.M.N.H. (7), U.I.M.N.H. (10); 11.5 kilo- meters north of Chapala, U.I.M.N.H. (11); 16 kilo- meters north of Chapala, A.M.N.H. (1); 3 kilometers north of Ciudad Guzman, U.M.M.Z. (5); 5 kilometers northeast of Ciudad Guzman, F.M.N.H. (6); 1.6 kilometers west of Ciudad Guzm;in, U.M.M.Z. (2); Guarente, K.U. (1); 5 kilometers northwest of Degollado, K.U. (3); Guadalajara, A.M.N.H. (2); F.M.N.H. (3), K.U. (1); U.I.M.N.H. (1); 20 kilo- meters south, 29 kilometers west of Guadalajara, K.U. (1); 21 kilometers south of Guadalajara, U.M.M.Z. (2); 38 kilometers south of Guadalajara, U.M.M.Z. ( 1 ) ; 29 kilometers southwest of Guadalajara, U.M.M.Z. (19); 1 kilometer northwest of I.xtla- huacan, A.M.N.H. (24); 8 kilometers northwest of I.xtlahuacan, A.M.N.H. (1); 1.6 kilometers south of Jalostotitlin, K.U. (1); Janiay, A.M.N.H. (11); 3 kilometers east of Jocotepec, U.I.M.N.H. (1); 5 kilo- meters northwest of Jocotepec, A.M.N.H. (2), U.I.M.N.H. (2), U.S.N.M. (2); Lago Chapala, A.M.N.H. (2); Lagos de Morena, A.M.N.H. (1), C.A.S. (1); 13 kilometers northeast of Lagos de Morena, U.I.M.N.H. (6); 40 kilometers east of Lagos de Morena, K.U. (16, 2 skeletons, 1 tadpoles); 3 kilometers west-northwest of Lagos de Morena, K.U. (2); Laguna de Magdalena, A.M.N.H. (1); La Mesa Maria de Leon, K.U. (39); 3 kilometers northwest of Magdalena, K.U. (1), T.N.H.G. (1); 1.6 kilometers northwest of Mazaniitla, U.M.M.Z. (1); Ocotliin, A.M.N.H. (1); 21.7 kilometers west of Ojuelos, L'.I.M.N.H. (3); Rancho Primavera, near Guadala- jara, U.I.M.N.H. (1); 5 kilometers west of San Antonio, U.I.M.N.H. (1); 35 kilometers west of Soyatkin, T.G.W.C. (10); 10 kilometers north, 6 kilo- meters east of Tepatitlan, K.LI. (15); 12 kilometers northeast of Tepatitlan, U.M.M.Z. (17); TIaquepaque, A.M.N.H. (14); Tonola, A.M.N.H. (3); between Tonoki and TIaquepaque, A.M.N.H. (5); Villa Corona, north end of Lago Atotonilco, K.U. (21); Villa de Guadalupe, C.A.S. (1); 1.6 kilometers north- east of Villa Hidalgo, K.LI. (3); 11 kilometers south, 1.6 kilometers east of Yahualica, K.U. (1); Zapotiltic, F.M.N.H. (4), U.I.M.N.H. (3). Mexico: 5.1 kilo- meters south of Acolman, T.N.H.G. (3); Ameyal, L'.I.M.N.H. (5); 5 kilometers south of Bosencheve, U.M.M.Z. (32); Chalco, F.M.N.H. (1); Chapingo, I.P.N. (1); 45 kilometers west of Ciudad Me.\ico, T.C.W.C. (2); Ixtapan de la Sal, A.M.N.H. (11), F.M.N.H. (3), T.N.H.G. (11); 5 kilometers north of Ixtapan de la Sal, T.N.H.G. (2); 6 kilometers north of I.xtapan de la Sal, U.M.M.Z. (23, 2 tadpoles); Laguna Agua Buena, 27 kilometers southwest of Toluca, U.M.M.Z. (6); Lagrma de Ojuelos, 8 kilo- meters west of Toluca, A.M.N.H. (3); La Marquesa, I.P.N. (1); Lengua de Vaca, 16 kilometers east of Zitacuaro (Michoacan), U.M.M.Z. (1); Lerma, F.M.N.H. (2), U.I.M.N.H. (2); Nevada de Toluca, F.M.N.H. (2), U.I.M.N.H. (2); Rancho Guadalupe, 51 kilometers west of Toluca, U.I.M.N.H. (1); Rio Frio, U.I.M.N.H. (1); 5 kilometers north, 11 kilome- ters west of San Jose Allende, K.U. (1); San Juan Teotihuacan, M.C.Z. (1); 16 kilometers from San Martin, F.M.N.H. (1); U.LM.N.H. (1); 8 kilometers south of Tenancingo, K.U. (2); 5.6 kilometers south of Tenango, T.G.W.C. (10); 6 kilometers west of Tepexpan, U.M.M.Z. (15); Toluca, A.M.N.H. (1), F.M.N.H. (1), U.S.N.M. (1); 3 kilometers west of Toluca, F.M.N.H. (10); 24-32 kilometers west of Toluca, U.M.M.Z. (1); 10 kilometers north-northwest of Toluca, F.M.N.H. (3); Tonatico, I.P.N. (1); 1.6 kilometers south of Valle de Bravo, K.U. (4); 19 kilo- meters west of Villa Victoria, U.S.N.M. (4). Michoa- can: 11 kilometers west of Ciudad Hidalgo, U.M.M.Z. (36); Cojumatlan, F.M.N.H. (1), U.I.M.N.H. (1); 6 kilometers north of Copuyo, T.C.W.C. (4); 6 kilo- meters south of Cuitzeo, U.M.M.Z. (1); 30 kilometers north of Jacona, U.I.M.N.H. (1); Jiquilpan, U.I.M.N.H. (1), U.M.M.Z. (1); 8.3 kilometers east- southeast of Jiquilpan, T.N.H.G. (1); Lago de Game- cuaro, U.M.M.Z. (1); Lago de Patzcuaro, A.M.N.H. (7), F.M.N.H. (2), U.I.M.N.H. (1); 2.5 kilometers .south of Los Reyes, K.U. (1); Morelia, F.M.N.H. (1); 11 kilometers west of Morelia, A.M.N.H. (9); Patz- cuaro, A.M.N.H. (6); 10 kilometers north of Patzcuaro, U.I.M.N.H. (92); 3 kilometers northeast of Patzcuaro, T.N.H.G. (1); 8 kilometers northeast of Patzcuaro, U.M.M.Z. (6); 5 kilometers south of Patzcuaro, C.A.S. (1); 25 kilometers south of Patz- cuaro, U.M.M.Z. (3); Sahuayo, U.S.N.M. (3); 5 kilo- meters west Tangamandapio, LI.M.M.Z. (4); Temaz- cal, I.P.N. ( 1 ); 3 kilometers west Temazcal, U.M.M.Z. (36); Tupataro, U.S.N.M. (1); Tu.xpan, U.M.M.Z. (15), 25 kilometers west of Tu.xpan, U.I.M.N.H. (3); bet%veen Tzintzuntzan and Patzcuaro, L'.M.M.Z. (25); Undameo, U.M.M.Z. (2); Uruapan, F.M.N.H. (6), U.I.M.N.H. (4); Villamor, A.M.N.H. (2); Zacapu, U.I.M.N.H. (1); Zamora, C.A.S. (1), F.M.N.H. (2), U.I.M.N.H. (1); 1.6 kilometers southeast of Zamora, T.N.H.G. (2); 14.4 kilometers east of Zamora, U.M.M.Z. (22); 1.6 kilometers northeast of Zinape- cuaro, K.U. (10). Morelos: 18 kilometers southeast of Cuautla, T.N.H.G. (4); 1.6 kilometers northwest of Cuautlixco, L'.M.M.Z. (1); 5 kilometers northwest of Cuautlixco, U.M.M.Z. (5); 3.5 kilometers west of Cuautlixco, (3); Cuernavaca, F.M.N.H. (2), T.C.W.C. (9), U.I.M.N.H. (5); 2.7 kilometers east of Cuernavaca, T.N.H.G. (1); 5.6 kilometers south of Cuernavaca, F.M.N.H. (1); 2 kilometers south of Jonacatepee, T.C.W.C. (9); Progreso, T.C.W.C. (30), U.F. (10); Temisco, F.M.N.H. (1), U.I.M.N.H. (2); Tepoztlan, F.M.N.H. (3). Nayarit: 3 kilometers south of Acaponeta, U.M.M.Z. ( 1 ); Arroyo de Rifilion, 9 kilometers nordi of Compostela, L.B.S.C. (1); 13 kilometers north of Compostela, L.B.S.C. (1); 3 kilo- meters south of Compostela, K.U. (1); Ixtliin del Rio, 704 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 U.M.M.Z. (1); 3 kilometers northwest of Ixtlan del Rio, T.C.W.C. (3); Laguna San Pedro, 16 kilometers east of Conipostela, A.M.N.H. (3); Petaquilla, A.M.N.H. (4); Rio San Cayetano, 5.6 kilometers southeast of Tepic, A.M.N.H. (2); Santa Teresa, U.S.N.M. (4); Tepic, F.M.N.H. (6); S.U. (2), U.I.M.N.H. (3), U.M.M.Z. (2 tadpoles), U.S.N.M. (2); 19 kilometers southeast of Tepic, K.U. (2); 22 kilometers southeast of Tepic, A.M.N.H. (2); 8.6 kilo- meters south-southeast of Tepic, U.M.M.Z. (14); 32 kilometers south-southeast, 6 kilometers east of Tepic, L.B.S.C. (1); 37 kilometers south-southeast of Tepic, L.B.S.C. (3); 6 kilometers south of Tepic, L.B.S.C. (1); 8 kilometers south of Tepic, L.B.S.C. (2); 8,8 kilometers south of Tepic, A.M.N.H. (1); 10 kilome- ters south of Tepic, L.B.S.C. ( 12); 35 kilometers south of Tepic, L.B.S.C. (3). Fuebla: 2 kilometers south of Ahuazotepec ,U.LM.N.H. (5); 6 kilometers south of Amazoc, F.M.N.H. (13), U.I.M.N.H. (10); 3 kilo- meters north of Cholula, F.M.N.H. (10), U.I.M.N.H. (11); 17 kilometers southeast of Huachinango, T.C.W.C. (1); 16 kilometers southwest of Huachin- ango, A.M.N.H. (1); 5.6 kilometers west of Huachin- ango, U.M.M.Z. ( 17, 1 tadpoles); 10 kilometers south- west of Iziicar de Matamoros, K.U. (1); Los Molinos, U.M.M.Z. (2); Fuebla, A.M.N.H. (1), U.S.N.M. (1); 11 kilometers south, 5 kilometers east of Fuebla, K.U. (2); 1.6 kilometers southwest of Fuebla, U.M.M.Z. (19); 4.2 kilometers southwest of Fuebla, U.M.M.Z. (3); 6 kilometers southwest of Fuebla, U.M.M.Z. (21); Reyes, near Santa Catarina, A.M.N.H. (1); Santa Catarina, A.M.N.H. (1); Tepeaca, F.M.N.H. (1), U.I.M.N.H. (2); 9 kilometers south of Tepeojuma, U.M.M.Z. (1); Tezuitlan, U.S.N.M. (1). Queretaro: 4 kilometers north of Quere- taro, U.I.M.N.H. (1); 13 kilometers northw^est of Queretaro, K.U. (20); 1.6 kilometers south of Santa Rosa, U.I.M.N.H. (2). San Luis Potosi: Alvarez, M.C.Z. (2), mountains north of Alvarez, M.C.Z. (1); 38 kilometers east of San Luis Potosi, U.M.M.Z. (12). Tamaulipas: La Joya de Salas, U.M.M.Z. (43). Tlaxcala: Apizaco, U.I.M.N.H. (20), U.M.M.Z. (18); Atlihuetzia, 10 kilometers northeast of Tlaxcala, U.I.M.N.H. (4); 1 kilometer northwest of Ocoto.xco, U.I.M.N.H. (3); 5 kilometers northwest of Sanctori- um, T.N.H.C. (3); 3 kilometers south of San Ildefonso Hiieyotlipan, U.I.M.N.H. (1). Veracruz: Banderilla, U.I.M.N.H. (1); Cuautlapan, K.U. (2); Jacales, K.U. (2); 10 kilometers southwest of Jacales, K.U. (69); Las Vigas, U.M.M.Z. (1); Mirador, U.S.N.M. (3); Tierra Colorada, U.I.M.N.H. (1); PVolcan San Martin, U.I.M.N.H. (2); Xuchil, F.M.N.H. (I). Zacatecas: 16.6 kilometers northwest of Jalpa, K.U. (2); 9.6 kilometers east of Monte Escobedo. K.U. (2); 5 kilometers northwest of Teul, U.M.M.Z. ( II ). Hyla fimbrimembra COSTA RICA: Alajuela: Cinchona, R.C.T. (1). Heredia: Isla Bonita, R.C.T. ( 1 ). Hyla godmani MEXICO: Puchla: Maria Andrea, A.M.N.H. (8); 1 0 kilometers southwest of Mecatepec ( Veracruz ) , U.I.M.N.H. (13). Veracruz: 5 kilometers east-south- east of Cordoba, A.M.N.H. (7), K.U. (2 tadpoles), T.N.H.C. (10); 7.2 kilometers east-southeast of Cor- doba, K.U. (1); U.M.M.Z. (22); 6.4 kilometers east of Encero, A.M.N.H. (1), F.M.N.H. (1), S.U. (1), U.I.M.N.H. (18); Jalapa, U.M.M.Z. (2); 23 kilo- meters southeast of Jalapa, K.U. (13, 2 skeletons), 2 kilometers east-northeast of Mata Oscura, K.U. (34, 13 skeletons); Mirador, U.M.M.Z. (1); Misantla, B.M.N.H. (1); Falma Sola, U.S.N.M. (20); Potrero Vicjo, F.M.N.H. (5), M.C.Z. (14), U.I.M.N.H. (48), U.M.M.Z. (16); 37 kilometers west of Fosa Rica, U.M.M.Z. (3). Hyla hazelae MEXICO: Oaxaca: Canon Tlalixtac, 6 kilometers southeast, 17 kilometers northeast of Oaxaca, K.U. (1); Cerro Machin, U.I.M.N.H. (3); Cerro San Felipe, F.M.N.H. (4), U.S.N.M. (2); 3 kilometers east of Ixtldn de Juarez, A.M.N.H. (1); 13 kilometers northwest of Ixtlan de Juarez, T.N.H.C. (4) near Oaxaca, U.I.M.N.H. (2); 2 kilometers south of E. Punto, K.U. (2, 1 skull). Hyla lancasteri COSTA RICA: Cartago: 7.4 kilometers west of Juan Vina, U.M.M.Z. (1); Moravia, K.U. (46, 3 skeletons), U.S.C. (2); 3 kilometers south of Pavones, K.U. (65, 2 skeletons, 3 tadpoles). U.M.M.Z. (5), U.S.C. (7); Peralta, M.C.Z. (1); Rio Chitaria, 3 kilo- meters north-northeast of Pavones, K.U. (2), M.C.Z. (1), U.S.C. (5); Turrialba, K.U. (9, 1 skeleton), M.C.Z. (2); south slope of Volcan Turrialba, U.M.M.Z. (5 tadpoles). Limon: El Tigre, 9-14 kilo- meters southwest of Siguirres, U.S.C. (4). F.\NAMA: Bocas del Toro: North slope of Cerro Pando, 1450 meters, K.U. (21, 2 skeletons, 3 tadpoles, 1 eggs); north slope of Cerro Pando, 1810 meters, K.U. (2); north slope of Cerro Pando, 1920 meters, K.U. (9, 1 eggs); Rio Changena, 650 meters, K.U. (2); Rio Changena, 830 meters, K.U. (8); Rio Clara near junction v\ith Rio Changena, 910 meters, K.U. (3, 4 tadpoles, 1 eggs). Hyla legleri COSTA RICA: Puntarcnas: Finca El Helechales, 15 kilometers northeast of Potrero Crande, U.S.C. (2); Finca Loma Linda, 2 kilometers south-southwest of Canas Gordas, U.M.M.Z. (4), U.S.C. (20). San Jose: south slope Cerro de la Muerte, 1540 meters, U.S.C. (3); 14 kilometers north of San Isidro el General, K.U. (2, 3 skeletons), U.M.M.Z. (I); 15 kilometers north of San Isidro el General, K.U. (2, 1 tadpoles); 15 kilometers west-southwest of San Isidro el General, K.U. (12, 5 tadpoles), U.S.C. (6), U.M.M.Z. (1). PANAMA: Chiriqui: Finca Santa Clara, K.U. (3, 1 tadpoles ) . 1970 DUELLMAN: HYLID FROGS 705 Hyla loquax MEXICO: Campeche: Lagiina Alvarado, 65 kilo- meters south of Xpujil, K.U. (21); 10 kilometers east of Lasima Silvitiic, K.U. (2); Tres Brazos, F.M.N.H. (1), U.I.M.N.H. (1). Chiapas: El Suspiro, U.M.M.Z. (1)'; Laguna Ocotal, M.C.Z. (1); 16 kilometers south of Teapa (Tabasco), U.I.M.N.H. (6). Oaxaca: 3.7 kilometers north of Donaji, U.M.M.Z. (7); 43 kilome- ters north of Matias Romero, U.I.M.N.H. (21); Rio Chicapa, near El Atravesado, A.M.N. H. (1). Tabasco: Teapa, T.N.H.C. (4); 12.4 kilometers north of Teapa, U.M.M.Z. (2); 27 kilometers north of Teapa, U.M.M.Z. (10); 3.5 kilometers south of Villahermosa, U.M.M.Z. (16). Veracruz: 19 kilometers north of Acayucan, U.I.M.N.H. (4); 4.5 kilometers south of Aquilera, U.M.M.Z. (3); Cuautotlapam, U.M.M.Z. (3); 8 kilometers southwest of Coatzacoalcos, U.M.M.Z. (36); Encinal, U.M.M.Z. (4); 5 kilometers west of Juan Diaz Covarrubias, U.M.M.Z. (8); 8 kilometers south of Lago Catemaco, U.I.M.N.H. (1); 4 kilometers northeast of Minatitlan, T.N.H.C. (I); 2 kilometers south of Naranja, U.M.M.Z. (12); San Lorenzo, U.S.N. M. (3). BRITISH HONDURAS: Cayo: 2 kilometers southwest of Cayo, U.M.M.Z. (5); Hummingbird Highway, 16 kilometers from Belize-Cayo road, U.M.M.Z. (6); Pine Ridge Road, 20.3 kilometers from Belize-Cayo road, U.M.M.Z. (2); Pine Ridge Road, 57-58 kilometers from Belize-Cayo road, U.M.M.Z. ( 1 ) ; San Augustine, U.M.M.Z. ( 1 ) . GUATEMALA: Aha Verapaz: Finca Chama, C.A.S. (1), U.M.M.Z. (85). El Peten: No specific locality, U.S.N.M. (1); La Libertad, F.M.N.H. (2), K.U. (17, I skeleton), M.C.Z. (2), U.I.M.N.H. (1), U.M.M.Z. (8), U.S.N.M. (1); Piedras Negras, F.M.N.H, (3), U.I.M.N.H. (1), U.S.N.M. (1); 8 kilo- meters south of Piedras Negras, U.S.N.M. (32); Tikal, U.M.M.Z. (3); Toocog, 15 kilometers southeast of La Libertad, K.U. (9, 5 skeletons, 1 eggs). Izabal: Puerto Barrios, F.M.N.H. (2); 8 kilometers south of Puerto Barrios, K.U. (9). HONDURAS: Francisco Morazan: Cerro de Guaimaca, U.M.M.Z. (1); Cerro Uyuca, A.M.N.H. (1), U.M.M.Z. (2). Yoro: Subirana Valley, F.M.N.H. (1), M.C.Z. (4). NICARAGUA: Boaco: 14 kilometers north, 13 kilometers east of Boaco, K.U. (7). Esteli: Finca Daraili, 5 kilometers north, 14 kilometers east of Con- dega, K.U. (2); Finca Venecia, 7 kilometers east of Condega, K.U. (6). Matagalpa: 14.4 kilometers southeast of Jinotega, K.U. (1); 19 kilometers north of Matagalpa, U.M.M.Z. (10); Santa Maria de Ostuma, K.U. (1). Zelaya: Isla Grande del Maiz, K.U. (5). COSTA RICA: Cartago: EI Silencio, 14.4 kilo- meters northeast of Turrialba, K.U. (2); Moravia de Turrialba, K.U. (13, 1 skeleton, 1 tadpoles, I eggs), U.S.C. (25); 11 kilometers southwest of Moravia, K.U. (2); 1 kilometer east-northeast of Pacuare, K.U. (17); Peralta, K.U. (5); Tuis, K.U. (I); Turrialba, F.M.N.H. (12), K.U. (48, 3 skeletons), U.S.C. (9), L'.M.M.Z. (5). Guanacaste: Finca San Bosco, U.S.C. (21); Silencio, U.S.C. (14); Tilaran K.U. (8), 2 kilometers east of Tilaran, K.U. (27, 2 skeletons); 4 kilometers east of Tilaran, U.S.C. (4); 8 kilometers northeast of Tilaran, K.U. (15). Heredia: Finca La Selva, U.S.C. (3); Puerto Viejo, K.U. (3, 3 skeletons). Hyla melanomma bivocata MEXICO: Chiapas: ,32 kilometers north of Jitotol, U.I.M.N.H. (3); 18 kilometers north of Pueblo Nuevo Solistahuacan, K.U. (I), U.M.M.Z. (4); 5.6 kilome- ters south of Rayon Mescalapa, K.U. (1, 1 skeleton); 6.2 kilometers south of Rayon Mescalapa, K.U. (5). Hyla melanomma melanomma MEXICO: Guerrero: Acahuitzotla, T.C.W.C. (3); Agua del Obispo, A.M.N.H. (3), K.U. (1, 1 tadpoles); 11 kilometers east of Chilpancingo, F.M.N.H. (17), M.C.Z. (1), U.I.M.N.H. (10), U.M.M.Z. (1), U.S.N.M. (6); 22 kilometers south of Chilpancingo, C.A.S. (8). Oaxaca: 29 kilometers south-southeast of Juchatengo, K.U. (I); 56 kilome- ters north of Pochutla, U.M.M.Z. (1); 62 kilometers north of Pochutla, U.M.M.Z. (6); 6 kilometers north- northwest of San Gabriel Mixtepec, K.U. (1, 1 tad- poles); 12 kilometers north-northwest Mixtepec, K.U. (16,2 skeletons). Hyla microcephala microcephala COSTA RICA: Puntarcnas: Golfito, K.U. (36); 3 kilometers east of Golfito, K.U. (1), U.S.C. (2); Pal- mar Sur, K.U. (24, 5 skeletons), U.S.C. (14), U.U. (26); 3 kilometers northwest of Piedras Blancas, K.U. (11); Villa Neilly, U.S.C. (12); 10.5 kilometers west- northwest of Villa Neilly, K.U. ( 19, I eggs). PANAMA: Canal Zone: Albrook Air Base, T.N.H.C. (2); Balboa, A.N.S.P. (2); Fort Clayton, U.I.M.N.H. (5); 2.8 kilometers southwest of Fort Kobbe, K.U. (11); Frijoles, M.C.Z. (I); Gamboa M.C.Z. (1); 8.3 kilometers north of Gatun Locks, T.N.H.C. (1); Juan Diaz, M.C.Z. (1); Juan Mina, A.M.N.H. (2), A.N.S.P. (2), U.M.M.Z. (7), U.U. (7); Madden Dam, F.M.N.H. (1); 8-14 kilometers north of Miraflores Locks, T.N.H.C. (116); Rio Chagres, A.M.N.H. (2); Rio Cocoli, 3.5 kilometers north of Miraflores Locks, T.N.H.C. (1); Summit, A.N.S.P. (7), F.M.N.H. (4), K.U. (5). Chiriqui: 5.5 kilometers east of Concepcion, A.M.N.H. ( I ); 14.4 kilometers east of Concepcion, A.M.N.H. (6); 2 kilo- meters south of David (1); Progreso, U.M.M.Z. (5); Rio Gariche, 8.3 kilometers east-southeast of Paso Canoas, K.U. (4). Code: 1 kilometer southeast of El Caiio, K.U. (10); El Valle de Anton, A.M.N.H. (II), A.N.S.P. (4), K.U. (14), M.V.Z. (6), U.I.M.N.H. ( 1 ) . Colon: Cement Plant, Transisthmian Highway, F.M.N.H. (1). Darien: El Real, K.U. ( 15), U.M.M.Z. (10), U.S.N.M. (2); Rio Canclon at Rio Chucunaque, U.M.M.Z. (1); Rio Chucunaque, near Yavisa, A.M.N.H. (1). Los Santos: Tonosi: K.U. (4). Panama: 5 kilometers south of Bejuco, A.M.N.H. (1); 706 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 3 kilometers west of Chepo, K.U. (3, 2 tadpoles); 6 kilometers west-southwest of Chepo, K.U. (1); Chico, Rio La Jagua, U.S.N.M. (1); La Joya, A.M.N. H. (5); Nueva Gorgona, A.M.N.H. (2); L6 kilometers west of Nueva Gorgona, A.M.N.H. (1); 9 kilometers north- east of Pacora, K.U. (1); 1.5 kilometers west of Pacora, K.U. (25); Panama, K.U. (1); Rio La Laja, near Chame, A.N.S.P. ( 1 ); Rio Tapia, A.M.N.H. (4); 18 kilometers east of Tocumen, M.V.Z. (1). Vera- guas: Rio Coroba, U.S.N.M. (1). Hyla m. microcephala .\ underwoodi COSTA RICA: Puntarciias: Parrita, U.S.C. (9); 6.1 kilometers northeast of mouth of Rio Tarcoles, U.S.C. (4). Hyla microcephala underwoodi MEXICO: Campeche: Balchacaj, F.M.N. H. (1), U.I.M.N.H. (3); Encarnacion, F.M.N.H. (3), M.C.Z. (2), U.I.M.N.H. (12), U.S.N.M. (2); Escarcega, U.M.NLZ. (1); 7.5 kilometers west of Escarcega, K.U. (15); Lagima Alvarado, 65 kilometers south of Xpujil, K.U. (6); Pacaitiin, Rio Candelaria, F.M.N.H. (3); Tres Brazos, F.M.N.H. (22), U.I.M.N.H. (1); 10 kilometers west of Xpujil, K.U. (2). Chiapas: Palenque, U.I.M.N.H. (13), U.S.N.M. (6). Oaxaca: 5 kilometers north of Chiltepec, K.U. (9); 3 kilometers north of Donaji, U.M.M.Z. (14); 43 kilometers north of Matias Romero, U.I.M.N.H. (19); 3.5 kilometers north of Palomares, T.N.H.C. (40); 4.6 kilometers north of Sarabia, U.M.M.Z. (2); 6.1 kilometers north of Sarabia, U.M.M.Z. (11); 3 kilometers north of Tolo- cita, K.U. (1); Tuxtepec, K.U. (17); 3 kilometers south of Tuxtepec, U.I.M.N.H. (100). Tabasco: 24 kilometers north of Frontera, M.C.Z. (6), U.I.M.N.H. (4); 0.8 kilometers east of the Rio Tonala, T.N.H.C. (1); Teapa, U.M.M.Z. (4); 2.7 kilometers north of Teapa, U.M.M.Z. (4); 10 kilometers north of Teapa, U.M.M.Z. (6); 11.5 kilometers north of Teapa, U.M.M.Z. (I); 15.2 kilometers north of Teapa, U.M.M.Z. (4); 17.6 kilometers north of Teapa, U.M.M.Z. (12); 3.3 kilometers south of Villahemiosa, U.M.M.Z. (12); 17.6 kilometers south of Villaher- mosa, U.M.M.Z. (12). Veracruz: 2.1 kilometers north of Acayucan, U.I.M.N.H. (3); 6.4 kilometers northwest of Acayucan, U.M.M.Z. (14); 1.6 kilometers east-southeast of Alvarado, U.M.M.Z. (39); 2.4 kilo- meters east-southeast of Alvarado, U.M.M.Z. (2); 4.5 kilometers south of Aquilera, U.M.M.Z. (21); 8 kilo- meters southwest of Coatzacoalcos, U.M.M.Z. (10); 13 kilometers north of Corral Nuevo, U.I.M.N.H. (7); 2.2 kilometers east of Cosaleacaque, U.M.M.Z. (26); 10 kilometers .southeast of Hueyapan, U.M.M.Z. (1); 0.8 kilometer south of Lerdo de Tejada, U.M.M.Z. (1); 3.6 kilometers northeast of Minatitlan, T.N.H.C. (3); 1.9 kilometers south of Naranja, U.M.M.Z. (3); 4.5 kilometers northeast of Novillero, U.M.M.Z. (I); San Andres Tu.xtla, F.M.N.H. (5), U.I.M.N.H. (2). Yucatan: Chichen-Itza, F.M.N.H. (1), M.C.Z. (2). BRITISH HONDURAS: Cayo: 6.2 kilometers south of El Cayo, M.C.Z. (8). Stann Creek: Stann Creek, F.M.N.H. (1). GUATEMALA: Alia Verapaz: 28.3 kilometers north of Campur, K.U. (13); Chinaja, K.U. (1); Cubilquitz, U.M.M.Z. (5); Finca Chama, U.M.M.Z. (163); Finca Tinaja, B.Y.U. (1); Panzos, U.M.M.Z. (2). Chiqiiimula: Chiquimula, U.M.M.Z. (1); 2 kilo- meters north of Esquipulas, U.M.M.Z. ( 1 ). El Peten: La Libertad, K.U. (51, 5 skeletons), M.C.Z. (1), U.M.M.Z. (48); Piedras Negras, F.M.N.H. (1), U.I.M.N.H. (1); 5 kilometers south of Piedras Negras, U.S.N.M. (22); Tikal, U.M.M.Z. (2); Toocog, 15 kilo- meters southeast of La Libertad, K.U. (21). El Quiche: Finca Tesoro, U.M.M.Z. (5). Huehuete- nango: Finca San Rafael, 16 kilometers southeast of Barillas, F.M.N.H. (3). Izahal: Murcielago, Lago Izabal, U.I.M.N.H. (5); Puerto Barrios, F.M.N.H. (4); 8 kilometers south of Puerto Barrios, K.U. (31, 1 eggs, 1 tadpoles); Quirigua, C.A.S. (45); 2.5 kilometers northeast of Rio Blanco, K.U. (1); San Felipe, F.M.N.H. (1). Zacapa: 14 kilometers east- northeast of Mayuelas, K.U. (5); 8 kilometers east- northeast of Rio Hondo, K.U. (4). HONDURAS: Atlantidad: La Ceiba, U.M.M.Z. (2), U.S.N.M. (8); Lancetilla, M.C.Z. (1). Cortes: Lago Yojoa, A.M.N.H. (5), K.U. (15). El Paraiso: Valle de Janiastran, A.M.N.H. (6). F rancisco-Mora- zdn: El Zamorano, A.M.N.H. (9), K.U. (1), U.M.M.Z. (1); Montana de Guaimaca, A.M.N.H. (8); Rancho San Diego, 19 kilometers southwest of Guai- maca, A.M.N.H. (1). Itibucd: Vieja Itibuca, A.M.N.H. (2). NICARAGUA: Boaco: 14 kilometers north, 13 kilometers east of Boaco, K.U. (19). Chontalcs: 11.7 kilometers east of Santo Tonuis, K.U. ( 1 ); 3 kilometers southwest of Santo Tomas, K.U. (10, 1 skeleton). Esteli: Finca Venecia, 7 kilometers north, 16 kilome- ters east of Condega, K.U. (1); 2.4 kilometers north of Esteli, M.C.Z. (5). Managua: 12-13 kilometers east of Managua, K.U. (5); 10 kilometers southwest of Tipitapa, U.M.M.Z. (10). Matagalpa: Finca Tepeyac, 10.5 kilometers north, 9 kilometers east of Matagalpa, K.U. (2); Hacienda La Cumplida, K.U. (17, 3 skeletons), U.M.M.Z. (42); 14.4 kilometers southeast of Jinotega, K.LI. (1). Nueva Segovia: 5 kilometers north, 2.5 kilometers east of Jalapa, K.U. (38). Rivas: Finca Amayo, 13 kilometers south, 14 kilometers east of Rivas, K.U. (4); 16 kilometers south of Rivas, M.C.Z. (7); 20.5 kilometers southeast of Rivas, K.U. (3); 5 kilometers southeast of San Pablo, K.U. (4). Zelaija: El Recreo, K.U. (1). COSTA RICA: Guanacaste: Arenal, U.S.C. (2); 3 kilometers west of Bagaces, U.S.C. (10); Finca San Bosco, U.S.C. (9); Guayabo de Bageces, U.S.C. (8); 12 kilometers south of La Cruz, U.S.C. (2); Laguna Arenal, U.S.C. (1); 27 kilometers north of Las Canas, U.S.C. (6); 16 kilometers east of Las Caiias, K.U. (7); 16 kilometers south-southeast of Las Canas, K.U. (6); 20 kilometers .southeast of Las Canas, K.U. (1); Liberia, U.S.C. (9); 7.3 kilometers north of Liberia, U.S.C. (4); 10 kilometers north of Liberia, U.S.C. (9); 7.5 kilometers southeast of Liberia, K.U. (7, 2 skeletons); 14.7 kilometers south of Liberia, U.S.C. 1970 DUELLMAN: HYLID FROGS 707 (3); 4 kilometers west of Liberia, K.U. (11); 2 kilo- meters south of Nicoya, U.S.C. (1); 3-10 kilometers east-southeast of Playa del Coco, U.S.C. (30); 21.6 kilometers east-southeast of Playa del Coco, U.S.C. (13); Pefias Blancas, K.U. (4); Rio Bebedero, 5 kilo- meters south of Bebedero, K.U. (1); Rio Higueron, U.S.C. (2); Santa Cruz, U.S.C. (2); Silencio, U.S.C. (1); Tenorio, K.U. (1); Tilaran, K.U. (3), 2 kilome- ters east of Tilaran, K.U. (1); 5 kilometers northeast of Tilaran, K.U. (7), U.S.C. (1). Puntamias: Bar- ranca, K.U. (8); 5 kilometers west-northwest of Bar- ranca, U.M.M.Z. (2); 3 kilometers northeast of Boca del Barranca, U.S.C. (21); 10 kilometers east of Esparta, K.U. (3, 1 tadpoles); 1 kilometer west- northwest of Esparta, K.U. (1); 4 kilometers west- northwest of Esparta, K.U. (1); 10 kilometers west-northwest of Esparta, K.U. (25, 5 skeletons); 12 kilometers west-northwest of Esparta, K.U. (5), U.S.C. ( 1 ); 21.8 kilometers west of San Ramon U.S.C. (15). Hyla miliaria NICARAGUA: No specific locaUty, U.S.N.M. (1). COSTARICA; Cartago: Turrialba, K.U. ( 1 ). PANAMA; Chiriqui: Finca Santa Clara, K.U. (2). Code: El Valla, A.M.N.H. (1). Hyla miotympanum MEXICO; Chiapas: No specific locality, U.S.N.M. (1); 16.5 kilometers north of Pueblo Nuevo Solista- huacan, U.M.M.Z. (15). Guerrero: PAcapulco, F.M.N.H. (1), U.I.M.N.H. (I). Hidalgo: 9.4 kilo- meters north of Metzquititlan, K.U. (20, 1 tadpoles); Rio Chinameca, 7.2 kilometers north of Tianguistengo, K.U. (2, 4 tadpoles); Tianguistengo, C.A.S. (1), F.M.N.H. (19), U.I.M.N.H. (9). 3 kilometers south- east of Xochicoatlan, K.U. (5, 1 tadpoles); Zacualti- pan, A.N.S.P. (1); 8.5 kilometers southeast of Zacualtipan, K.U. (1); 6 kilometers south of Zacualti pan, F.M.N.H. (12), U.I.M.N.H. (5). Nuevo Leon Hacienda Pablillo, above Galeana, A.N.S.P. (1) Paraje de los Osos, Villa de Santiago, K.U. (19); Salto Cola de Caballo, A.M.N.H. (2), A.N.S.P. (I), I.P.N (6), K.U. (9), M.V.Z. (1), T.N.H.C. (20) U.I.M.N.H, (9), U.M.M.Z. (15); Santiago, Vista Hermosa, F.M.N.H. (10); Zaragosa, K.U. (5) Oaxaca: Cerro San Felipe, F.M.N.H. (1), U.I.M.N.H (1); PTehuantepec, U.S.N.M. (3). Puebla 4.5 kilo- meters northeast of Huachinango, U.M.M.Z. (1) 14.4 kilometers west of Huachinango, K.U. (71, 4 skeletons), U.M.M.Z. (19, 1 skeleton, 1 tadpoles) 15.6 kilometers west of Huachinango, K.U. (1); 16 kilometers west of Huachinango, A.M.N.H. (2) Plziicar de Matanioros, Z.M.B. (2); Lago Necaxa U.M.M.Z. (1); Rio Frio, 10.7 kilometers north-north- east of Tezuitlan, K.U. (I, I tadpoles); Rio Octapa 3.7 kilometers north-northeast of Tezuitlan, K.U. ( 36, I tadpoles); Rio San Marcos, U.M.M.Z. (1); Rio Texcapa, 5 kilometers east of Huachinango, K.U. (3) San Diego de Tehuacan, U.M.M.Z. (14), U.S.N.M. (4). San Luis Potosi: 8 kilometers west of La Meca, T.CW.C. (2); Naranjo, F.M.N.H. (8), U.I.M.N.H. (4); Palictla, A.M.N.H. (3), M.C.Z. (18); Patilla, 8 kilometers north of Tamazunchale, U.M.M.Z. (2); Tamazunchale, U.I.M.N.H. (1), U.M.M.Z. (11); 3 kilometers north of Tamazunchale, T.CW.C. (1), U.I.M.N.H. (1); 5 kilometers north of Tamazunchale, T,C,W,C, (2); 12,4 kilometers north of Tamazun- chale, T.N.H.C. (5); 50 kilometers south of Valles, F,M,N,H, (9), U,I.M.N.H. (4); Xilitla, A.M.N.H. (1), K.U. (1), U.S.N.M. (1 tadpoles); 1.6 kilometers northwest of Xilitla, U.I.M.N.H. (1, 1 tadpoles). Tamaulipas: Acuna, U.M.M.Z. (1 tadpoles); 5 kilometers northwest of Acuiia, U.M.M.Z. (16); 2 kilometers southeast of La Joya de Salas, U,M,M,Z, (22); Las Yucas, north of Aldama, M,C.Z, (5); 5 kilometers northwest of San Jose, U,M,M,Z, (1); Santa Barbara, U.M,M,Z, (2); 1,6 kilometers north of Santa Inez, U,I,M,N,H, (2), Veracruz: Acultzingo, U,I,M,N,H, (2), U.M.M.Z. (3), U.S.N.M. (1); 8 kilometers east of Acultzingo, U.S.N.M. (15); 2 kilometers west of Acultzingo, U.I.M.N.H. (5), U.S.N.M. (15); Arroyo del Meco, near Linales, M.C.Z. (2); Banderilla, F.M.N.H. (7), U.I.M.N.H. (4); Barranca Metlac, F.M.N.H. (38), M.C.Z. (41), K.U. (1, 11 skeletons, 1 tadpoles, I eggs); U.I.M.N.H. (60), U.M.M.Z. (211, 2 tadpoles); Cordoba, F.M.N.H. (5), U.S.N.M. (2); Coscoma- tepec, K.U. (43); 5.5 kilometers north-northeast of Coscomatepec, U.M.M.Z. (3); 4.3 kilometers west of Coscomatepec, U.M.M.Z. (1); 7 kilometers west of Coscomatepec, U.M.M.Z. (1); Cuautlapan, F.M.N.H. (3), K.U. (32), U.I.M.N.H. (54), U.S.N.M. (15). Cumbres de Acultzingo, F.M.N.H. (11), M.C.Z. (2), U.I.M.N.H. (8), U.M.M.Z. (3); Fortln de las Flores, U.I.M.N.H. (1), U.M.M.Z. (5); 1,6 kilometers north of Fortin de las Flores, U,I,M,N,H, (56); 3 kilometers north of Fortin de las Flores, U,I,M,N,H, (1); 9 kilo- meters southwest of Fortin de las Flores, U,M,M,Z, ( 2 ) ; 5 kilometers west of Fortin de las Flores, U,I,M,N,H, (9), U,S,N,M, (15), Huatusco, K.U, (6), 7 kilometers northeast of Huatusco, U,M,M.Z, (18); 1,6 kilometers south of Huatusco, U,I,M,N,H, (1); 3 kilometers southwest of Huatusco, K,U, (5 skeletons), LI,M.M,Z, (57); 7,5 kilometers southwest of Huatusco, U.M.M.Z, (9); 12 kilometers southwest of Huatusco, U.M.M.Z, (1); Jalapa, B.M,N,H, (5), F,M,N,H, (14), U,I,M,N,H, (8); 5 kilometers north of Jalapa, KU, (11), T,C,W,C, (1); 10 kilometers south of Jalapa, U.M.M.Z. ( 1 ); La Perla, M.CZ. (93), U,I,M,N.H, (21); Laguna Encatada. Sierra de los Tuxtlas, U,M,M,Z, (8); Los Chaneques, 2 kilometers north of Santiago, Tu.xtla, U,I.M.N,H, (3), U,M,M,Z, (22, I tadpoles); Mirador, U,S.N,M, (2); Orizaba, M.CZ, (5), U.M.M.Z. (13); 2 kilometers north of Paraje Nuevo, K.U. (1); Potrero, F.M.N.H. (1), U.I.M.N.H. (4), U.S.N.M. (3), U.M.M.Z. (1); Potrero Viejo, U.M.M.Z. (1); Rio Sordo, 3 kilometers west of Jalapa, F.M.N.H. (5); 3 kilometers north- northeast of San Andres Tuxtla, T.CW.C. (3), U.M.M.Z, (5, 1 tadpoles); 6 kilometers southeast of Tebanca, T.C,W,C, ( 1 ); between Tebanca and Volcan Santa Maria, U,M,M,Z, (1); Tequeyutepec, M,C,Z, 708 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 (3); Teocelo, K.U. (18); 3 kilometers north of Teo- celo, F.M.N.H. (3); 15 kilometers east-northeast of Tlacotepec, K.U. (1); 4 kilometers west of Tlapaco- yan, K.U. (1); Volcan Pajapan, U.I.M.N.H. (3); southeast slope of Volcan San Martin, K.U. (45, 3 skeletons, 2 tadpoles, 2 eggs), U.I.M.N.H. (19), U.M.M.Z. (10); Xico, U.I.M.N.H. (21), U.S.N.M. ( 1 ) ; between Xometla and La Perla, Pico de Orizaba, K.U. ( 14); Zongolica, I.P.N. (1). Hyla mixe MEXICO: Oaxaca: 4.2 kilometers south of Campamento Vista Hermosa, K.U. (2, 1 skull, 1 tad- poles ). Hyla mixomaculata MEXICO: Veracruz: Barranca metlac, U.M.M.Z. (2); Coscomatepec, K.U. (7, 2 skeletons); 7.2 kilo- meters southwest of Coscomatepec, U.M.M.Z. (1); Huatusco, K.U. (2); 3 kilometers southwest of Hua- tusco, K.U. (1 tadpoles), U.M.M.Z. (1 tadpoles); 7.5 kilometers southwest of Huatusco, U.M.M.Z. (1); 12 kilometers southwest of Huatusco, U.M.M.Z. (2); Sumidero, M.C.Z. (1); 1.6 kilometers west of Xico, U.M.M.Z. (3). Hyla nubicola MEXICO: Veracruz: Barranca Metlac, U.M.M.Z. (1, 1 skeleton); 7 kilometers northeast of Huatusco, U.M.M.Z. (1); 1.9 kilometers south of Huatusco, U.I.M.N.H. (1); 3 kilometers southwest of Huatusco, K.U. (1), U.M.M.Z. (1). Hyla pachydeima MEXICO: Veracruz: Pan de Olla, south of Tezuitlan, Veracniz, U.S.N.M. (4). Hyla pellita MEXICO: Oaxaca: 30 kilometers north of San Gabriel Mi.xtepec, K.U. (2); 33 kilometers north of San Gabriel Mixtepec, K.U. (3, 1 skeleton). Hyla pentheter MEXICO: Oaxaca: 29 kilometers south-southeast of Juchatengo, K.U. (1); Pluma Hidalgo, A.M.N.H. ( 1 ) ; 37 kilometers north of San Gabriel Mixtepec, K.U. (3, 1 skeleton, 1 tadpoles), U.M.M.Z. (5). Hyla phlebodes NICARAGUA; Zelaya: Isla Grande del Maiz, M.C.Z. (1); Rio Mice, Recrero, K.U. (1), U.M.M.Z. (6). COSTA RICA: Alajuela: 12.4 kilometers north of Florencia, M.V.Z. (3), U.S.C. (1); Las Playuelas, 11 kilometers south of Los Chiles, U.S.C. ( 1 ) ; Los Chiles, U.S.C. (2); 3 kilometers northeast of Muelle de Arenal, U.S.C. (2); "San Carlos," U.S.N.M. (1). Cartage: Chitaria, K.U. (1); El Silencio, 14.4 kilo- meters northeast of Turrialba, K.LI. (2); 1.6 kilome- ters east of the Rio Reventazon bridge, east of Tur- rialba, U.M.M.Z. (2); Tunnel camp, near Peralta, K.U. (13, 1 skeleton); Turrialba, F.M.N.H. (3), K.U. (46, 6 skeletons, 1 eggs, 1 tadpoles), M.C.Z. (5), U.M.M.Z. (10), U.S.C. (16), U.S.N.M. (1). Guarja- caste: Arenal, U.S.C. (1); Finca San Bosco, U.S.C. (7); Guayabo de Bagaces, U.S.C. (4); Laguna Arenal, U.S.C. (4); 3 kilometers northeast of Tilaran, U.S.C. (1); 6 kilometers northeast of Tilaran, U.M.M.Z. (6, 1 skeleton), U.S.C. (9). Heredia: Puerto Viejo, K.U. (46, 5 skeletons, 4 tadpoles); 4.2 kilometers west of Puerto Viejo, K.U. (2); 5.9 kilo- meters west of Puerto Viejo, K.U. (5); 7.5 kilometers west of Puerto Viejo, K.U. (1). Limon: Batan, U.M.M.Z. (2); La Castilla, A.N.S.P. (1); Puerto Limon, K.U. (7). PANAM.-V: Bocaa del Toro: 3.2 kilometers north- west of Almirante, K.U. (1); Cayo de Agua, K.LL (5); Fish Creek, K.U. (3); Isla Escudo de Veraguas, K.U. (2); mouth of Rio Cahuita, K.U. (2). Canal Zone: Barro Colorado Island, A.M.N.H. (1), A.N.S.P. (7), F.M.N.H. (4); Juan Mina, A.M.N.H. (1), U.U. (1); 8.6-13.8 kilometers north of Miraflores Locks, T.N.H.C. (63); Rio Chagres, A.M.N.H. (4); Rio Cocoli, 3.5 kilometers north of Miraflores Locks, T.N.H.C. (11); Summit, A.N.S.P. (1); Three Rivers Plantation, S.U. (1). Cocle: El Valle de Anton, A.M.N.H. (5), A.N.S.P. (4). Colon: Acliiote, K.U. (64); Ciricito, C.A.S. (4). Darien: Rio Canclon at Rio Chucunaque, U.M.M.Z. (1); Rio Chucunaque, near Yavisa, A.M.N.H. (1). Panama: Cerro La Campana, F.M.N.H. (4) San Bias: Sasardi, K.U. (1). Veraguas: mouth of Rio Concepcion, K.U. (1). Hyla picadoi COSTA RICA: Alajuela: Poasito, U.F. (1); 13 kilometers northwest of Poasito, U.S.C. (1); Rio Poasito, 1 kilometer west of Poasito, K.U. (2, 1 skele- ton). Cartago: 1 kilometer southeast of La Chonta, U.S.C. (1); 6 kilometers north of Las Cruces, (6); Volcan Turrialba, U.M.M.Z. (1). Heredia: Volcan Barba, A.N.S.P. (3), M.C.Z. (1), U.M.M.Z. (1). San lose: El Empalme, K.U. (1), U.S.C. (1). PANAMA: Bocas del Toro: north slope Cerro Pando, 1920 meters, K.U. (2). Hyla picta MEXICO: Campcche: 7.5 kilometers west of Escarcega, K.U. (16); Matamoros, F.M.N.H. (11); Paicatiin Rio Candelaria, F.M.N.H. (27); Tres Brazos, F.M.N.H. (1). Chiapas: Palenque, U.I.M.N.H. (3); 18 kilometers .south of Teapa (Tabasco), U.I.M.N.H. (1). Oaxaca: 5 kilometers north of Chiltepec, K.\J . (3); 3 kilometers north of Donaji, U.M.M.Z. (8); 78 kilometers north of La Venta, T.N.H.C. (4); 85 kilo- meters north of La Venta, T.N.H.C. (1); 3.7 kilome- ters north of Sarabia, U.M.M.Z. (11); 1.6 kilometers south of Tolo.sita, U.M.M.Z. (14); Tuxtepec, K.U. (2), U.I.M.N.H. (14); 3 kilometers south of Tu.\- 1970 DUELLMAN: HYLID FROGS 709 tepee, U.I.M.N.H. ( 1 ) 13 kilometers south of Tuxte- pec, U.I.M.N.H. (3); 1 kilometer south of Ubero, U.M.M.Z. (6); 2 kilometers south of Valle Nacional, K.U. (1). Pucbla: 9 kilometens south of Tepeojuma, U.M.M.Z. (5); San Diego, A.M.N.H. (15); Villa Juarez, T.N.H.C. (18), U.I.M.N.H. (4). San Luis Fotosi: 16 kilometers north of Tamazunchale, U.I.M.N.H. (3); Valles, F.M.N.H. (2). Tabasco: Frontera, U.S.N.M. (8); 6 kilometers west of Palo Mulato, U.M.M.Z. (3); Teapa, U.M.M.Z. (34); 10 kilometers north of Teapa, U.M.M.Z. (4); 15 kilome- ters north of Teapa. U.M.M.Z. (1); Villahermosa, U.I.M.N.H. (2); 17 kilometers south of Villahermosa, U.M.M.Z. (3). Veracruz: 19 kilometers north of Acayucan, U.I.M.N.H. (4); 2.5 kilometers east- southeast of Alvarado, U.M.M.Z. (5); 3.2 kilometers southeast of Alvarado, U.M.M.Z. (5); 4.5 kilometers south of Aquilera, U.M.M.Z. (1); Barranca Metlac, U.I.M.N.H. (6); 10 kilometers south of Catemaco, U.M.M.Z. (2); 8 kilometers southwest of Coatzacoal- cos, U.M.M.Z. (1); 5 kilometers east-southeast of Cordoba, T.N.H.C. (39); 6.4 kilometers east-south- east of Cordoba, A.M.N.H. (16), U.M.M.Z. (14); Coyame, U.I.M.N.H. (1), U.M.M.Z. (6); Cuautla- pan, F.M.N.H. (1), U.I.M.N.H. (9), U.S.N.M. (6); 1.6 kilometers east-northeast of Encinal, U.M.M.Z. (2); 1 kilometer south of Encinal, U.M.M.Z. (4); Hacienda Tamiahua, Cabo Rojo, K.U. (1); 3 kilome- ters southwest of Huatusco, U.M.M.Z. (60); 10 kilo- meters southeast of Hueyapan, U.M.M.Z. (7); 20 kilometers east-northeast of Jesvis Carranza, K.U. (1); north side Lago de Catemaco, K.U. (3); 21.6 kilome- ters south of Las Choapas, T.C.W.C. (4); 5 kilome- ters northwest of Lerdo de Tejada, U.M.M.Z. (1); 17 kilometers east of Martinez del Torre, U.I.M.N.H. ( 1 ); 6.4 kilometers west of Martinez del Torre, U.I.M.N.H. (2); 2 kilometers east-northeast of Mata Oscura, K.U. (19); 4 kilometers northeast of Miniatitlan, T.N.H.C. (1); Orizaba, U.S.N.M. (1); Potrero Viejo, F.M.N.H. (5), M.C.Z. (2), U.I.M.N.H. (56), U.M.M.Z. (68), U.S.N.M. (25); Tierra Colorada, F.M.N.H. (6), U.I.M.N.H. (3); Tlalpan, F.M.N.H. (2); 2.7 kilo- meters northwest of Tulsa, U.M.M.Z. (3); 8 kilome- ters south of Veracruz, U.M.M.Z. (2); 3 kilometers west of Veracruz, U.I.M.N.H. (15); 7 kilometers west of Veracruz, K.U. (1). BRITISH HONDURAS: Cayo: 5 kilometers southwest of Cayo, U.M.M.Z. (6); 6.4 kilometers south of Cayo, M.C.Z. (3); 5 kilometers west of Pine Ridge road on Belize-Cayo highway, U.M.M.Z. (1). Stauu Creek: 10 kilometers east of Stann Creek, U.M.M.Z. (1). GUATEMALA: Aha Verapaz: 5.1 kilometers northeast of Campur, K.U. (6); Finca Chama, U.M.M.Z. (72); 13 kilometers south of Sebol, T.N.H.C. (1). ElPeten: Dolores, U.M.M.Z. (1); La Libertad, U.M.M.Z. (7); Toocog, 15 kilometers south- east of La Libertad, K.U. ( 8 ) . Izabal: Puerto Barrios, T.C.W.C. (5); 2.5 kilometers northeast of Rio Blanco, K.U. (12, 2 skeletons). HONDURAS: Atlantidad: Ceiba, U.S.N.M. ( 1 ); Lancetilla, M.C.Z. (6). Hyla pictipes COSTA RICA: Alajuela: Rio Poasito, 1 kilometer north of Poasito, K.V. (68, 4 skeletons, 4 tadpoles), M.C.Z. (2); Volcan Poas, U.M.M.Z. (1 tadpoles). Cartago: 0.5 kilometers east of Tierra Blanca, U.M.M.Z. (1 tadpoles). Heredia: Paso Llano, south slope of Volcan Barba, K.U. ( 13 tadpoles); Rama Sur of Rio Las Vueltas, south slope of Volcan Barba, K.U. (7, 1 skeleton), U.S.C. (29); Volcan Barba, U.S.C. (6). San Jose: south slope of Cerro de la Muerte, U.S.C. (4); La Palma, A.N.S.P. (8), U.M.M.Z. (1); 2 kilometers north of Las Nubes, K.U. (1); 3 kilo- meters southeast of Rancho Redondo, U.M.M.Z. (2), U.S.C. (2). Hyla pinorum MEXICO: Guerrero: Agua del Obispo, F.M.N.H. (1), K.U. (1 tadpoles), U.I.M.N.H. (1); 1.6 kilome- ters east of San Andreas de la Cruz, K.U. (3, 1 tad- poles), U.M.M.Z. (3, 1 skeleton, 1 tadpoles); 3.3 kilometers north of San Vicente, K.U. (5). Oaxaca: San Vicente, U.I.M.N.H. (1). Hyla plicata MEXICO: No specific locality, M.N. H.N. (1). Distrito Federal: Canon Contreras, 10 kilometers southwest of Ciudad Mexico, U.M.M.Z. (2); Desierto de los Leones, A.M.N.H. (4, 1 tadpoles), M.C.Z. (1); Pico Santo Rosa, U.M.M.Z. (1); 15.5 kilometers southwest of Villa Obregon, U.M.M.Z. (1). Hidalgo: El Chico Parque Nacional, A.M.N.H. (5), K.U. (21, 1 skeleton, 1 tadpoles), U.I.M.N.H. (5), U.S.N.M. (3); Guerrero, M.C.Z. (3), U.M.M.Z. (2); San Miguel Regla, F.M.N.H. (2), U.M.M.Z. (4); Velasco, A.M.N.H. (12), M.C.Z. (1). Mexico: 23 kilometers west of Ciudad Me.xico, M.V.Z. (I); 55 kilometers southeast of Ciudad Me.xico, T.C.W.C. (9); 5 kilo- meters west of Las Cruces, U.I.M.N.H. (2); Llano Grande near Rio Frio, F.M.N.H. (1), M.C.Z. (2), T.C.W.C. (21), U.I.M.N.H. (28), U.S.N.M. (23); Nevado de Toluca, A.M.N.H. (1), F.M.N.H. (1); Salazar, I.P.N. (2); San Juan Teotihuacan, A.M.N.H. (2), M.C.Z. (2); 5.6 kilometers south of Tenango, T.C.W.C. (10); 18 kilometers north of Tenancingo, K.U. (1); Tlamacas, I.P.N. (1). Michoacdn: Cerro San Andres, U.M.M.Z. (26); 8 kilometers south- southeast of Opopeo, K.U. (4); 12 kilometers south- southeast of Opopeo, U.M.M.Z. (4); 3 kilometers east of San Gregorio, K.U. (56, 1 skeleton). Morelos: 3 kilometers west of Huitzilac, T.C.W.C. (5); 5 kilo- meters west of Huitzilac, U.M.M.Z. (1); 6 kilometers west of Huitzilac, K.U. (1); Lagunas de Zempoala, A.M.N.H. (8), F.M.N.H. (11), I.P.N. ( 11 ), T.C.W.C. (21), U.I.M.N.H. (58), U.S.N.M. (3); 7.3 kilome- ters southeast of Santa Martha, U.I.M.N.H. (2). Puebla: 14.4 kilometers northeast of Acatzingo, K.U. (1); Cruz Alto, south of Aquixtla, U.M.M.Z. (2); Rio Otlati, 15 kilometers northwest of San Martin, T.C.W.C. (1). Tlaxcala: 5 kilometers northwest of Sanctorium, T.N.H.C. (1); 13 kilometers northeast of Tlaxcala, T.C.W.C. (1). Veracruz: 12 kilometers 710 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 southwest of Huatusco, U.M.M.Z. (2); Las Vigas, F.M.N.H. (2),U.I.M.N.H. (1), U.M.M.Z. (2). Hyla pseudopuma infucata PANAMA: Bocas del Tow: Rio Claro near junc- tion with Rio Changena, K.U. (31, 5 skeletons); Rio Changena, 830 meters, K.U. (11, 1 skeleton), M.C.Z. (2). Hyla pseudopuma pseudopuma COSTA RICA: Ahjucla: 25 kilometers northwest of Barba U.M.M.Z. (2); Chinchona, K.U. (8, 1 skele- ton, 4 tadpoles), U.S.C. (2); between Chinchona and Salto El Angel, U.S.C. (1); Salto El Angel, U.S.C. (1); 21 kilometers north of Varablanca, U.M.M.Z. (1); Volcan, Poas, K.U. (9), U.M.M.Z. (1 tadpoles); 1.6 kilometers south of Zapote, U.S.C. (2). Cartago: 29 kilometers south of Cartago, U.S.C. (1); 1 kilome- ter southeast of La Chonta, U.S.C. (3); Moravia de Turrialba, K.U. (15, 7 skeletons); Rio Playas, U.S.C. (2); Tapanti, K.U. (52, 7 skeletons, 15 tadpoles, 1 eggs); Tuis, K.U. (1). Heredia: Finca El Conde de Tattenbach, south slope of Volcan Barba K.U. (4); Hacienda Cavuga, 1 kilometer north of Montana Azul, K.U. (1); Isla Bonita, F.M.N.H. (1), K.U. (1); 9 kilometers north of La Concordia; K.U. (2); Montana Azul, K.U. (1); 3 kilometers north of Montana Azul, K.U. ( 1 ) ; Paso Llano, south slope of Volcan Barba, K.U. (3); Rama Sur of Rio Las Vueltas, south slope of Volcan Barba, K.U. (16, 1 skeleton, 5 tadpoles), U.S.C. (1); Varablanca, K.U. (12); Volcan Barba, M.C.Z. (5), U.M.M.Z. (1 tadpoles), U.S.C. (1). Puntarenas: Monteverde, U.S.C. (19); 3.6 kilometers east of Monteverde, U.S.C. (14). San Jose: El Copev, M.C.Z. (1); El Empalme, U.S.C. (64), K.U. (1 tad- pole); 3 kilometers northwest of El Empalme, K.U. (1); 7 kilometers southeast of El Empalme, K.U. (1); La Estrella, M.C.Z. (3); La Palma, A.N.S.P. (26), F.M.N.H. (3), K.U. (29, 3 tadpoles), M.C.Z. (2), U.M.M.Z. (10), U.S.C. (58), U.S.N.M. (1); Rio Claro at Rio La Hondura, U.S.C. (7); San Pedro, A.M.N.H. (3), U.S.C. (3); San Isidro, A.N.S.P. (2); 6.4 kilometers north of San Isidro el General, M.C.Z. (5); 14 kilometers north of San Isidro el General, K.U. (1); 15 kilometers north of San Isidro el Gen- eral, K.U. (6, 6 tadpoles); 18 kilometers north of San Isidro el General, K.U. (1), U.M.M.Z. (1); 12.4 kilo- meters southwest of San Isidro el General, U.S.C. ( 1 ). PANAMA; Chiriqui: Boquete, U.M.M.Z. (4). Hyla regilla curta MEXICO: Baja California Sur: Cabo San Lucas, M.C.Z. (1), U.S.N.M. (17); Canon Cantiles, U.S.N.M. (1); Comondu, U.S.N.M. (2); Isla Coronado, M.C.Z. (1); La Paz (5); Miraflores, A.M.N.H. (16); Rancho de Parras, 19 kilometers south of Loreto, A.M.N.H. (2); San Ignacio, A.M.N.H. (1), M.C.Z. (1), U.M.M.Z. (28); Soria, U.S.N.M. (12); Todos Santos, K.U. (11). Hyla regilla hvpochondriaca MEXICO: Baja California Norte: Canon de las Palmas, Sierra de Juarez, U.S.N.M. (1); 4 kilometers north of Descausa, U.M.M.Z. (1), 77 kilometers southeast of Ensenada, L.B.S.C. (2); Isla Cedros, A..\I.N.H. (2), U.S.N.M. (52); La Grulla, U.S.N.M. (1); Laguna Hanson, Sierra de Juarez, L.B.S.C. (1); Mattomi, F.M.N.H. (1); Playa Estero, 14.4 kilome- ters south of Ensenada, A.M.N.H. (8); Punta Clara, U.M.M.Z. (1); Rosario, U.M.M.Z. (2); San Quintin, U.S.N.M. (1); Sierra San Pedro Martir, U.S.N.M. (19); Te.xate, U.S.N.M. (15); Tijuana, A.M.N.H. (2). Hyla rivularis COSTA RICA: Alaiuela: Chinchona, K.U. (18, 7 tadpoles), U.S.C. (7); between Chinchona and Salto El Angel, U.S.C. (4); Rio Poasito, 1 kilometer north of Poasito, K.U. (21, 4 skeletons, 1 tadpoles); Salto El Angel, U.S.C. (2); 10 kilometers south of Varablanca, U.S.C. (2); 22.5 kilometers northwest of Varablanca, U.M.M.Z. (1 tadpoles); 1.6 kilometers south of Zapote, U.S.C. (10); 8 kilometers north of Zarcero, U.S.C. (3). Cartago: 1.6 kilometers north- east of Casa Mata, U.S.C. (1); 1 kilometer east of Pacayas. U.S.C. (1); Santa Cruz, U.S.C. (7); Volcan Turrialba, 1380 meters, U.M.M.Z. (14); Volcan Turrialba, 2175 meters, U.M.M.Z. (1). Heredia: El Gallito, Volcan Barba, A.N.S.P. (1), U.S.N.M. (3); Hacienda Cayuga, 1 kilometer north of Montana Azul, K.U. (3, 1 tadpoles) 5 kilometers south of Los Car- tagos, K.U. (3, 4 tadpoles); 3 kilometers north of Montana Azul, K.U. (1 tadpole); Rama Sur of Rio Las Vueltas, K.U. (11. 1 tadpoles), U.S.C. (19); San Jose de la Montana, K.U. (1); 2.7 kilometers north of San Jose de la Montana, K.U. (3 tadpoles), U.S.C. (1); 1.6 kilometers north-northeast of Uvita, U.S.C. (25); Volcan Barba, A.N.S.P. (4). Puntarenas: Monteverde, U.S.C. (26). San Jose: Cerro de la Muerte, M.C.Z. (2), U.S.C. (5); El Copey, A.N.S.P. (1); La Hondura, U.M.M.Z. (1); 2 kilometers north of Las Nubes, K.U. (3, 1 tadpoles); 18.7 kilometers north of San Isidro el General, U.M.M.Z. (15). P.\NAMA: Bocas del Toro: north slope of Cerro Pando, 1920-2100 meters, K.U. (41). Chiriqui: Finca Bambito, 6 kilometers east-northeast of El Volcan, K.U. (3); Finca Ojo de Agua, southeast slope of Cerro La Pelota, K.U. (1); Quebrada Chevo, south slope of Cerro La Pelota, K.U. (9); south slope of Cerro Santa Catalina, 8 kilometers northwest of El Volcan, K.U. (9,1 tadpole). Hyla robertmerfensi MEXICO: Chiapas: Acacojagua, U.S.N.M. (8); 2 kilometers west of Acacoyagua, U.M.M.Z. (203); 32 kilometers north of Arriaga, K.U. (6, 2 skeletons), 32 kilometers southeast of Arriaga, U.I.M.N.H. (2); Asuncion, F.M.N.H. (5), U.I.M.N.H. (7), U.S.N.M. (1); 5 kilometers east of Hui.\tla, U.I.M.N.H. (15); La E.speranza, U.S.N.M. (16); 17 kilometers south of Las Cruces, K.U. (25, 1 eggs); 8.5 kilometers north 1970 DUELLMAN: HYLID FROGS 711 of Puerto Madero, U.M.M.Z. (2); 11.7 kilometers north of Puerto Madero, U.M.M.Z. (1); Tapaclnila, F.M.N.H. (1), U.I.M.N.H. (1); 11 kilometers south of Tapachula, K.U. (14, 1 skeleton); Tonala, F.M.N.H. (7), U.I.M.N.H. (1); 16 kilometers south- east of Tonala, U.I.M.N.H. (1). Oaxaca: Tapana- tepec, U.M.M.Z. (2); 1.6 kilometers east of Tapana- tepec, U.M.M.Z. (14); 4.3 kilometers east of Ta- panatepec, U.I.M.N.H. (2); 7.5 kilometers west of Tapanatepec, U.M.M.Z. (39); 12.8 kilometers west of Tapanatepec, K.U. (8); 7.2 kilometers west-northwest of Zanatepec, U.M.M.Z. (77); 13.6 kilometers west- northwest of Zanatepec, T.N.H.C. (10); 22.7 kilome- ters west-northwest of Zanatepec, T.N.H.C. (7). GUATEMAL.\: Jiitiapa: Jutiapa, U.M.M.Z. (1); La Trinidad, U.M.M.Z. (23). RctaUuieleu: Casa Blanca, U.M.M.Z. (1), EL SALVADOR: La Libertad: 16 kilometers northwest of Santa Tecla, K.U. (1). San Salvador: 21.9 kilometers north of San Salvador, U.M.M.Z. (6). Hyla robertsorum MEXICO: Hidalgo: 16 kilometers west of Agua Buena, U.M.M.Z. (6); El Chico Parque Nacional, F.M.N.H. (2), K.U. (49, 5 skeletons), M.C.Z. (2), U.I.M.N.H. (45), U.M.M.Z. (6, 1 tadpoles), U.S.N. M. (25); 3.3 kilometers north of Zacualtipan, K.L'. (1, 1 tadpoles); 8.5 kilometers southeast of Zacualtipan, K.LI. (1 tadpoles). Pitebla: Honey, U.M.M.Z. (1). Hyla rosenbergi COSTA RICA: Puntarenas: Cerro Puntado, 2 kilometers northeast of Jaco, U.S.C. (1); Dominical, U.U. (3); Golfito, K.U. (20, 2 skeletons), U.M.M.Z. (3), U.S.C. (6); 3 kilometers east of Golfito, K.U. (4), II kilometers east of Golfito, K.U. (2 skeletons), U.S.C. (1); Palmar Sur, K.U. (3); 2.5 kilometers southeast of Palmar Sur, K.U. (3); 4 kilometers south- east of Palmar Sur, K.U. (2); Rincon de Osa, U.M.M.Z. (4); U.S.C. (8), 4.5 kilometers west of Rincon de Osa, K.LI. (4); Rio Ferruviosa, 4.5 kilome- ters south of Rincon de Osa, U.M.M.Z. (2), U.S.C. (1). Villa Neilly, K.U. (1); 1 kilometer west-north- west of Villa Neilly, U.S.C. (1); 7 kilometers west- northwest of Villa Neilly, U.S.C. (2); 10.5 kilometers west-northwest of Villa Neilly, K.U. (2, 1 skeleton); 22 kilometers west-northwest of Villa Neilly, U.S.C. (5). PANAMA: Canal Zone: Alhajuela, U.M.M.A. (1); Camp Chagres, K.U. (83; 5 kilometers northwest of Gamboa, K.U. (1); Madden Dam, U.M.M.Z. (3); San Pablo, M.C.Z. (I); Summit Gardens, K.U. (I). Chihqui: Puerto Armuelles, A.M.N.H. (2), A.N.S.P. (2). Darien: Camp Creek, below Yavisa, A.M.N.H. (25, 4 tadpoles); Cana, U.S.N.M. (1); Chalichimans Creek, Rio Subcuti, A.M.N.H. (1); Rio Esnape, M.C.Z. (1); Rio Membrillo, mouth, A.M.N.H. (1); Rio Chucunaque, 7 kilometers above Rio Morti, K.U. (3); Rio Chucunaque, 10 kilometers below Rio Sub- cuti, K.U. (3); Rio Chucunaque at Rio Ucurgan- ti, U.S.N.M. (2); Rio Sanson, A.M.N.H. (1); Rio Tuira at Rio Mono, K.U. (25, 2 skeletons); Tacarcuna, K.U. (6); Three Falls Creek, below Yavisa, A.M.N.H. ( 2 ) . Los Santos: Tonosi ( 3 ) . PanatJid: Bejuco, Rio Bejuco, A.M.N.H. (1); 6 kilometers west-southwest of Chepo, K.U. (5); Rio Bayano, F.M.N.H. (1), U.S.N.M. (1). Hyla rostrata PANAMA: Canal Zone: No specific locality, A.M.N.H. (3), T.N.H.C. (6); between Gatuncillo and Guayabalito, A.M.N.H. (1) ; 11 kilometers north- west of Miraflores Locks, T.N.H.C. (1); Road K2, T.N.H.C. (2). Panama: 3 kilometers west-southwest of Chepo, K.LI. (9, 2 tadpoles); 6 kilometers west- southwest of Chepo, K.U. (4), M.C.Z. (2); La Jolla, A.M.N.H. (1); 1.5 kilometers southwest of Naranjal, K.U. ( 1, 1 skeleton); 9 kilometers northeast of Pacora, K.U. (1); 2 kilometers north of Tocumen, K.U. (5, I ske'eton ) ; 8 kilometers northeast of Tocumen, K.U. (9). San Bias: Sasardi, K.U. ( 1 ). Hyla rubra P.'VNAMA: Canal Zone: No specific locality, U.S.N.M. (1); Madden Dam, F.M.N.H. (1); San Pablo, M.C.Z. (2). Colon: Achiote, U.F. (13); Cerro Bruja, M.C.Z. (I). Darien: El Real, U.S.N.M. (2); Yavisa, M.V.Z. (8). Panama: luan Diaz, M.C.Z. (1); Las Sabanas, M.C.Z. (1); Rio Trinidad, U.S.N.M. (1); 17 kilometers east of Tocumen, M.V.Z. (1). Hyla rufioculis COSTA RICA: Alajuela: Cinchona, K.U. (21), U.M.M.Z. (2); 5 kilometers south of Ciudad Quesada, U.S.C. (1). Cartago: Moravia de Turrialba, K.U. (107, 3 skeletons), M.C.Z. (3); Morehouse Finca, 7 kilometers south of Turrialba. F.M.N.H. (4), K.U. (17); Rio Chitaria, 3 kilometers north-northeast of Pavones, U.S.C. (3); Turrialba, K.U. (13, 1 skele- ton). Guanacaste: El Silencio, U.S.C. (4). Heredia: Isia Bonita, F.M.N.H. (6), K.U. (13), M.C.Z. (1). Limon: El Tigre, 12-20 kilometers southwest of Siquirres, U.S.C. (6); Rio Lari at Rio Dipnari, 21 kilometers southwest of Aniubri, L.A.C.M. (1). Pun- tarenas: Finca Loma Linda, 2 kilometers southwest of Cafias Gordas, ( 1 ) ; 3.6 kilometers east of Monte- verde, U.S.C. (1). San Jose: south slope of Cerro de la Muerte, 1540 meters, U.S.C. (3); Rio Claro at Rio La Hondura, U.S.C. (10); 13 kilometers north of San Isidro el General, K.U. (12), U.M.M.Z. (3), U.S.C. (1); 14 kilometers north of San Isidro el Gen- eral, K.U. (13), U.M.M.Z. (13); 15 kilometers west- southwest of San Isidro el General, K.U. (45, 5 skele- tons, 5 tadpoles), U.M.M.Z. (11), U.S.C. (58); 17.2 kilometers west-southwest of San Isidro el General, U.S.C. (2). Hyla rufitela NICARAGUA: No specific locality, U.S.N.M. (2). Zelaya: El Recreo, K.U. (1); Machuca, A.N.S.P. (2); Maselina Creek, A.M.N.H. (2). COSTA RICA: Heredia: Puerto Viejo, U.C.R. 712 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 (2). Limon: La Castilla, A.N.S.P. (15); Rio Tortu- guero, 3 kilometers from mouth, A.M.N.H. (1). Pun- tarenas: Golfito, K.U. (2, 1 tadpoles); 4.5 kilometers west of Rincon de Osa, K.U. ( 1 tadpoles). PANAMA: Bocas del Tom: Cayo de Agua, G.M.L. (1), K.U. (3); Isla Colon, K.U. (1); Penin- sula Valiente, Bluefields, K.U. (2); Rio Cahuita, mouth, K.U. (8). Canal Zone: Barro Colorado Island, A.N.S.P. (7), F.M.N.H. (7), K.U. (11, 1 skeleton, 7 tadpoles, 1 eggs), T.N.H.C. (1), U.M.M.Z. (3). Colon: Achiote, K.U. (10). Panama: Rio Puente, M.C.Z. (1). Veragtias: Rio Concepcion, mouth, K.U. (1). Hyla salvadorensis EL SALVADOR: Santa Ana: Hacienda Monte- cristo, Cerro Metapan, K.U. (1 tadpoles); Hacienda Los Planes, U.I.M.N.H. (1); Rancho San Jose, K.U. (3, 1 skeleton, 2 tadpoles). HONDURAS: Franciico-Morazdn: West slope of Cerro Uyuca, A.M.N.H. (14), K.U. (6, 1 skeleton), U.M.M.Z. (1). Hyla sartori MEXICO: Guerrero: 23.2 kilometers north of Acapulco, U.I.M.N.H. (4); 5 kilometers east of Aca- pulco, A.M.N.H. (2); Colonia Buenos .\ires, 23 kilo- meters east of Tecpiin de Galeana, U.M.M.Z. (7); El Limoncito, F.M.N.H. (16), U.M.M.Z. (1), U.S.N.M. (1); El Treinte, F.M.N.H. (1), U.I.M.N.H. (3); La- guna Coyuca, A.M.N.H. (1); La Venta, M.C.Z. (1); Marijonares, U.I.M.N.H. (11); 1,6 kilometers north of Organos, F.M.N.H. (2), U.I.M.N.H. (2); 19.2 kilometers south of Petaquillas, U.I.M.N.H. (1); 6.1 kilometers east of Tecpan de Caleana, T.N.H.C. (13); 11.2 kilometers west-northwest of Tierra Colorado, U.I.M.N.H. (1); 11.8 kilometers west-northwest of Tierra Colorada, U.M.M.Z. (51, 3 skeletons); Zacual- pan, U.M.M.Z. (6). Jalisco: 6.4 kilometers northeast of La Resolana, K.U. (17); 24 kilometers northeast of La Resolana, K.U. (4). Oaxaca: 3 kilometers north of Pochutia, K.U. (1); 11.3 kilometers north of Poch- utla, U.I.M.N.H. (4); 13.4 kilometers north of Poch- utia, U.M.M.Z. (40). Hyla siopela MEXICO: Veracruz: west slope of Cofre de Perote, K.U. (36, 3 skeletons), U.I.M.N.H. (15). Hyla smaragdina MEXICO: Colima: Paso del Rio, U.M.M.Z. (2). Michoacdn: 6 kilometers east of Cojumatlan, F.M.N.H. (19), M.C.Z. (1), U.M.M.Z. (1); 1.6 kilo- meters north of Copuyo, T.C.W.C. (4); 8 kilometers north of Copuyo, T.C.W.C. (3); 17 kilometers east of Dos Aguas, U.M.M.Z. (22); Ostula, U.M.M.Z. (8); Pomaro, U.M.M.Z. (3); Salitre de Estopilas, U.M.M.Z. (7). Morelos: Tepoztlan, U.I.M.N.H. (1). Nayarit: Santa Barbara, L.A.C.M. (1 tadpoles). Sinaloa: Co- pala, K.U. (7); Potrerillos, K.U. (2 tadpoles); Santa Lucia, K.U. (43, 3 skeletons), L.B.S.C. (31). Hyla smithii MEXICO: Colima: Armeria, U.M.M.Z. (1); 3 kilometers southwest of Colima, U.M.M.Z. (3); 5 kilometers south of Colima, K.V. (1); 72 kilometers southwest of Colima, M.V.Z. (2); Manzanillo, A.M.N.H. (1, 1 skeleton); 16 kilometers north of Manzanillo, C.A.S. (2); 5 kilometers east of Manza- nillo, .\.M.N.H. (2); 41.7 kilometers northwest of Manzanillo, M.V.Z. (I); Paso del Rio, F.M.N.H. (1), U.I.M.N.H, (1), U.M.M.Z. (9); Periquillo, U.M.M.Z. (23); Queseria, F.M.N.H. (7), M.C.Z. (2), U.I.M.N.H. (7), U.M.M.Z. (17); Rio Astillero, C.A.S. (1); 7 kilometers southwest of Tecolapa, LI.M.M.Z. (3); 10 kilometers north of Trapicliillos, U.M.M.Z. (1). Guerrero: Acahuitzotla, T.C.W.C. (32); Aca- pulco, T.C.W.C. (2), U.M.M.Z. (1); 13 kilometers north of .Acapulco, T.N.H.C, (2); 24 kilometers north of Acapulco, F.M.N.H. (39), U.I.M.N.H. (29); 26 kilometers north of Acapulco, T.N.H.C. (2); 27 kilo- meters northeast of Acapulco, U.I.M.N.H. (1); Agua del Obispo, F.M.N.H. (11), T.C.W.C. (5), U.I.M.N.H. (11), U.M.M.Z, (12), U.S,N,M. (16); 2 kilometers north of Agua del Obispo, T.C.W.C. (5); 4 kilometers south of Almolonga, T.C.W.C. (5); 5 kilometers west of Bajos de Ejido, U.M.M.Z. (1); Buena Vista, F.M.N.H, (1); 5 kilometers south of Buena Vista, A,M,N,H. (9); Clulpancingo, M.C.Z. (7); 19 kilometers south of Chilpancingo, F.M.N.H. (2); 1,6 kilometers southwest of Colotlipa, T,C.W.C. (2); 13.3 kilometers northwest of Coyuca, U.I.M.N.H. (7); El Limoncito, F.M.N.H. (3), U.I.M.N.H. (3); El Treinta, F.M.N.H. (21), U.I.M.N.H. (17); 22.4 kilometers south of I.xtapan de Sal (Mexico), K.U. (8); Laguna Coyuca, A.M.N.H. (2); 2.5 kilometers north of Mazatlan, U.I,M,N.H. (1), U.M.M.Z. (32); 14.4 kilometers south of Mazatlan, F.M.N.H. (8), U.I.M.N.H, (9); Monjonaros, U.I.M.N.H. (9); 4-6 kilometers north of Ocotito, K.U. (2), T.C.W.C. (10), U.M.M.Z, (5); Palo Blanco, F.M.N.H. (2), U.I.M.N.H. (2); 19 kilometers south of Petaquillas, U.I..M.N.H. (3); 10 kilometers west of Pie de la Cuesta, U.M.M.Z. (1); 19 kilometers south of Puente de I.\tla (Morelos), F.M.N.H. (34), U.I.M.N.H. (22); Rincon, T.C.W.C. (3); 2 kilometers southeast of San Andres de la Cruz, K.U. (2), U.M.M.Z, (1); San Vincente, K.U. (10); 17 kilometers south of Ta.\co, T.C.W.C. (2); 26 kilometers east of Tecpan de Galeana, T.N.H.C. (6); Tierra Colorada, T.C.W.C. (3), U.S.N.M. (7); 7 kilometers soudi of Tierra Colorada, U.M.M.Z. (2); 12 kilometers west-soutli- west of Tierra Colorada, U.M.M.Z. ( 10); 2 kilometers east of Ti.xda, K.U. (2); Xaltianguis, F.M.N.H. (I); 5 kilometers east of Zacatula, U.M.M.Z. (1). Jalisco: .\utlan road (kilometer 133), F.M.N.H. (4), U.I.M.N.H. (7); 5 kilometers east of Autlan, F.M.N.H. (I); south of Autlan, U.I.M.N.H. (6); Barro de Navidad, K.U. (14); 48 kilometers nortli- east of Barro de Navidad, M.V.Z. (8); 5 kilometers 1970 DUELLMAN: HYLID FROGS 713 east of Barro de Na\idad, U.M.M.Z. ( 1); 5 kilometers northwest of Barro de Navidad, K.U. (1); 6.4 kilo- meters northwest of Barro de Navidad, K.U. (4); 12.8 kilometers northeast of La Huerta, K.U. (5); 3 kilometers northeast of La Resolana, U.M.M.Z. (19); 6 kilometers northeast of La Resolana, K.U. (12); 3 kilometers southwest of La Resolana, K.U. (4); 8 kilometers east of Melaque, K.U. (10). Michoacdn: Augililla, U.M.M.Z. (14); Apatzingan, F.M.N.H. (15), M.C.Z. (2), U.l.M.N.H. (46), U.M.M.Z. (9), U.S.N.M. (25); 1.6 kilometers east of Apatzingan, U.M.M.Z. (4); 8.6 kilometers east of Apatzingan, U.M.M.Z. (5); 24.5 kilometers east of Apatzingan, U.M.M.Z. (1) between Apatzingan and Uruapan, C.A.S. (19); 1.6 kilometers north of Arte- aga, U.M.M.Z. (1); 13 kilometers south of Artega, U.M.M.Z. (1 tadpoles); 21 kilometers south of Ar- teaga, U.M.NLZ. (1 tadpoles); 2 kilometers south of Charapendo, U.M.M.Z. (5); 3 kilometers north-north- east of Coalcoman, U.M.M.Z. (5); El Sabino, F.M.N.H. (16), U.l.M.N.H. (13); La Playa de Jorullo, U.NLM.Z. (6); 11.2 kilometers south of Lom- bardia, U.M.M.Z. (1); Playa Azul, U.M.M.Z. (1); between Rio Marquez and Cuatro Caminos, K.U. (9); Salitre de Estiopilas, U.M.M.Z. (1). Morelos: Alpu- yeca, U.l.M.N.H. (4); Antiguo, F.M.N.H. (1); 3.5 kilometers west of Cuautlixco, K.U. (6); Cuernavaca, T.C.W.C. (8), U.l.M.N.H. (3), U.S.N.M. (1); 2.7 kilometers east of Cuernavaca, T.N.H.C. (9); 3 kilo- meters south, 8.8 kilometers east of Cuernavaca, T.C.W.C. (6); Huajintlan, F.M.N.H. (3), U.l.M.N.H. (2); 2 kilometers south of Jonacatepec, T.C.W.C. (21); Progreso, T.C.W.C. (13), U.l.M.N.H. (1); Puente de Ixtla, T.C.W.C. (1), U.l.M.N.H. (74), U.M.M.Z. (3), U.S.N.M. (25); 1 kilometer east of Puente de I.xtla, K.U. (6), T.C.W.C. (16); Temilpa, T.C.W.C. (26), Temoac, T.C.W.C. (4); 17 kilome- ters west of Yautepec, T.C.W.C. (2); Zacatepec, T.C.W.C. (18); 3 kilometers west of Zacatepec, T.C.W.C. (18); U.l.M.N.H. (2). Nayarit: 21.6 kilometers south of Acaponeta, LI.I.M.N.H. (20); 29.5 kilometers south of Acaponeta, U.l.M.N.H. (49); 47 kilometers south of Acaponeta, T.N.H.C. (2); Arrovo de Rifilion, 9 kilometers north of Compostela, C.A.S. (30); Cinco de Mayo, C.A.S. (4); 56 kilo- meters south of Escuinapa (Sinaloa), K.U. (3); 5 kilometers southeast of Huajicori, K.U. (1); 1.6 kilo- meters east of Ixtlan del Rio, U.M.M.Z. (1); 4.3 kilo- meters east of Ixtlan del Rio, C.A.S. (1); 6.4 kilome- ters east-southeast of Ixtlan del Rio, K.U. (12); La Libertad, 16 kilometers northeast of San Bias, U.M.M.Z. (11); Navarrete, L.B.S.C. (6); 2 kilome- ters east of Navarrete, C.A.S. (2), T.N.H.C. (1); 3.5 kilometers southwest of Navarrete, C.A.S. (15), T.N.H.C. (15); Petaquilla, A.M.N.H. (1); Rancho Buenas Aires, 25 kilometers west of Tepic, A.M.N.H. (2); Rio San Cayetano, 5 kilometers east of Tepic, A.M.N.H. (9); San Bias, C.A.S. (13), L.B.S.C. (1), M.V.Z. (1), U.LM.N.H. (4); 1-8 kilometers north- east of San Bias, K.U. (7); 4 kilometers northeast of San Bias, L.B.S.C. (1); 5 kilometers northeast of San Bias, C.A.S. (3); 7-16 kilometers northeast of San Bias, C.A.S. (8); 1.6 kilometers southwest of San Jose del Conde, U.M.M.Z. (9); 2.4 kilometers east of Santa Cruz, C.A.S. (6); 14.5 kilometers east of Santa Cruz, C.A.S. (17); 4.5 kilometers west of Santa Maria del Oro, C.A.S. (9); Tepic, A.M.N.H. (2), C.A.S. (16), F.M.N.H. (17), U.l.M.N.H. (15), U.M.M.Z. (5); 29 kilometers north of Tepic, U.F. (1); 35 kilometers north of Tepic, C.A.S. (32), M.V.Z. (2); 9 kilometers east of Tepic, A.M.N.H. (2); 19 kilometers southeast of Tepic, K.U. (11); 37 kilometers southeast of Tepic, L.B.S.C. (32); 5.5 kilometers south of Tepic, A.M.N.H. (5); 8 kilome- ters east of Villa Hidalgo, C.A.S. (1); 2 kilometers west of Yago, C.A.S. (3). Oaxaca: Chacalapa, K.U. (2); La Candelaria, K.U. (57); 2.5 kilometers south of La Candelaria, K.U. (2); Mira Leon, U.l.M.N.H. (1); 3 kilometers north of Pochutla, K.U. (36, 5 skeletons); 11 kilometers north of Pochutla, A.M.N.H. (1); 13.4 kilometers north of Pochutla, U.M.M.Z. (1); 22.2 kilometers north of Pochutla, U.M.M.Z. (1); 28.2 kilometers north of Pochuda, U.M.M.Z. (10); 17 kilometers north of San Gabriel Mixtepec, K.U. (7); 5.7 kilometers south of San Gabriel Mix- tepec, K.U. (1). Puebla: 10 kilometers southwest of Iziicar de Matamoros, K.U. (24). Sinaloa: 35 kilo- meters north of Acaponeta (Nayarit), U.l.M.N.H. (8); Chele, U.M.M.Z. (7); 4 kilometers northeast of Concordia, K.U. (1); 5 kilometers east of Concordia, C.A.S. (6), L.B.S.C. (1); 10 kilometers southwest of Concordia, K.U. (6); 18 kilometers northeast of Co- pala, U.l. (2); 3.2 kilometers southwest of Copala, K.U. (32); CuUacan, L.B.S.C. (3), U.F. (1); 12.1 kilometers north of Culiacan, U.M.M.Z. (1); 13.6 kilometers northwest of Cuhaean, A.M.N.H. (1); Eldorado, U.l.M.N.H. (12); El Vena- dillo, U.M.M.Z. (3); 34 kilometers southeast of Es- cuinapas, K.U. (5); 7.3 kilometers southwest of Matatan, K.U. (3); Mazatldn, L.B.S.C. (1); 11.3 kilometers north of Mazatlan, L.B.S.C. (1); 32.3 kilo- meters north-northwest of Mazatlan, LI.M.M.Z. (1); 14.7 kilometers south of Mazatlan, L.B.S.C. (1); U.l.M.N.H. (47); Plumosas, K.U. (3); Rio Paxtala, L.B.S.C. (2); 5 kilometers southeast of Rosario, U.LM.N.H. (1); 19 kilometers northeast of San Benito, K.U. (1); San Ignacio, K.U. (5); Teacapan, Isla Palmito del Verde, K.U. (1); 10 kilometers north- northwest of Teacapan, K.U. (1); Villa Union, K.U. (7); 10 kilometers northeast of Villa Union, K.U. (1); 41.6 kilometers northeast of Villa Union, L.B.S.C. (8); 3.7 kilometers east of Villa Union, K.U. (15, 1 eggs), L.B.S.C. (4). Hyla stauSeri altae PANAMA: Canal Zone: 2.8 kilometers southwest of Fort Kobbe, K.U. (1); Road K2, T.N.H.C. (37); Summit, A.N.S.P. (4), F.M.N.H. (1), M.C.Z. (1), U.M.M.Z. (1). Chiriqui: 14.4 kilometers east of Concepcion, A.M.N.H. (3); David, A.M.N.H. (1); 6 kilometers north of David, A.M.N.H. (1); 2.8 kilo- meters south of David, A.M.N.H. (1). Code: 1 kilo- meter northeast of El Cano, K.U. (3); El Valle, A.M.N.H. (21), A.N.S.P. (1), K.U. (15); 6 kilome- ri4 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 ters south-southwest of Penononie, K.U. (13); 7 kilometers south-southwest of Penonome, K.U. (3). Los Santos: Tonosi, K.U. (5, 1 tadpoles). Panama: 5 kilometers south of Bejuco, A.M.N.H. (2); 2 kilo- meters west-southwest of Chepo, K.U. (9), 6 kilo- meters west-southwest of Chepo, K.U. (4); Nueva Gorgona, A.M.N.H. (2); 1.6 kilometers west of Nueva Gorgona, A.M.N.H. (4); 9 kilometers northeast of Pacora, K.U. (4); 1.5 kilometers west of Pacora, K.U. (5); Panama, K.U. (3); 2 kilometers north of Tocu- men, K.U. (7); 8 kilometers northeast of Tocumen, K.U. (1). Hyla staufferi staufferi MEXICO: Campeche: Balchacaj, U.I.M.N.H. (5); Champoton, M.C.Z. (2), U.M.M.Z. (8); 5 kilo- meters south of Champoton, K.U. (2); Ciudad Car- men, U.I.M.N.H. (1); Encarnacion, F.M.N.H. (11), U.I.M.N.H. (21); 6 kilometers west of Escarcega, K.U. (2); 7.5 kilometers west of Escarcega, K.U. (9); 13 kilometers west, 1 kilometer north of Escarcega, K.U. (6); Matamoras, F.M.N.H. (1); Pacaitun, Rio Candelaria, F.M.N.H. (2); Tres Brazos, U.I.M.N.H. (3); Tuxpeiia, U.M.M.Z. (1). Chiapas: No specific locality, U.M.M.Z. (I); Acacoyagua, U.S.N.M. (6); 2 kilometers west of Acacoyagua, U.M.M.Z. (5); 32 kilometers north of Arriaga, K.U. (4); 32 kilometers south of Arriaga, U.I.M.N.H. (2); A.sunci6n, C.A.S. (1), U.I.M.N.H. (5); Berriozabal, U.M.M.Z. (1); Buena Vista, U.M.M.Z. (7); El Real, 34 kilometers northeast of Altimirano, T.C.W.C. (1); 6 kilometers northeast of Escuintla, U.M.M.Z. (1); 3 kilometers east of Finca Juarez, U.I.M.N.H. (22); 4 kilometers north of Ixtapa, K.U. (6); 3 kilometers southwest of Las Cruces, K.U. (1); 17 kilometers south of Las Cruces, K.U. (2); 24 kilometers south of Las Cruces, K.U. ( 1 tadpoles ) ; 25 kilometers east, 5.6 kilometers north of Ocozocoautla, U.I.M.N.H. (4); west of Oco- zocoautla, U.I.M.N.H. (14); Palenque, U.I.M.N.H. (4), U.S.N.M. (24); 1.6 kilometers south of Pichu- cualo, U.I.M.N.H. (1); Puerto Arista, U.I.M.N.H. (11); 8.5 kilometers north of Puerto Madero, K.U. (5); Rancho Monserrate, U.I.M.N.H. (3); Rancho San Bartolo, U.I.M.N.H. (13); Region Soconusco, U.I.M.N.H. (2); 2 kilometers south of Rio de las Sal- mas, U.M.M.Z. (5); 11 kilometers south of Tapachula, K.U. (8); Tonala, U.I.M.N.H. (7); 8 kilometers north- west of Tonala, T.N.H.C. (2); 6 kilometers west of Tu.xtla Gutierrez, U.M.M.Z. (3). Guerrero: Acapulco, U.M.M.Z. (2); 9 kilometers northwest of Acapulco, U.F. (1); 13 kilometers north of Acapulco, T.N.H.C. (2); 5 kilometers west of Bajos del Ejido, U.M.M.Z. (1); El Limoncita, F.M.N.H. (3), U.I.M.N.H. (10); Laguna Coyuca, A.M.N.H. (1); Me.\cala, U.I.M.N.H. (1); 1.6 kilometers north of Organos, U.I.M.N.H. (1); Puerto Marques, A.M.N.H. (4); 5 kilometers east of Zacatula, U.M.M.Z. (1). Morelos: 3 kilometers south, 8.8 kilometers east of Cuernavaca, T.C.W.C. (1). Oaxaca: Chivela, A.M.N.H. ( 1 ); 11,8 kilometers south of Chivela, U.M.M.Z. (1); Huilotepec, A.M.N.H. (5); Juchitan, U.M.M.Z. (3); La Mata, U.I.M.N.H. (1); La Venta, U.I.M.N.H. (10); 78 kilometers north of La Venta, T.N.H.C. (22); Loma Bonita, F.M.N.H. (1); Matias Romero, A.M.N.H. (3); 6.6 kilometers north of Matias Romero, U.I.M.N.H. (1); Nisa Pipi, 8 kilometers northwest of Tehuantepec, U.M.M.Z. (100); 3 kilometers north of Pochutla, K.U. (22, 4 skeletons); 13.4 kilometers north of Pochutla, U.M.M.Z. (1); 1.3 kilometers south of Pochutla, U.M.M.Z. (1); 2.5 kilometers south of Pochutla, K.U. (3); 5 kilometers south of Pochutla, K.U. (7); San Geronimo, U.I.M.N.H. (1); 4.4 kilometers north of Sarabia, U.M.M.Z. (11); 6 kilometers south of Sarab- ia, U.M.M.Z. (2); Sierra Madre, north of Zanatepec, U.I.M.N.H. (2); 1,6 kilometers east of Tapanatepec, U.I.M.N.H. (40); 3 kilometers east of Tapanatepec, K.U. (26); 4.3 kilometers southwest of Tapanatepec, U.I,M,N,H. (1); 17.6 kilometers west-northwest of Tapanatepec, K.U. (2); Tehuantepec, A.M.N.H. (14), U.I.M,N,H, (26), U,M,M,Z. (36), U,S.N.M. (18); 3 kilometers east of Tehuantepec, U.M.M.Z. (9); 8 kilometers northeast of Tehuantepec, A.M,N,H, (1); 4 kilometers west of Tehuantepec, U.M.M.Z. (1); Temascal, U.S.C, (8); 3 kilometers south of Tolocita, K.U. (2); Tu.xtepec, K.U. (9, 1 tadpoles), U.I.M.N.H. (60); 3 kilometers south of Tuxtepec, U.I.M.N.H. (41); 13 kilometers south of Tuxtepec, U.I.M.N.H. (1); 17 kilometers south of Tuxtepec, K.U. (3); 21 kilometers south of Tuxtepec, U,I,M.N,H. (8); 27 kilometers south of Tuxtepec, U.I.M.N.H. (3); 3 kilo- meters south of Ubero, U.M.M.Z. (1); 1 kilometer north of Valle Nacional, U.I.M.N.H. (1); 1 kilometer west of Zanatepec, K.U. (1 tadpoles); 7 kilometers west-northwest of Zanatepec, U.M.M.Z. (6). Ptiebla: San Diego, A.M.N.H. (1); 30 kilometers northeast of Villa Juarez, T.N,H,C, (2). Quintana Roo: Coba, U.M.M.Z. (1); Isla Cozumel, 3.5 kilometers north of San Miguel, K.U. (3). San Luis Potosi: Ciudad Valles, A.M.N.H. (1), U.I.M.N.H, (1). Tabasco: Teapa, U.M.M.Z. (13); 10 kilometers north of Teapa, U.M.M.Z. (1); 24 kilometers north of Teapa, U.M.M.Z. (6); 27 kilometers north of Teapa, U.M.M.Z. (1); Tenosique, U.I.M.N.H. (1); 3.5 kilo- meters south of Villahermosa, U.M.M.Z. (2); 17 kilo- meters south of Villahermosa, U.M.M.Z. (8). Tamau- lipas: 6 kilometers southeast of Altamira, U.I.M.N.H. (2); Antiguo Morelos, U.I.M.N.H. (1); 1.6 kilometers east of Chamal, U.M.M.Z. (1); 19 kilometers south of Ciudad Mante, T.N.H.C. (3); 25 kilometers north of El Limon, U.I.M.N.H, (1); 36.2 kilometers north of El Limon, U.I.M.N.H. (4); between El Limon and Llera, U.M.M.Z. (2); Gomez Farias, U.M.M.Z. (3); 5 kilometers southeast of Gomez Farias, U.M.M.Z. ( 1 ) ; Pano Ayuctle, 8 kilometers northeast of Gomez Farias, U.M.M.Z. (4); Rio Frio, 8 kilometers west of San Gerardo, U.M,M.Z. (5); 5 kilometers west of San Gerardo, U.M.M.Z. (7); 4 kilometers southeast of Tres Marias, U.I.M.N.H. (12). Veracruz: 21 kilo- meters north of Acayucan, U.I.M.N.H. (1); 6 kilo- meters northwest of Acayucan, U.M.M.Z. (7); below .\cultzingo, U.M.M.Z. (2); 1.6 kilometers east-south- east of Alvarado, U.M.M.Z. (2); 8.4 kilometers west of Alvarado, U.M.M.Z. (3); 24.5 kilometers northwest of Alvarado, U.I.M.N.H. (3); 2.5 kilometers south- 1970 DUELLMAN: HYLID FROGS 715 southwest of Amatitlan, U.M.M.Z. (3); 5 kilometers south of Aquilera, U.M.M.Z. (1); Arroyo de las Pal- mas, 10 kilometers north of Cordoba, U.M.M.Z. (1); Boca del Rio, U.I.M.N.H. (21); 1.6 kilometers south of Boca del Rio, U.M.M.Z. (2); 11 kilometers south of Boca del Rio, U.M.M.Z. (7); 3 kilometers south- west of Boca del Rio, K.U. (1); 5 kilometers south- west of Boca del Rio, K.U. (1); 6 kilometers west of Boca del Rio, U.I.M.N.H. (1); 8 kilometers east of Cerro Gordo, T.C.W.C. (7); Ciudad Aleman, U.M.M.Z. (2); 8 kilometers soutlivvest of Coatza- coalcos, U.M.M.Z. (1); Cordoba, U.M.M.Z. (1); 5 kilometers east-southeast of Cordoba, A.M.N.H. (1), K.U. (1 tadpoles), T.N.H.C. (3), U.F. (1); 7 kilo- meters east-southeast of Cordoba, U.M.M.Z. (8); 3 kilometers west of Corral Nuevo, U.I.M.N.H. (1); 2.2 kilometers east of Cosaleacaque, U.M.M.Z. (8); Cosa- maloapan, U.M.M.Z. (4); west of Cotaxtla, U.I.M.N.H. (1); Cruz Blanca, U.I.M.N.H. (3); Cua- tulapan, K.U. (6), M.C.Z. (2), U.I.M.N.H. (16), U.S.N.M. (1); Cuatotolapan, U.M.M.Z. (30); El Chico, 11 kilometers south-southeast of Jalapa, F.M.N.H. (2); 3 kilometers north of El Tropido, U.I.M.N.H. (6); 6 kilometers east of Encero, U.I.M.N.H. (23); 1.6 kilometers east-northeast of Encinal, U.M.M.Z. (3); Hacienda Tamiahua, Cabo Rojo, K.U. (1); Huatusco, U.I.M.N.H. (5); 10 kilo- meters southeast of Hueyapan, U.M.M.Z. (1); Jalapa, U.I. (1); 6 kilometers southeast of Jalapa, U.M.M.Z. (1); 1.6 kilometers north of La Laja, U.I.M.N.H. (1); 21.6 kilometers south of Las Choapas, T.C.W.C. (2); 5 kilometers northwest of Lerdo de Tejada, LI.M.M.Z. (1); 17 kilometers east of Martinez de la Torre, U.I.M.N.H. (1); 6 kilometers west of Martinez de la Torre, U.I.M.N.H. (1); 2 kilometers east-northeast of Mata Oscura, K.U. (1); 3 kilometers northeast of Novillero, U.M.M.Z. (5); 5.4 kilometers northeast of Novillero, U.M.M.Z. (1); Orizaba, U.S.N.M. (1); Otatitlan, U.I.M.N.H. (5); Palma Sola, U.S.N.M. (1); Paso del Macho, U.I.M.N.H. (3), U.M.M.Z. (1); 5 kilometers southeast of Paso del Toro, K.U. (1); Potrero, M.C.Z. (3), U.I.M.N.H. (2), U.M.M.Z. (37), U.S.N.M. (3); Potrero Viejo, F.M.N.H. (4), K.U. (32); U.I.M.N.H. (16), U.M.M.Z. (69), U.S.N.M. (21); Presidio, U.S.N.M. (2); Puente Nacional, U.I.M.N.H. (2); Rodriguez Clara, U.I.M.N.H. (I); San Andres Tu.xtla, U.I.M.N.H. (1); 38 kilometers north of San Andres Tu.xtla, U.M.M.Z. (1); 10 kilo- meters east of San Juan de la Punta, M.C.Z. (1), U.I.M.N.H. (8), U.S.N.M. (15); Santiago Huatusco, U.M.M.Z. (2); Sauzla, U.M.M.Z. (4); 62 kilometers south of Tampico ( Tamaulipas ) , U.I.M.N.H. (10); 19 kilometers north of Tempoal, U.I.M.N.H. (I); Tierra Colorada, U.I.M.N.H. (3); Tula, U.I.M.N.H. (3); 2.7 kilometers northwest of Tula, U.M.M.Z. (2); Veracniz, A.M.N.H. (I); 5 kilometers south of Vera- cruz, U.M.M.Z. (3); 24 kilometers west of Veracruz, U.I.NLN.H. (17); Yanga, U.I.M.N.H. (2). BRITISH HONDURAS: Belize: Belize, F.M.N.H. (1). Cayo: 6 kilometers south of Cayo, M.C.Z. (2); San Augustin, U.M.M.Z, (8). Stann Creek: 10 kilo- meters east of Stann Creek, U.M.M.Z. (1); between Stann Creek and Roaring Creek, U.M.M.Z. (1); 5 kilometers south of Waha Loaf Creek, M.C.Z. ( 1 ). GUATEMALA: Alta Verapaz: Chinaja, K.U. (1); Cubilquitz, U.M.M.Z. (8). Baja Verapaz: 1 kilome- ter south of San Geronimo, U.M.M.Z. (21). Chiqui- mula: 1.6 kilometers southeast of Chiquimula, U.M.M.Z. (2); Esquipulas, U.M.M.Z. (18). El Petcn: Dolores, U.M.M.Z. (1); La Libertad, F.M.N.H. (2), K.U. (I), M.C.Z. (2), U.M.M.Z. (66), U.S.N.M. (2); Paso de Caballo, U.M.M.Z. (1); 1 kilometer south of Poptun, U.M.M.Z. (1); Sacluc, U.S.N.M. (1); Santa Teresa, U.M.M.Z. (4). Escuintla: Cuyuta, 20 kilo- meters north of San Jose, A.M.N.H. (8). Guatemala: 16 kilometers northeast of Guatemala, K.U. (1). Iza- bal: Puerto Barrios, T.C.W.C. (14), U.M.M.Z. (10); 2.5 kilometers northeast of Rio Blanco, K.U. (2). Jalapa: Jalapa, U.M.M.Z. (44). Jutiapa: Finca La Trinidad, U.M.M.Z. (28); JuUapa, U.M.M.Z. (2). Zacapa: 14 kilometers east-northeast of Mayuelas, K.U. (1); 7 kilometers east-northeast of Rio Hondo, K.U. (2, 1 tadpoles). EL SALVADOR: Cuscatldn: 8 kilometers west- northwest of Cojutepeque, T.N.H.C. (3); 11.5 kilo- meters west-northwest of Cojutepeque, T.N.H.C. (4). La Libertad: Quetzaltepeque, C.A.S. (1); 16 kilo- meters northwest of Santa Tecla, K.U. (2). La Union: 2.4 kilometers east of Santa Rosa, T.C.W.C. (2). Morazdn: Divisadero, U.S.N.M. (5). San Salvador: San Salvador, F.M.N.H. (6), K.U. (17, 1 eggs), U.M.M.Z. (7); 1.6 kilometers northwest of San Sal- vador, K.U. (2); 21.9 kilometers north of San Salva- dor, U.M.M.Z. (1). HONDURAS: Atlantidad: Ceiba, U.S.N.M. (1); Tela, M.C.Z. (1). Choltitcca: Choluteca, K.U. (6); 1.9 kilometers east of Choluteca, U.M.M.Z. (7); 3 kilometers east of Choluteca, K.U. (2); 6.2 kilome- ters east of Choluteca, K.LI. (11); 10 kilometers east of Choluteca, K.U. (1); 5 kilometers south of Cholu- teca. Colon: Lagima Ebano, F.M.N.H. (1), M.C.Z. (1); Patuca, U.S.N.M. (1). Comaijagua: 6.9 kilo- meters northwest of Siguatepeque, T.N.H.C. (I); 12 kilometers northwest of Siguatepeque, K.U. (1). Cortes: Lago de Yojoa, K.U. (7), M.C.Z. (1), T.C.W.C. (2). El Paraiso: Valle de Jamastran, A.M.N.H. (5). Francisco Morazdn: 8.6 kilometers northwest of Comayaguela, K.U. (1); El Zamorano, A.M.N.H. (10), K.U. (1), M.C.Z. (4); 29.3 kilome- ters north of Tegucigalpa, T.N.H.C. (2). NICARAGUA: Chinandega: Finca San Isidro, 10 kilometers south of Chinandega, K.U. (23). Ma- nagua: 13 kilometers east of Managua, K.U. (1), U.M.M.Z. (8); 2 kilometers south of Tipitapa, K.U. (5). Rivas: Rivas, M C.Z. (2); 9.5 kilometers south- east of Rivas, K.U. (1); 16 kilometers southeast of Rivas, M.C.Z. (12); 18 kilometers southeast of Rivas, K.U. (1); 7.7 kilometers northeast of San Juan del Sur, K.U. (8); 16.5 kilometers northeast of San Juan del Sur, K.U. (6); 5 kilometers southeast of San Pab- lo, K.U. (11). Zelaya: El Reereo, K.U. (19); Isla Grande del Maiz, K.U. (4); Wounta Haulover, A.N.S.P. (2). 716 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 COSTA RICA: Alajuela: Los Chiles, U.S.C. (3). Guanacaste: 4 kilometers west of Bagaces, U.S.C. (5); Finca Taboga, 20 kilometers southeast of Las Caiias, K.U. (2); 1.6 kilometers north of Guayaho de Bagaces, U.S.C. (3); Guardia, Rio Tempisque, U.S.C. (1); 10 kilometers north of Guardia, K.U. (2); 12 kilometers south of La Cruz, U.S.C. (1); Las Caiias, K.U. (1 skeleton); 27 kilometers north of Las Cafias, U.S.C. (5) Liberia, K.U. (13); 6 kilometers north of Liberia, U.S.C. (1); 8 kilometers north of Liberia, K.U. (1); 14.5 kilometers north of Liberia, U.S.C. (3); 14.5 kilometers south of Liberia, U.S.C. (5); 4 kilometers K.U. ( 1 ) ; 8.6 kilometers east-southeast of Playa del Coco, U.S.C. (14); 21 kilometers east-southeast of Playa del Coco, U.S.C. (2); Santa Cruz U.S.C. (2); Tenorio, K.U. (1); Tilaran, K.U. (1). Puntarenas: 6 kilometers east of Esparta, K.U. (1); 4 kilometers west-northwest of Esparta, K.U. (1); 10 kilometers west-northwest of Esparta, K.U. (8, 2 skeletons); 12 kilometers west-northwest of Esparta, K.U. (1); Hotel Maribella, K.U. (2); 10.8 kilometers north, 3 kilome- ters west of Puntarenas, T.C.W.C. (8). Hyla subocularis PANAMA: Darien: Laguna, K.U. (13); Rio Chucunaque, A.M.N.H. (1); Rio Chucunaque at first creek above Rio Tuquesa, A.M.N.H. (1); Rio Ucur- ganti, 7 kilometers above mouth, K.U. (1, 1 tadpoles); Tacarcuna, K.U. (45, 3 skeletons), U.M.M.Z. (1). Hyla sumichrasti MEXICO: Chiapas: 10 kilometers northeast of Los Amates, U.I.M.N.H. (4); 19 kilometers north of Arriaga, U.M.M.Z. (3, 2 tadpoles); El Sumidero, U.I.M.N.H. (1); Finca San Bartolo, U.I.M.N.H. (8); Ocozocoautla, U.I.M.N.H. (4); 4.5 kilometers north- east of Ocozocoautla, U.I.M.N.H. (18); 26 kilometers east, 5.6 kilometers north of Ocozocoautla, U.I.M.N.H. (3). Pitutal, south of Ocozocoautla, U.I.M.N.H. (6), T.C.W.C. (1); 2 kilometers northwest of Pueblo Nuevo Solistahuacan, K.U. (42, 3 skeletons); Tonola, U.I.M.N.H. (6). Oaxaca: Arroyo Palmar, U.I.M.N.H. (1); Cerro San Pedro del Istmo, U.I.M.N.H. (2); Cerro Santa Lucia, U.I.M.N.H. 11.8 kilometers south of Chivela, U.M.M.Z. (18, I skeleton); Llano Ocotal, C.A.S. (1), F.M.N.H. (12), U.I.M.N.H. (4); PorUllo Nejapa, A.M.N.H. (2), K.U. (13, 1 tadpoles); Santa Efigenia, U.S.N.M. (5); 16 kilometers east of Tapana- tepec, U.I.M.N.H. (I); Tres Cumbres, U.I.M.N.H. (I); between Zapotitlan and Huamelula, F.M.N.H. (3), U.I.M.N.H, (1). Hyla taeniopus MEXICO: Hidalgo: Tianguistengo, F.NLN.H. (4); 2.5 kilometers southwest of Tianguistengo, K.U. (3); 4 kilometers southwest of Tianguistengo, K.U. (I); 3 kilometers west of Xochicoatlan, K.U. (8, 2 skeletons). Pucbla: 8.7 kilometers southwest of Hua- chinango, U.M.M.Z. (1); 11.7 kilometers southwest of Huachinango, U.M.M.Z. (1); Rio Octapa, 3.7 kilo- meters north-northeast of Tezuitlan, K.U. ( 15, 4 skeletons, 1 tadpoles); 1.6 kilometers west of Teteles, T.N.H.C. (1); 8 kilometers northeast of Tezuitlan, K.U. ( 1 ); 1.5 kilometers southwest of Tlatlauquitepec, K.U. (I); 3 kilometers northwest of Zacapoa.xtla, U.M.M.Z. (2). Veracruz: Barranca Texola, 16 kilo- meters southeast of Jalapa, L'.I.M.N.H. ( 1 ); Huatusco, K.U. (1); 3 kilometers southwest of Huatusco, K.U. (3), U.M.M.Z. (3, 1 skeleton); 7.5 kilometers south- west of Huatusco, U.M.M.Z. (9, 1 skeleton); Jalapa, B.M.N.H. (1); 2 kilometers west of Jico, K.U. (5), U.M.M.Z. (2). Hyla thorectes MEXICO: Oaxaca: 30 kilometers north of San Gabriel Mixtepec, K.U. (1); 37 kilometers north of San Gabriel Mi.xtepec, K.U. (11, 2 skeletons, 3 tad- poles, 3 eggs), U.M.M.Z. 10. Hyla thysanota PANAMA: Darieu: Cerro Mali, U.S.N.M. (1). Hyla tica COSTA RICA: Alajuela: Cinchona, K.U. (9), M.C.Z. (3); 5 kilometers south of Ciudad Quesada, U.S.C. (1); Rio Maria-Aguilar, 3 kilometers west of Cariblanco, K.U. (1); 1.6 kilometers south of Zapote, U.S.C. (I); east slope of Volcan Poas, 21.3 kilometers north of Varablanca, U.M.M.A. (1). Cartago: Rio Playas, U.S.C. (4); Tapanti, K.U. (21, 1 skeleton, 2 tadpoles), U.S.C. (9); Volcan Turrialba, 1385 meters, U.M.M.Z. (5). Heredia: 2 kilometers north of Cinco Esquinas, K.L'. (1); San Jose de la Montaiia, U.M.M.Z. (1); 5.6 kilometers south of Varablanca, T.N.H.C. (1). Puntarenas: 1 kilometer northeast of Monteverde, U.S.C. (5); I kilometer west of Monte- verde, U.S.C. (1). San ]ose: 2 kilometers north of Las Nubes, K.U. (1); Rio Claro at Rio La Hondura, U.S.C. (10); Rio Tarrazu, 1 kilometer south of San Cristobal, U.S.C. (1); 15 kilometers nortli of San Isi- dro el General, K.U. (2). P.^N.^M.^: Chiriqui: south slope of Cerro Santa Catalina, 8 kilometers northwest of El Volcan, K.U. (1, 1 skeleton); Finca Bambito, 6 kilometers east- northeast of El Volcan, K.L'. ( I ); Finca Ojo de Agua, .southeast slope of Cerro La Pelota, K.U. (2); Finca Palosanto, 7 kilometers north-northwest of El Volcan, K.U. (7); Quebrada Chevo, south slope of Cerro La Pelota, K.U. (17); Rio Colorado, 17.5 kilometers northwest of El Volcan, K.U. (1); 14.5 kilometers north-northwest of El Volcan, K.U. (1); 16 kilometers north-northwest of El Volcan, K.U. ( 1 ). Hyla uranochroa COSTA RICA: Alajuela: Cinchona, K.U. (6, 2 skeletons, 8 tadpoles), U.S.C. (5); between Cinchona and Salto El Angel, U.S.C. (1); Ciudad Quesada, U.S.C. (1); San Carlos, U.S.N.M. (I); north .slope of Volcan Poas, 22.5 kilometers north of Varablanca, U.M.M.Z. (1 tadpole); 1.6 kilometers south of Zapote, U.S.C. (2). Cartago: Moravia de Turrialba, K.U. ( 10, 1970 DUELLMAN: HYLID FROGS 717 1 skeleton); 1 kilometer east of Pacayas, U.S.C. (1); 3 kilometers south of Pavones, K.U. (18, 1 tadpoles); Rio Chitaria, 3 kilometers north-northeast of Pavones, K.U. (1 tadpoles); Rio Izaquito, near Pavones, U.S.C. (1); 4.3 kilometers northeast of Rio Reventazon bridge, U.M.M.Z. (2 tadpoles); 3 kilometers north of Santa Rosa, K.U. (1 tadpoles); 1 kilometer north of Tapanti, U.S.C. (2); Turrialba, K.U. (1); Volcan Turrialba, U.M.M.Z. (1 tadpoles). Hcredia: Cari- blanco, K.U. (1); Hacienda Cayuga, 1 kilometer north of Montaiia Azul, K.U. (1 tadpole), Isla Bonita, F.M.N. H. (1), K.U. (2, 1 tadpoles); Montana Azul, K.U. (1 tadpoles); San Jose de la Montaiia, K.U. (1); 2.7 kilometers north of San Jose de la Montana, K.U. (2 tadpoles); 1.6 kilometers north-northeast of Uvita, U.S.C. (22). Limon: El Tigre, 12-20 kilometers southwest of Siquirres, U.S.C. (1); Pico Blanco, U.S.N.M. (1). Puntarenas: Esparta, M.C.Z. (1); Monteverde, U.S.C. (19). San ]ose: south slope of Cerro de la Muerte, 1.524 meters, U.S.C. (1); La Es- trella, M.C.Z. (1); 1 kilometer west of La Hondura, U.S.C. (1); La Palma, A.N.S.P. (11), M.C.Z. (1), U.M.M.Z. (1), U.S.C. (4), U.S.N.M. (1); Rio Tirivi, U.M.M.Z. (1); 14 kilometers north of San Isidro el General, L'.M.M.Z. (1); 1.5 kilometers north of San Isidro el General, K.U. (3, 2 skeletons), U.M.M.Z. (2); 18.5 kilometers north of San Isidro el General, K.U. (5, 1 tadpoles), U.M.M.Z. (1). PANAMA: Bocas del Tow: north slope of Cerro Pando, 1450 meters, K.U. (26, 2 skeletons, 2 tad- poles); La Loma, M.C.Z. (6); Rio Changena, 650 meters, K.U. (4); Rio Changena, 830 meters, K.U. (4). Hyla valancifer MEXICO: Veracruz: Volcan San Martin, K.U. (1);U.I.M.N.H. (1), U.M.M.Z. (2). Hyla walkeri MEXICO: Chiapas: 10 kilometers northwest of Comitan, K.U. (4); 14 kilometers northwest of Comi- tan, K.U. (3); 18 kilometers northwest of Comitan, K.U. (16, 2 skeletons, 1 tadpoles), M.C.Z. (2); El Suspiro, U.M.M.Z. ( 1 ); 2.5 kilometers south of Jitotol, K.U. (3, 1 tadpoles); 2 kilometers northwest of Pueblo Nuevo Solistahuacan, U.M.M.Z. (23); San Cristobal de las Casas, U.I.M.N.H. (2); 8.8 kilometers southeast of San Cristobal de las Casas, K.U. (1); 12.8 kilome- ters southeast of San Cristobal de las Casas, K.U. (2); 6.4 kilometers northwest of San Cristobal de las Casas, K.U. (1); 8.8 kilometers northwest of San Cristobal de las Casas, U.M.M.Z. (11); 30 kilometers northwest of San Cristobal de las Casas, U.M.M.Z. (20). GUATEMALA: El Quiche: La Primavera, be- tween Sacapulas and Santa Cruz Quiche, L'.M.M.Z. (30); Ututlan, U.M.M.Z. (1). Huehuetermngo: 3 kilometers north of San Juan I.xcoy, U.M.M.Z. (32), 4 kilometers east of San Juan l.\coy, U.M.M.Z. (5); Soloma, U.M.M.Z. (28). Jalapa: Aserradero San Lorenzo, U.M.M.Z. (5). Hyla xanthosticta COSTA RICA: Hcredia: south fork of Rio Las Vueltas, south slope of Volcan Barba, K.U. ( 1 ). Hyla zeteki COSTA RICA: Hcredia: Isla Bonita, K.U. (1); Varablanca, K.U. (1). San Jose: La Hondura, A.N.S.P. (5, 1 tadpoles); La Palma, K.U. (2), U.S.C. (2). PANAMA: Chiriqui: Boquete, M.C.Z. (1), U.M.M.Z. (12, 1 skeleton), U.S.N.M. (1). Hyla sp. GUATEMALA: Alta Verapaz: Finca Los Alpes, K.U. (2 tadpoles), U.M.M.Z. (1 tadpoles). Pachymedusa dacnicolor MEXICO: Colinw: No specific locality, F.M.N.H. (1), U.S.N.M. (1); CoUma, A.M.N.H. (3), M.C.Z. (1), S.U. (1), U.I.M.N.H. (1), U.M.M.Z. (47); 3.7 kilometers north of Colima, A.M.N.H. (3, 1 skeleton); 1.6 kilometers southwest of Colima, A.I.M.N.H. (1); Hacienda Albarradita, U.M.M.Z. (4, 1 eggs); east of Lo de Villa, A.M.N.H. (1); Manzanilla, U.M.M.Z. (1); 33 kilometers southeast of Manzanillo, A.M.N.H. (2), K.U. (1), U.M.M.Z. (1); Paso del Rio, U.I.M.N.H. (3); Periquillo, U.M.M.Z. (25); 1.6 kilo- meters south of Puebla Juarez, U.M.M.Z. (2); Que- seria, U.M.M.Z. (2); Rio Armeria, U.M.M.Z. (1); Santiago, U.M.M.Z. (5); 8 kilometers southwest of Tecolapa, U.M.M.Z. (5, 1 eggs); 4 kilometers north- west of Tecoman, U.M.M.Z. (1); 5-8 kilometers northwest of Villa Alvarez, L'.M.M.Z. (1). Guerrero: No specific locality, U.M.M.Z. (2); Acahuitzotia, K.U. (1), T.C.W.C. (1); U.M.M.Z. (1); Acapulco, A.M.N.H. (3), M.C.Z. (3), U.F. (1), U.M.M.Z. (1); 6.4 kilometers north of Acapulco, U.I.M.N.H. (1); 7.8 kilometers north of Acapulco, U.F. (1); 27 kilo- meters northeast of Acapulco, U.I. (3); 5 kilometers south of Buena Vista, A.M.N.H. (3); between Chilapa and Tixtla. K.U. (1); Chilpancingo, F.M.N.H. (1), M.C.Z. (25), U.M.M.Z. (7); Cocula, A.M.N.H. (2); 13.3 kilometers northwest of Coyuca, U.I.M.N.H. (1); Colonia Buenas Aires, 27 kilometers east of Tecpan, K.U. (1 skeleton), U.M.M.Z. (1); 1.6 kilometers southwest of Colotlipa, T.C.W.C. (6); 12 kilometers north of El Naranjo, U.M.M.Z. ( 1 ); 3 kilometers south of Garrapata, U.I.M.N.H. (1); Iguala, T.C.W.C. (4); 9 kilometers south of Mazatlan, U.I.M.N.H. (1); 8 kilometers north of Me.xcala, U.I.M.N.H. (1); 1.6 kilometers southeast of Mochitlan, T.C.W.C. (2); Mojonares, U.I.M.N.H. (8); Ocotito, T.C.W.C. (4), U.I.M.N.H. (1); 5.4 kilometers north of Ocotito, U.M.M.Z. (3); Ometepec, U.S.N.M. (1); 1.6 kilo- meters north of Organos, U.I.M.N.H. (2); Palo Blan- co, U.I.M.N.H. (3); 9.6 kilometers west of Pie de la Cuesta, U.M.M.Z. (3); Puerto Marquez, A.M.N.H. (2); Rincon, T.C.W.C. (1); Rio AguacaUllo, 30 kilo- meters north of Acapulco, T.C.W.C. (1); 1.7 Idlome- 118 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 ters south of San Andreas de la Cniz, K.U. ( 1 ) ; 17 kilometers south of Taxco, T.C.W.C. (3); 21 kilo- meters south of Taxco, T.C.W.C. (4); 32 kilometers east-southeast of Tecpan, U.I.M.N.H. (1); Tierra Colorada, U.S.N.M. ( 1 ); 10 kilometers north of Tierra Colorada, U.F. ( 1 ); 1.6 kilometers southwest of Tierra Colorada, T.C.W.C. (1); 8 kilometers southwest of Tierra Colorada, T.C.W.C. (7); 2 kilometers east of Tixtla, K.U. (7), T.N.H.C. (1); 5 kilometers east of Tixtla, U.F. (3). Jalisco: 5 kilometers northeast of Autlan, U.I.M.N.H. (2); 15 kilometers northwest of Cihuatlan, K.U. (1); 3 kilometers north of La Reso- lana, U.M.M.Z. (2), 6.6 kilometers northeast of La Resolana, K.U. (2); 32 kilometers southwest of La Resolana, K.U. (1); 16 kilometers southeast of Las Anonas, T.C.W.C. (5); 8 kilometers east of Melaque, K.U. (6); 3 kilometers west of Tamazula, A.M.N.H. (2). Michoacdn: Aquililla, U.M.M.Z. ( 13); Apatzin- gan, F.M.N.H. (1), U.M.M.Z. (2); 1.6 kilometers east of Apatzingan, U.M.M.Z. (3); 10.4 kilometers east of .\pat7.ingan, U.NLM.Z. (1); 14.4 kilometers east of Apatzingan, U.M.M.Z. (1); 23 kilometers south of Apatzingan, K.U. (1); 2 kilometers soutli of Chara- pendo, U.M.M.Z. (1); Coahuayana, U.M.M.A. (3); Coalcoman, K.U. (5, 1 skeleton), U.M.^LZ. (.55); El Sabino, U.I.M.N.H. (1); Huetamo road, U.I.M.N.H. (2); La Placita, U.M.M.Z. (1); 32 kilometers east of Nueva Italia, U.M.M.Z. (2); between Rio Marquez and Cuatro Caminos, K.U. (2); Salitre de Estopilas, U.M.M.Z. (2). Morelos: Alpuyeca, T.C.W.C. (1); 4 kilometers south of Alpuyeca, T.C.W.C. (6); 12 kilo- meters northwest of Axochiapan, T.C.W.C. (1); 3.5 kilometers west of Cuautlixco, K.U. (10, 4 skeletons); 5 kilometers northwest of Cuautlixco, U.M.M.Z. (2); 14 kilometers south of Cuernavaca, U.M.M.Z. (1); El Rodeo, T.C.W.C. (15); 2 kilometers south of Jonacatepec, T.C.W.C. (7); Puente de I.xtla, U.I.M.N.H. (1); 1 kilometer northeast of Puente de Ixtla, K.U. (3); T.C.W.C. (21); 19 kilometers south of Puente de Ixtla, U.I.M.N.H. (2); Temilpa, T.C.W.C. (4); 1.6 kilometers south of Temixco, U.M.M.Z. (2); Tequesquitengo, A.M.N.H. (1); Zaca- tepec, T.C.W.C. (12); 3 kilometers west of Zacatepec, T.C.W.C. (10). Nayarit: Acaponeta, A.M.N.H. (2), U.S.N.M. (2), T.C.W.C. (1); 30-50 kilometers south of Acaponeta, A.M.N.H. (2); 6.4 kilometers north of Compostela, A.M.N.H. (7); 6 kilometers south of Ixtlan del Rio, U.M.M.Z. (1); Jesus Maria, A.M.N.H. (1); 8.6 kilometers northeast of Navarrete, U.F. (1); 10 kilometers south-southwest of Navarrete, M.V.Z. (1); Peiiitas, A.M.N.H. (4); Rosaniorada, A.M.N.H. (4); San Bias, A.M.N.H. (8), K.U. (4), U.I.M.N.H. (4), U.M.M.Z. (1); 2 kilometers north of San Bias, S.U. (2); 23 kilometers east of San Bias, U.I.M.N.H. (5); 8.6 kilometers south-southeast of San Bias, U.M.M.Z. (4); 1.6 kilometers southwest of San Jose del Conde, U.M.M.Z. (1); San Juan Peyotan, L.A.C.M. (1 tadpoles); 5 kilometers north of Santa Isabela, U.M.M.Z. (1); Santiago Escuintla, A.M.N.H. (2); Tepic, U.I.M.N.H. (2), U.M.M.Z. (6); 3 kilo- meters south of Tepic, A.M.N.H. (10); 5.5 kilometers south of Tepic, A.M.N.H. ( I ); 37 kilometers northwest of Tepic, T.C.W.C. ( 1 ); 4 kilometers east of Tuxpan, K.U. (2); 11 kilometers southeast of Tuxpan, U.I.M.N.H. (2). Oaxaca: Chacalapa, K.U. (1); 4.2 kilometers north of Chacalapa, U.F. (2); Escurano, U.I. (1); La Candelaria, K.LI. (4, 1 eggs); Mirador, A.M.N.H. (1); Pochutla, U.I.M.N.H. (8); 3 kilome- ters north of Pochutla, K.U. (1); 32.9 kilometers north of Pochutla, U.M.M.Z. (21), 41.4 kilometers north of Pochutla, U.M.M.Z. (1); 2.0 kilometers south of Pochutla, K.U. (3); U.M.M.Z. (10), 5 kilometers south of Pochutla, K.U. (4); Tehuantepec, U.S.N.M. ( 1 ) . Sinaloa: 8 kilometers north of Carrizalejo, K.U. (15); Chele, U.M.M.Z. (1); Concepcion, K.U. (1); Concordia, A.M.N.H. (1); 5 kilometers southwest of Concordia, K.U. (5), 3.2 kilometers southwest of Copala, K.U. (1); Costa Rica, 25 kilometers south of Culiacan, U.I.M.N.H. (2); Culiacan, U.M.M.Z. (1); 3 kilometers north of Culiaciin, A.M.N.H. ( 1 ); 12 kilo- meters north of Culiacan, K.U. (1), U.M.M.Z. (1); 21 kilometers south of Culiacan, M.V.Z. (2); Eldora- do, A.M.N.H. (2); 1.6 kilometers northeast of El Fuerte, F.M.N.H. (22); 13 kilometers northeast of El Fuerte, F.M.N.H. (1); Elota, K.U. (1); Escuinapa, A.M.N.H. (3); 22 kilometers southeast of Escuinapa, T.C.W.C. (1); 25 kilometers southeast of Escuinapa, U.M.M.Z. (5); El Venadillo. U.M.M.Z. (1); Guana- caste, 1.6 kilometers southwest of Palmar, M.V.Z. (1); Isia Palmito del Verde, middle, K.U. ( 1 ); 5 kilometers northeast of Las Trancas, K.U. (4); 5 kilometers north of Los Mochis, K.U. (1); 21 kilometers north-north- east of Los Mochis, U.I.M.N.H. (3); 7.3 kilometers southwest of Matatiin, K.U. (9); Mazatkin, A.M.N.H. (1), U.I.M.N.H. (10), U.S.N.M. (1); 3-40 kilometers north-northwest Mazatkin, A.M.N.H. (3), A.N.S.P. (2), M.C.Z. (12), M.V.Z. (11), U.M.M.Z. (6); 3 kilometers east of Mazatlan, T.C.W.C. (1); 1 kilo- meter southeast of Mazatkin, M.C.Z. (1); 4 kilometers southeast of Navolato, K.U. (I); Presidio, S.U. (1), U.S.N.M. (1); Rosario, U.I.M.N.H. (10), U.S.N.M. (1); 13 kilometers west-northwest of Rosario, U.M.M.Z. (2); San Franci.squito, A.M.N.H. (1); 5 kilometers southwest of San Ignacio, K.U. (4, 3 tad- poles); Teacapiin, K.U. (1); Villa Union, K.U. (37, 1 tadpoles), S.U. (6), U.M.M.Z. (1); 9.1 kilometers northeast of Villa Union, K.U. (4); 3.7 kilometers east of Villa Union, K.U. (1). Soiwra: Alamos, A.M.N.H. (5), K.U. (2); Guiracoba, A.NLN.H. (5), M.V.Z. (11); 2.5 kilometers north of Navajoa, U.M.M.Z. (1); 5 kilometers northwest of Navajoa, U.M.M.Z. (1); Presa Obregon, K.U. (1); Rio Alamos, 14.4 kilome- ters southeast of Alamos, K.U. (15). Plirynohyas venulosa MEXICO: Campcchc: Beain, M.C.Z. (1), U.NLM.Z. (1); Champoton, U.M.M.Z. (4); 5 kilo- meters south of Champoton, K.U. (5); 2.5 kilometers west of Esc;ircega, K.U. (1); 7.5 kilometers west of Esc;ircega, K.U. (2), U.M.M.Z. (1); 12 kilometers west of Esciircega, K.U. (1); 13 kilometers west, 1 kilometer north of Esc;ircega, K.U. (3); La- guna Silvituc, K.U. (1); Pacaitun, Rio Candelaria, F.M.N.H. (2); Ruinas Edzna, K.U. (I); Tres Brazos, 1970 DUELLMAN: HYLID FROGS 719 F.M.N.H. (1), U.I.M.N.H. (1). Chiapas: Acacoya- gua, U.S.N.M. (1); Colonia Soconusco, U.S.N.M. (2); Cruz de Piedra, U.S.N.M. (10), Escuintla, U.M.M.Z. (10); La Esperanza, U.M.M.Z. (2), U.S.N.M. (3); 8 kilometers north of Puerto Madero, U.M.M.Z. (1); 18 kilometers south of Teapa ( Tabasco ), U.I.M.N.H. ( 1 ) . Colima: 1.6 kilometers north of Coiinia, U.M.M.Z. ( 1 ); 11-32 kilometers northwest of Manzanillo, M.V.Z. ( 1 ) ; Paso del Rio, U.M.M.Z. ( 1 ) ; Rio Astillero, C.A.S. (1). Guerrero: La Venta, F.M.N.H. (3); Puerto \hirquez, A.M.N.H. (2). Michoacdii: Barranca de Bejuco, U.M.M.Z. (1). Nayarit: 74 kilometers south of Esquinapa (Sinaloa), K.U. (2); east of San Bias, U.I.M.N.H. (1); 8 kilometers east of San Bias, U.I.M.N.H. (1); 23 kilometers east of San Bias, K.U. (1); 29 kilometers north-northwest of Tepic, U.F. (2); 34 kilometers north-nortliwest of Tepic, K.U. (1 tadpoles), L.B.S.C. (16, 1 skeleton). Oaxaca: Matias Romero, A.M.N.H. (4); 45 kilometers north of Matias Romero, U.I.M.N.H. (1); Tapanatepec, M.C.Z. (3); Temascal, U.S.C. (1); Tu.xtepec, U.S.N.M. ( 1 ); between Zanatepec and Tapanatepec, U.I.M.N.H. (1). Quintana Roo: 4 kilometers north-northeast of Felipe Carrillo Puerto, K.U. (1); 8 kilometers west of Puerto Juarez, K.U. (1); 13 kilometers west of Puerto Juarez, K.U. (1). Sati Luis Potosi: 1 kilometer east of El Naranjo, U.M.M.Z. (2); Pujal, L.S.U. (1); Rio Coy near Pujal, U.S.N.M. (1); Tamazunchale, U.M.M.Z. (1); 4 kilometers north of Tamazunchale, U.M.M.Z. (1); 29 kilometers east of Tamuin, U.F. (1); 16 kilometers south of Valles, A.M.N.H. (1); 16 kilometers northwest of Xilitia, A.M.N.H. (1). Sina- loa: Presidio, B.M.N.H. (2). Tabasco: Cardenas, U.M.M.Z. (1); 60 kilometers west of Cardenas, K.U. (3); 4 kilometers northeast of Comalcalco, A.M.N.H. (3); Frontera, B.M.N.H. (2); 24 kilometers east of Frontera, M.C.Z. (1); 2.3 kilometers northeast of Huimanguillo, U.M.M.Z. (1); 5 kilometers north of Teapa, U.M.M.Z. (2); 13 kilometers north of Teapa, U.M.M.Z. (1); 21 kilometers north of Teapa, U.M.M.Z. (4); 25 kilometers north of Teapa, U.M.M.Z. (1); 29 kilometers north of Teapa, U.M.M.Z. (3); Tenosique, U.S.N.M. (1); 10 kilome- ters north, 24 kilometers west of Villahermosa, K.U. (2); 3.5 kilometers south of Villahermosa, U.M.M.A. (3). Tainaulipas: 6 kilometers north of El Mante, U.M.M.Z. (1); Rio Sabinas, 5 kilometers northeast of Gomez Farias, U.M.M.Z. (1); Tampico, B.M.N.H. (5). Veracruz: 29 kilometers southeast of Alvarado, U.M.M.Z. (14); 38.4 kilometers southeast of Alvarado, U.M.M.Z. (1); 2.5 kilometers south-southwest of Amatitlan, U.M.M.Z. (12); Barranca Metlac, K.U. (1 skeleton ) ; 5 kilometers southwest of Boca del Rio, K.U. (6), U.I.M.N.H. (5); 6.6 kilometers southwest of Boca del Rio, K.LI. (5); 7 kilometers north-north- west of Cerro Gordo, K.U. (1); Chacaltianguis, U.M.M.Z. (1); Cienega de Macuile, U.M.M.Z. (13); Ciudad Aleman, U.M.M.Z. (6); Cordoba, U.S.N.M. (2); 10 kilometers north of Cordoba, U.M.M.Z. (1); 17 kilometers east-southeast of Cordoba, T.N.H.C. (1); 21 kilometers north of Corral Nuevo. U.I.M.N.H. (12); Cosamaloapan, U.M.M.Z. (1); Cuatotolapan, U.M.M.Z. (2); Cuautlapan, F.M.N.H. (2), K.U. (48), U.M.M.Z. (50); 11 kilometers east of Ebano (San Luis Potosi), T.N.H.C. (2); El Potrero, M.C.Z. (2); Encinal, K.U. (1 tadpoles); 5.5 kilometers east- northeast of Encinal, U.M.M.Z. (1); Huatusco, U.I.M.N.H. (2); Jalapa, B.M.N.H. (1); 19 kilometers east of Jalapa, U.I.M.N.H. (3); 20 kilometers east- northeast of Jesus Carranza. K.U. (2); 20 kilometers south of Jesus Carranza, K.U. (4); 10 kilometers north of Jose Cardel, M.V.Z. (4); La Laja, U.I.M.N.H. (22); 2 kilometers northwest of Lerdo de Tejada, U.M.M.Z. (2); 16 kilometers west-northwest of Los Conejos, K.U. (3); 6 kilometers west of Martinez de la Torre, U.I.M.N.H. (1); Minatitlan, A.M.N.H. (1); 5 kilometers west-southwest of Minatitlan, U.M.M.Z. (1); Misantla, B.M.N.H. (5); 1.6 kilometers northeast of No\illero, U.M.M.Z. (1); 5 kilometers northeast of Novillero, U.M.M.Z. (1); Ozuluama, K.U. (1); Panu- co, M.C.Z. (1), U.M.M.Z. (6); 5.5 kilometers north- east of Panuco, T.N.H.C. (1); Paraje Nuevo, U.M.M.Z. (16); Paso del Macho, U.I.M.N.H. (4), U.M.M.Z. (4); Peiiuela, A.M.N.H. (1); Potrero Viejo, F.M.N.H. (1), K.U. (10), U.I.M.N.H. (2), U.M.M.Z. (23); Rodriguez Clara, F.M.N.H. (1); Salinas, T.C.W.C. (1); 20 kilometers northwest of San Andres Tu.xtla, U.M.M.Z. ( 1 ); 48 kilometers northwest of San Andres Tu.xtla, U.M.M.Z. (1); San Isidro, K.U. (1); 3 kilometers west of San Marcos, K.U. (3); Sausal, U.M.M.Z. (9); Tecolutla, U.I.M.N.H. (13); 5 kilo- meters south of Tehuatlan, K.U. (2); Tierra Colora- da, F.M.N.H. (1), U.I.M.N.H. (1), To.xtlacuaya, F.M.N.H. (2); Tuxpam, A.M.N.H. (2), K.U. (1); Veracruz, A.M.N.H. (1), I.P.N. (1), U.F. (10), U.M.M.Z. (3); 6 kilometers west-southwest of Zacu- alpilla, K.U. (1). Yucatan: Chichen Itza, F.M.N.H. (6), U.M.M.Z. (2). BRITISH HONDURAS: Orange Walk: 3 kilo- meters south of Corozal, M.C.Z. ( 1 ). GUATEMALA: El Peten: La Libertad, F.M.N.H. (2), M.C.Z. (1), U.M.M.Z. (30), U.S.N.M. (2). Escuintla: Cuvuta, A.M.N.H. (72). Retalhuleu: Caballo Blanco, F.M.N.H. (1); Casa Blanca, U.M.M.Z. (1). Suchitepequez: Mazatenango, C.A.S. (6). Za- capa: 23 kilometers west of Zacapa, T.C.W.C. (20). HONDURAS: Comayagua: La Mision, A.M.N.H. (1), M.C.Z. (1); 8 kilometers above La Mision, M.C.Z. (1). Cortes: Agua Azul, A.M.N.H. (2); 7 kilometers southwest of La Lima, K.U. (1); 1.6 kilo- meters west of La Lima, T.C.W.C. (5). El Pariso: Valle de Jamastran, A.M.N.H. ( 1 ). NICARAGUA: No specific locality, U.S.N.M. (2). Chinandega: Finca San Isidro, 10 kilometers south of Chinandega, K.U. (4). Managua: 8 kilome- ters northwest of Managua, K.U. (2). Matagalpa: Hacienda La Cumplida, K.U. (1). Zelaya: Wounta Haulover, A.N.S.P. (1). COSTA RICA: Alajuela: Los Chiles, U.S.C. (4). Guanacaste: Bebedero, B.M.N.H. ( 1 ); Finca Tabogo, 20 kilometers southeast of Las Caiias, K.U. (1); Hacienda La Mojica, 3 kilometers south, 18 kilometers west of Las Caiias, T.C.W.C. (8); Las Huecas, 720 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 U.M.M.Z. (1); 17 kilometers north of Liberia, U.S.C. (1); 20 kilometers north of Liberia, U.S.C. (5); 33 kilometers north of Liberia, U.S.C. (1); Rio Tenorio, 5 kilometers south, 16 kilometers west of Las Caiias, T.C.W.C. (1). Funiarenas: 4 kilometers west-north- west of Esparta, K.U. (26, 5 skeletons); Palmar Norte, K.U. (S, 1 skeleton); Palmar Sur, K.U. (3); Parrita, U.S.C. (1). PANAMA: Canal Zone: No specific locality, M.C.Z. (1). Ancon, A.N.S.P. (1); La Pita, M.C.Z. (1); Madden Dam, S.U. (1); Madden Forest, K.U. (2). Chiriqui: Suma, Rio San Pablo, A.M.N. H. (1). Code: AguaduJce, K.U. (1); Santa Clara, F.M.N.H. (1). Colon: Ciricito, C.A.S. (1). Darien: El Real, K.U. (2); Rio Canclon at Rio Chucunaque, U.M.M.Z. (12); Rio L'curganti, 7 kilometers above mouth, K.U. ( 1 ) . Panama: 5 kilometers south of Bejuco, A.M.N.H. (2); 10 kilometers west-southwest of Chepo, K.U. (16); Nueva Gorgona, A.M.N.H. (3, 1 tadpoles), K.U. (1 tadpoles); Punta Paitilla, M.C.Z. (1); Tapia, A.M.N.H. (1). ters south of San Mateo Lxtatan, K.U. (1, 1 tadpoles); 3 kilometers south of Paqui.x, U.M.M.Z. (7); 8 kilo- meters south of Paquix, K.U. (Ill, 9 skeletons, 3 tadpoles); M.C.Z. (3); 1.5 kilometers east of San Mateo lxtatan, K.U. (1 tadpoles); Todos Santos, U.M.M.Z. (2); 2.5 kilometers north of Toquia, K.U. (2). Jalapa: 8 kilometers east of Mataquescuintla, La Soledad Grande, F.M.N.H. (11). Quetzaltenango: 37 kilometers .southeast of Malacatancito, K.U. ( 1 tadpoles ) ; 6 kilometers north of San Carlos Sija, K.U. (7). San Marcos: 1 kilometer east of Ixchiguan, U.M.M.Z. (1 tadpoles); 5 kilometers west of Ixchi- guan, U.M.M.Z. (1 tadpoles); 2 kilometers northwest of Ixhiguan, M.C.Z. (1), U.M.M.Z. (6); Volcan Tajamuico, F.M.N.H. (1). Solold: Los Encuentros, U.M.M.Z. (13). Totonicapdn: Desconsuelo, U.M.M.Z. (1); Maria Tucuni, U.M.M.Z. (1, 2 tad- poles), K.U. (1 tadpoles); 13.4 kilometers north of San Carlos Sija, K.U. (2). EL SALVADOR: M.V.Z. (1). Clialatenango: Los Esemiles, Phyllomedusa lemur COSTA RICA: Alajuela: Cinchona, K.U. (8, 1 tadpoles); 5 kilometers south of Giudad Quesada, U.S.C. (1). Cartago: La Suiza, U.S.C. (1); Moravia, K.U. (22, 2 skeletons); Tapantl, K.U. (53, 4 skele- tons), U.S.C. (1); 10 kilometers north of Rio Reven- tazon bridge, U.S.C. (1). Hcrcdia: Cariblanco, M.C.Z. (1); Isla Bonita, F.M.N.H. (1). Limon: El Tigre, 12-20 kilometers southwest of Siquirres, K.U. (1 tadpoles), LI.S.C. (3, 1 tadpoles); junction of Rio Lari and Rio Dipari, 21 kilometers south of Amubre, U.S.C. (1). Puntarenas: 3.6 kilometers east of Mon- teverde, U.M.M.Z. (I), U.S.C. (1). San Jose: La Palma, A.N.S.P. (2), K.U. (22), M.C.Z. (2), U.M.M.Z. (5), U.S.C. (4), U.S.N.M. (1); Rio Claro at Rio Hondura, U.S.C. ( 1 ). PANAMA; Bocas del Tow: Rio Changena, 650 meters, B.Y.U. (2), K.U. (5); Rio Changena, 830 meters, K.U. (1); Rio Claro near junction with Rio Changena, 910 meters, K.U. (7). Code: El Valle, A.N.S.P. (1). Darien: Cerro Mali, U.S.N.M. (1). Panama: Cerro La Campana, K.U. (3). Phyllomedusa venusta PANAMA: Darien: Rio Tuira at Rio Mono, K.U. (4, 1 skeleton). Plectrohyla avia MEXICO: Chiapas: El Chiciquite, Volcan Ta- cana, U.I.M.N.H. (1); Region de Soconusco, K.U. (I skeleton), U.I.M.N.H. (1); Volcan Tacana, 8 kilo- meters north of Union Juarez, K.U. (2). GUATEMALA: Quetzaltenango: Granja Lorena, U.M.M.Z. (I). Plectrohyla glandulosa GUATEMALA: No .specific locality, B.M.N.H. (2). Huehuetenango: Laguna de Vejcha, 5.5 kilome- Plectrohyla guatemalensis MEXICO: Chiapas: Chicomuselo, U.M.M.Z. (2); El Chiciquite. Volcan, Tacana, U.I.M.N.H. (1); Letrero, U.M.M.Z. (1); 3.6 kilometers south of Rayon Mesdalapa, K.U. (1 tadpoles); 5.6 kilometers south of Rayon Mescalapa, K.U. ( 1, 2 tadpoles); 6.2 kilo- meters south of Rayon Mescalapa, K.U. (7, 1 skele- ton, 2 tadpoles); Region de Soconusco, U.I.M.N.H. ( I ) ; Rio Hondo, 9.5 kilometers south of Pueblo Nuevo Solistahuacan, K.U. (1); 18 kilometers north of Pueblo Nuevo Solistahuaciin, K.U. (8), U.M.M.Z. (4); San Cristobal de las Casas, A.M.N.H. (1), U.I.M.N.H. (1); 10 kilometers southeast of San Cris- tobal de las Casas, M.V.Z. (4); 4 kilometers west of San Cristobal de las Casas, U.M.M.Z. (2); Volcan Tacana, 8 kilometers north of Union Juarez, K.U. (2, 1 tadpoles). GUATEMALA: Aha Vcrapaz: Finca Chichen, U.M.M.Z. (1 tadpoles); Finca Los Alpes, K.U. (II, 2 skeletons). Baja Verapaz: Cubulco, B.Y.U. (1). Chimaltenango: Tecpan, A.M.N.H. (1 tadpoles). El Quidic: Nebaj, U.M.M.Z. (1 tadpoles). Gi/a(c- mala: 11 kilometers east of San Jose Pinula, U.M.M.Z. (2 tadpoles). Huehuetenango: San Juan Ixcoy, K.U. (1); 3 kilometers east of San Juan I.xcoy, LI.M.M.Z. (1). jalapa: Aserradero San Lorenzo, U.NLM.Z. (3); 8 kilometers east of Mataquescuintla, La Soledad Grande, F.M.N.H. (I). Quetzaltenango: Granja Lorena, K.U. (I, I skeleton), U.M.M.Z. (1, 2 tad- poles); 10.5 kilometers west-southwest of San Martin Sacatepequez, K.U. (1). San Marcos: Rio Achute below Tacana, U.M.M.Z. (1 tadpoles); Tacana, U.M.M.Z. (I); Tejutla, U.M.M.Z. (I). Solold Pana- jachel, M.C.Z. (I); 2 kilometers northwest of Pana- jachel, K.U. (I), U.M.M.Z. (2, I tadpoles); 1.6 kilo- meters southeast of Solola, K.U. (1). Totonicapdn: Momos tcnan go, U.M.M.Z. (2); Totonicapan, U.S.N.M. (1 tadpoles). 1970 DUELLMAN: HYLID FROGS 721 HONDURAS: Yoio: Fortillo Grande, F.M.N.H. (4). EL SALVADOR: Santa Ana: Ceno Metapaii, K.U. (1); Ceno Trifinio, K.U. (1); Hacienda Los Planes. K.U. (1); Hacienda Montecristo, K.U. (7); Miramnndo, F.M.N.H. (3). Plectrohyla hartwegi MEXICO: Chiapas: Bavrejonel, U.M.M.Z. (1); Paraje El Triunfo, K.U. (1). Oaxaca: Ceno Azul, U.LM.N.H. (1). Plectrohyla ixil MEXICO: Chiapas: 3.6 kilometers south of Rayon Mescalapa, K.U. (1 tadpoles); 5.6 kilometers south of Rayon Mescalapa, K.U. (8, 1 skeleton); 6.2 kilometers south of Rayon Mescalapa, K.U. (28, 3 skeletons, 1 tadpoles), M.C.Z. (2); 4 kilometers north- west of Pueblo Nue\o Solistahuacan, L'.M.M.Z. (2); 15 kilometers north of Pueblo Nuevo Soliasthuacan, U.M.M.Z. (13); 18 kilometers north of Pueblo Nuevo Solistahuacan, K.U. (22), U.M.M.Z. (10); 28 kilo- meters north of Pueblo Nuevo Solistahuacan, U.M.xM.Z (1). GUATEMALA: El Quiche: Finca San Francisco, U.NLM.Z. (5. 1 tadpoles); Finca Tesoro, U.M.M.Z. (1 tadpoles ) . Plectrohyla lacertosa MEXICO: Chiapas: Region de Soconusco, U.LM.N.H. (1). Plectrohyla matudai MEXICO: Chiapas: Cerro Ovando, M.C.Z. (1), U.LM.N.H. (1, 1 tadpoles), U.M.M.Z. (9, 1 skeleton, 3 tadpoles), U.S.N.M. (28); Cerro Tres Picos, U.LM.N.H. (I); El Chiciquite, U.LM.N.H. (2); El Feni.x, U.M.M.Z. (3); El Rastrojo, U.LM.N.H. (1); Monte Cristo, U.M.M.Z. (1); Region de Soconusco, C.A.S. (1), M.C.Z. (1); U.LM.N.H. (9); Rodilla, 16 kilometers south of Ciltepec, U.M.M.Z. (3); LInion Juarez, U.LM.N.H. (3). Oaxaca: Cerro Baul, U.LM.N.H. (2); Rio Ostuta, A.M.N.H. (4), K.U. (1); Sierra Madre above Zanatepec, LI.I.M.N.H. (8); 19 kilometers north-northeast of Zanatepec, L.S.U. (1). GLIATEMALA: Chimaltenango: Acatenango, U.S.N.M. (12); Finca Recreo, U.M.M.Z. (1 tad- poles). Guatemala: 11 ki'ometers east of San Jose Pinula, K.U. (2 tadpoles). Huehuetenango: Finca Injerta, U.M.M.Z. (I). San Marcos: El Porvenir, F.NLX.H. (12); Finca La Paz, K.U. (3, 1 skeleton, 2 tadpoles), U.M.M.Z. (6, 2 tadpoles); Volcan Taju- mulco, F.M.N.H. (2 tadpoles). Suchitepequez: Finca El Naranjo, west slope Volcan Santa Clara, U.LM.N.H. (10). Plectrohyla pycnochila MEXICO: Chiapas: 5 kilometers north-northwest of San Cristobal de las Casas, T.C.W.C. (1). Vera- eru~: Coyame, A.M.N.H. (1) [probably erroneous locality). Plectrohyla quecchi GUATEMALA: Alta Verapaz: Finca Chichen, U.M.M.Z. (2, 1 tadpoles); Finca Los Alpes, K.U. (9, 1 skeleton, 3 tadpoles), U.M.M.Z. (4, 1 tadpoles). El Quiche, Finca San Francisco, U.M.M.Z. ( 1 tad- poles ) . Plectrohyla sagorum MEXICO: Chiapas: Cerro Ovando, M.C.Z. (1), U.LM.N.H. (2), U.M.M.Z. (16, 1 skeleton, 2 tad- poles), U.S.N.M. (18); Cerro Paschtal, U.M.M.Z. (1); Chicomuselo, U.M.M.Z. (10); El Chiciquite, U.LM.N.H. (3); Monte Cristo, U.M.M.Z. (1); Paraje El Triunfo, K.U. (1); Region de Soconusco, U.LM.N.H. (5); Volcan Tacana, 8 kilometers north of Union Juarez, K.U. (13). GUATEMALA: Quctzaltenango: Granja Lorena, K.U. (3, 1 tadpoles), U.M.M.Z. (6, 2 tadpoles); 10.4 kilometers west-southwest of San Martin Sacatepe- quez, K.U. (2 tadpoles). San Marcos: Volcan Taja- mulco, F.M.N.H. (5). EL SALVADOR: Chalatcnango: Los Esemiles, M.V.Z. (1). Pseudacris clarkii MEXICO: Tamaulipas: 8 kilometers west of Matamoros, S.U. (2). Ptemohyla dentata MEXICO: AguascalietUes: 15 kilometers east of Agnasca'ientes, K.U. (4, 4 skeletons), U.LM.N.H. (137); 20.6 kilometers east of Aguascalientes, K.U. (1); 26.2 kilometers east of Aguascalientes, U.LM.N.H. (1). JaUsco: 13 kilometers northeast of Lagos de Moreno, U.LM.N.H. ( 1 ). Pternohyla fodiens MEXICO: Colinia: between Buena Vista and Salvador, U.M.M.Z. (1); Colima, M.C.Z. (2); Que- seria, U.M.M.Z. (1). Jalisco: 3-6 kilometers south of Acatlan, LI.M.M.Z. (2); 8 kilometers west-south- west of Acatlan, K.U. (1); 26.4 kilometers northeast of Ameca L'.I.M.N.H. (3); 3 kilometers northeast of Autlan, U.LM.N.H. (4); 4 kilometers west of Ayoel Chica, U.LM.N.H. (1); 51 kilometers northwest of Ayut'a, K.U. (1); Chapala, A.M.N.H. (10, 1 skele- ton); 1.6 kilometers north of Chapala, A.M.N.H. (1); 11.5 kilometers north of Chapala, U.LM.N.H. (5); 16 kilometers north of Chapala, A.M.N.H. (1); 5 kilometers northwest of DegoUado, K.U. (4); 16 kilo- meters northwest of Degollado, K.U. (2); Guadala- jara, K.U. (1); 16 kilometers east of Guadalajara, U.LM.N.H. (9); 28 kilometers south of Guadalajara, r22 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 U.I.M.N.H. (12); 21 kilometers south, 24 kilometers west of Guada'ajara, K.U. (1); 29 kilometers north- west of Guadalajara, U.I.M.N.H. (2); 1.6 kilometers west of Ixtlahuacan, A.M.N.H. (1); 8 kilometers west of I.xt'ahuacan, A.M.N.H. (1); Jamay, A.M.N.H. (69); Magdalena, A.M.N.H. (1), U.I.M.N.H. (11); 3 kilometers east-northeast of Magdalena. K.U. (1); 18 kilometers north of Santa Cruz, K.U. (7); 8 kilo- meters south of Santa Cruz, T.C.W.C. (20); 14 kilo- meters northeast of Tepatitlan, U.I.M.N.H. (22); 6 kilometers .southwest of Tepatitlan, U.I.M.N.H. (3); 16 kilometers northwest of Tequila, T.C.W.C. (2); 19 kilometers northeast of Union Tula, K.U. (1); 8 kilometers southwest of Union Tula, K.U. ( 17, 1 skeleton). Michoacdn: between Rio Marquez and Cuatro Caminos, K.U. (2). Naijarit: Acaponeta, U.S.N.M. (1); 29-50 kilometers south of Acaponeta, A.M.N.H. (10); Ahuacatliin, T.C.W.C. (2); 56 kilo- meters south of Esquinapa (Sinaloa), K.U. (3); Ixtlan del Rio, U.M.M.Z. (1); 1.6 kilometers east of Lxtliin del Rio, K.U. (1); 5 kilometers southeast of Mirador, K.U. (I); Penitas, A.M.N.H. (4); Rio Acaponeta, 4 kilometers south-southwest of Acaponeta, A.M.N.H. (1); Rio San Cayateno, 5.6 kilometers southeast of Tepic. A.M.N.H. (5); 3 kilometers southwest of Rosa- morada, K.U. (3); Tepic, U.I.M.N.H. (2); 37 kilo- meters east of Tepic, M.\'.Z. (1); 8.6 kilometers south-southeast of Tepic, U.M.M.Z. (9); 3 kilometers south of Tepic, S.U. (1); 34 kilometers north-north- west of Tepic, K.U. (I tadpoles); 11 kilometers southeast of Tuxpan, U.I.M.N.H. (27). Sinaloa: Concordia, T.C.W.C. (1); 4 kilometers northea.st of Concordia, K.U. (1); Costa Rica, 25 kilometers .south of Culiacan, U.I.M.N.H. (7); Eldorado, A.M.N.H. ( 1 ); 1.6 ki'ometers northeast of El Fuerte, U.I.M.N.H. (4); 34 kilometers southeast of Esquinapas, K.U. (7); 21 kilometers northeast of Los Mochis, U.I.M.N.H. (5); Matatan, K.U. (4), 7.3 kilometers southwest of Matatan, K.U. (2); Mazatlan, M.C.Z. (6, I skeleton), U.I.M.N.H. (6); 1.6 kilometers north of Mazatlan, K.U. (2 tadpoles); 5.6 kilometers north of Mazatlan, U.M.M.Z. (28); 6-12 kilometers north of Mazatkrn, U.M.M.Z. (I); 14.4 kilometers north of Mazatkrn, U.I.M.N.H. (1); 31.4 kilometers nortli of Mazatlan, U.M.M.Z. (I); Rosario, K.U. (1), U.I.M.N.H. (4), U.S.N.M. (1); 5 kilometers southwest of San Ignacio, K.U. (1); 1.6 east-northeast of San Lorenzo, K.U. (2); Villa Union, K.U. (14); 1 kilometer north of Villa Union, K.U. (6 skeletons); 10 kilometers north- east of Villa Union, K.U. (I); 3.7 kilometers east of Villa Union, K.U. (6). Sonora: 5 kilometers north- northwest of Alamos, K.U. (1); 13 kilometers north- northwest of ,-\lamos, U.I.M.N.H. (1); 13 kilometers north of Ciudad Obregon, K.U. (2); El Bamuri, S.U. (4); 13 kilometers north of El Oasis, U.M. (4); 45 kilometers east of Hermosillo, A.M.N.H. (2); 18.4 kilometers south of Hermosillo A.M. (2); 25 kilome- ters west of La Playa, S.U. (3); Magdalena, U.F. (2); 21 kilometers south of Masiaca, T.C.W.C. (5); 64 kilometers south of Na\ajoa, K.U. (1); 5 kilometers northwest of Navajoa, U.M.M.Z. (7); 8 kilometers north of Noria, S.U. (I), U.M.M.Z. (41); Tricheras, A.M.N.H. (5). Ptychohyla euthysanofa euthysanota MEXICO: Chiapas: Cascarada. 30 kilometers west of Ciltapec, U.M.M.Z. (2); Cerro Ovando, U.M.M.Z. (2); Chicomuselo, U.M.M.Z. (2); Finca Juarez, 28 kilometers north of Escuintla, U.S.N.M. (4); Las Nubes, Cerro Ovando, U.S.N.M. (9); Salto de Agua, U.S.N.M. (13). Oaxaca: Cerro Pecho Blan- co, U.I.M.N.H. (1); between La Gloria and Cerro Azul, U.I.M.N.H. (2); Rio Grande, A.M.N.H. (2); Santo Tonias Tecpan, U.I.M.N.H. (1); Sierra Madre above Zanatepec, U.I.M.N.H. (3). GU.A.TEMALA: San Marcos: Finca La Paz, 2 kilometers west of La Reforma, K.U. ( 14, 1 skeleton, 3 tadpoles), M.C.Z. (1), U.M.M.Z. (I, 7 tadpoles); Finca Pirineos, Rio Samala, F.M.N.H. (I). Santa Rosa: Finca La Gloria, U.M.M.Z. (2 tadpoles). Solo- la: Finca Santo Tomas, U.M.M.Z. (1 tadpoles); Olas de Moca, near Moca, F.M.N.H. ( 1 ). EL S.^LVADOR: Chalatcnango: Los Esemiles, U.S.N.M. (I). Santa Ana: Miramundo, F.M.N.H. (I). Ptychohyla euthysanota macrotympanum ME.XICO: Chiapas: 6 kilometers northeast of Chiapa de Corzo, T.C.W.C. (I); 16 kilometers east of Chiapa de Corzo, T.C.W.C. (1); 16 kilometers east of Chiapa de Corzo, A.M.N.H. (1); Linda Vista, 2 kilometers northwest of Pueblo Nuevo Solistahuaciin, K.U. (2, 1 skeleton); Rio Hondo, 9.5 kilometers south of Pueblo Nue\o Solistahuacan, K.U. (2, 4 tadpoles); 18 kilometers northwest of Pueblo Nuevo Solistahua- can, K.U. (I); San Fernando, M.Z.T.G. (2); Tonina (ruins), K.U. (I). GUATEMALA: Huchiictcnanpo: Finca La De- mocracia, U.M.M.Z. (1, 2 tadpoles); Jacaltenango, U.M.M.Z. (3, 1 tadpoles); 2 kilometers west of San Pedro Necta, U.M.M.Z. (1 tadpoles). Ptychohyla ignicolor MEXICO: Oaxaca: Campamento \'ista Hermosa, K.U. (4, 1 tadpoles), U.M.M.Z. (1); 4.2 kilometers south of Campamento Vista Hermosa, K.U. (34, 1 skeleton, 2 tadpoles, 1 eggs), M.C.Z. (4), U.I.M.N.H. (2); 6 kik)meters south of Campamento Vista Her- mosa, K.U. (8, 1 skeleton), U.M.M.Z. (3); 8 kilome- ters .south of Campamento Vista Hermosa, U.M.M.Z. (7). Ptychohyla Iconhardschultzei MEXICO: Guerrero: Agua del Obispo, F.M.N.H. (4), M.C.Z. (I), U.I.M.N.H. (2), U.S.N.M. (1); Malinaltepec, Z.M.B. (2); 1.6 kilometers southeast of San Andreas de la Cruz, U.M.M.Z. (3). Oaxaca: Campamento Vista Hermosa, K.U. (8, 3 tadpoles), U.M.M.Z. (1); 2.5 kilometers north of La Soledad, K.U. (1); 30 kilometers north of San Gabriel Mixte- pec, K.U. (22); 33 kilometers north of San Gabriel Mixtepec, K.U. (20), 37 kilometers north of San Gabriel Mixtepec, K.U. (7), M.C.Z. (5); San Lucas Camotlan, U.I.M.N.H. (I), U.S.N.M. (2); 10.4 kilo- 1970 DUELLMAN: HYLID FROGS 723 meters south of Valle Nacional, U.M.M.Z. ( 1 tad- poles); 22.7 kilometers south of Valle Nacional, U.M.M.Z. (5); 32.6 kilometers south of Valle Nacion- al, U.M.M.Z. (1); 5 kilometers south of Yetla, K.U. (1 tadpoles); 7.5 kilometers south of Yetla, K.U. (9, 2 skeletons. 4 tadpoles), U.M.M.Z. (2, 1 tadpoles); 9 kilometers .south of Yetla, K.U. ( 1 tadpoles). Plectrohyla schmidtoruni chamulae MEXICO: Chiapas: 32 kilometers north of Jitotol, U.I.M.N.H. (1); 15 kilometers north of Pueblo Nuevo Solistahuacan, U.M.M.Z. (4); 16.4 kilometers north of Pueblo Nuevo Solistahuacan, U.M.M.Z. (10); 18 kilometers north of Pueblo Nuevo Solistahuacan, K.U. (6), U.M.M.Z. (18); 18.6 kilometers north of Pueblo Nuevo Solistahuacan, K.U. (5), U.M.M.Z. (4); 5.6 kilometers south of Rayon Mescalapa, K.U. (1, 1 tad- poles); 6.2 kilometers south of Rayon Mescalapa, K.U. (17, 1 skeleton, 1 tadpoles), M.C.Z. (7); 5.6 kilome- ters south of Solusuchiapa, T.C.W.C. ( 1 ). Plectrohyla schmidtoruni schmidtorum MEXICO; Chiapas: Finca Irlandia, U.M.M.Z. (2); Finca San Jeronimo, U.I.M.N.H. (2). Oaxaca: Sierra Madre above Zanatepec, U.I.M.N.H. (4). GUATEMALA: San Marcos: El Porvenir, F.M.N.H. (3), U.M.M.Z. (1); Finca La Paz, 2 kilo- meters west of La Reforma, K.U. (29, 2 skeletons, 3 tadpoles), M.C.Z. (2), U.M.M.Z. (4 tadpoles). Plectrohyla spinipollex GUATEMALA: Alta Verapaz: Finca Chichen, U.M.M.Z. (1 tadpoles); Finca Los Alpes, K.U. (.30, 3 skeletons, 2 tadpoles), M.C.Z. (2), U.M.M.Z. (4, 1 tadpoles); La Primavera, U.M.M.Z. (1 tadpoles); Panzamala, U.M.M.Z. (1 tadpoles). Baja Verapaz: 32 kilometers north of Moraziin, K.U. (1 tadpoles); Santa Elena, U.M.M.Z. (3). Huehuetenango: 1 kilo- meter east of Barillas, U.M.M.Z. (2 tadpoles). Pro- greso: Finca Bucaral, U.M.M.Z. (3, I skeleton, 1 tad- poles ) . HONDURAS: Atlantidad: mountains behind Ceiba, M.C.Z. ( 1 ). Francisco Morazdn: Cerro Uyuca, A.M.N.H. (3), K.U. (1), U.M.M.Z. (2). Yoro: Por- tillo Grande, M.C.Z. (1). NICARAGUA: Matagalpa: Finca Tepeyac, 10 kilometers north, 9 kilometers east of Matagalpa, K.U. (3 tadpoles); 2.5 kilometers east of Matagalpa, U.M.M.Z. (1); Santa Maria de Ostuma, K.U. (1). Smilisca baudinii MEXICO: No specific locality, M.N.H.N. (1). Campcche: Balchacaj, F.M.N.H. (4), U.I.M.N.H. (15); Champoton, U.M.M.Z. (4); 16 kilometers east of Champoton, U.M.M.Z. (1); 5 kilometers south of Champoton, K.U. (7); 10 kilometers south of Cham- poton, K.U. (4, 2 tadpoles); 24 kilometers south of Champoton, U.M.M.Z. (2); Chuina, K.U. (3); Ciudad del Carmen, U.I.M.N.H. (6); Dzibalchen, K.U. (19); Encarnacion, F.M.N.H. (14), U.I.M.N.H. (16); 1 kilometer west of Escarcega, K.U. (6); 7 kilometers west of Escarcega, K.LI. (22); 14 kilometers west of Escarcega, K.U. (2); 3 kilometers north of Hopelchen, K.U. (3); Matamoras, F.M.N.H. (1); Pital, U.I.M.N.H. (1); 1 kilometer southwest of Puerto Real, Isla del Carmen, K.U. (20); Ruinas Edzna, K.U. (I); San Jose Carpizo, U.M.M.Z. (1); Tres Brazos, F.M.N.H. (1), U.I.M.N.H. (3); Tu.xpena Camp, U.M.M.Z. (1). Chiapas: Acacoyagua, U.S.N.M. (7); 5 kilometers east of Arroyo Minas, U.I.M.N.H. (5); Berriozabal, U.M.M.Z. (7); Chiapa de Corzo, U.M.M.Z. (2); Cintalapa, U.I.M.N.H. (1); Colonia Soconusco, U.S.N.M. (5); 5 kilometers west of Colonia Soconusco, U.M.M.Z. (7); Comitan, K.U. (1); El Suspiro, U.M.M.Z. (11); Escuintla, U.M.M.Z. (10); 6 kilometers northeast of Escuintla, U.M.M.Z. (26); 3 kilometers east of Finca Juarez, U.I.M.N.H. (1); Finca P-russia, U.M.M.Z. (I); Honduras, U.M.M.Z. (4); La Grada, U.M.M.Z. (I); 21 kilometers south of La Trinitaria, U.I.M.N.H. (2); 14.4 kilometers south- west of Las Cruces, K.U. (6), Palenque, U.I.M.N.H. (1), U.S.N.M. (12); 2 kilometers northwest of Pueblo Nuevo Solistahuacan, K.U. (3), U.M.M.Z. (9); 1.3 kilometers north of Puerto Madero, K.U. (4); 4 kilo- meters north of Puerto Madero, K.U. (2); 8 kilometers north of Puerto Madero, U.M.M.Z. (2); 12 kilometers north of Puerto Madero, K.U. (1); 17.6 kilometers north of Puerto Madero, U.M.M.Z. (1); Rancho Mon- serrata, U.I.M.N.H. (2), U.M.M.Z. (2); Region Soco- nusco, U.I.M.N.H. (15); San Bartola, U.I.M.N.H. (12); San Geronimo, U.I.M.N.H. (1); San Juanito, U.S.N.M. (2); San Ricardo, F.M.N.H. (1); Solosu- chiapa, K.U. (2); Tapachula, F.M.N.H. (4), U.I.M.N.H. (3); Tonala, A.M.N.H. (1); F.M.N.H. (2), U.I.M.N.H. (5), U.S.N.M. (1); Tuina, K.U. (1 skeleton); Tuxtia Gutierrez, F.M.N.H. (2); 6 kilo- meters east of Tu.xtla Gutierrez, U.I.M.N.H. (1); 10 kilometers east of Tu.xtla, Gutierrez, U.M.M.Z. (1). Chihuahua: 2.4 kilometers southwest of Toquina, K.U. (2); Riito, K.U. (1). Coahuila: mountain near Saltillo, U.I.M.N.H. (2). Colima: No specific locality, F.M.N.H. (1); Colima, A.M.N.H. (2); Hacienda Albarradito, U.M.M.Z. (2); Hacienda del Colomo, A.M.N.H. (1); Los Mezcales, U.M.M.Z. (1); Man- zanillo, A.M.N.H. (2); Paso del Rio, F.M.N.H. (3), U.I.M.N.H. (3), U.M.M.Z. (3); Periquillo, U.M.M.Z. (17); 1.6 kilometers southwest of Pueblo Juarez, U.M.M.Z, (1); Quesaria, F.M.N.H. (4), U.I.M.N.H. (3); U.M.M.Z. (14); Santiago, U.M.M.Z. (1); 7.2 kilometers southwest of Tecolapa, U.M.M.Z. (1). Guerrero: Acahuitzotia, U.F. (4); U.M.M.Z. (3); 3 kilometers south of Acahuitzotia, K.U. (5); Acapulco, A.M.N.H. (1), U.M.M.Z. (4), U.S.N.M. (1); 3 kilo- meters north of Acapulco, U.M.M.Z. (1); 8 kilome- ters northwest of Acapulco, U.F. (7); 27 kilometers northeast of Acapulco, U.I.M.N.H. (14); Agua del Obispo, F.M.N.H. (5), K.U. (4), U.I.M.N.H. (3); Atovac, K.U. (4); Buena Vista, F.M.N.H. (4), U.I.M.N.H. (1); Caculutla, K.U. (1); 20 kilometers south of Chilpancingo, F.M.N.H. (5); Colonia Buenas Aires, U.M.M.Z. (1); El Limoncito, F.M.N.H, (4); El Treinte, F.M.N.H. (5), U.I.M.N.H. (3), U.S.N.M. 724 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 (3); Laguna Coyuca, U.M.M.Z. (2); 3 kilometers north of Mazatlan, U.I.M.N.H. (3); 9 kilometers south of Mazatlan, F.M.N. H. (4), U.I.M.N.H. (2); Mex- cala, F.M.N.H. (5), U.I.M.N.H. (2); Ocotito, K.U. (10); 5.4 kilometers north of Ocotito, U.M.M.Z. (4); 1,6 kilometers north of Organos, U.I.M.N.H. (12); Palo Blanco, F.M.N.H. (5), U.I.M.N.H. (4); Pie de la Cuesta, A.M.N.H. (5); Puerto Marquez, A.M.N. H. (13); 5.6 kilometers south of San Andreas de la Cruz, K.U. (2); San Vincents, K.U. (1); Zacualpan, U.M.M.Z. (1). Hidalgo: below Tianguistengo, F.M.N.H. (1). Jalisco: Atenqueque, K.U. (2); 5 kilometers northeast of Autlan, U.I.M.N.H. (1); 5 kilometers east of Barro de Navidad, U.M.M.Z. (1); Charco Hondo, U.M.M.Z. (1); 6.4 kilometers east- northeast of La Huerta, K.U. (2); between La Huerta and Tecomates, K.U. (1); 3 kilometers southeast of La Resolana, K.U. ( 1, 1 skeleton). 11 kilometers south, 1.6 kilometers east of Yahualica, K.U. (1); Zapotilitic, F.M.N.H. (1). Michoacdn: Aguililla, U.M.M.Z. (5); Apatzingan, F.M.N.H. (25); K.U. (1 skeleton); 7 kilometers east of ,\patzingan, U.M.M.Z. (1); 11 kilo- meters east of Apatzingan, U.M.M.Z. (3); 27 kilome- ters south of Apatzingan, K.U. (3); 1.6 kilometers north of Arteaga, U.M.M.Z. (1); Charapendo, U.M.M.Z. (1); Coahuayana, U.M.M.Z. (1); El Sabino, F.M.N.H. (7), U.I.M.N.H. (2); La Placita, U.M.M.Z. (1); La Playa, (1), 30 kilometers east of Nueva Italia, U.M.M.Z. (2); 4 kilometers south of Nueva Italia, U.M.M.Z. (1); Ostula, U.M.M.Z. (4); Salitre de Estopilas, U.M.M.Z. (1); San Jose de la Montana, U.M.M.Z. (2); 11 kilometers south of Tum- biscatio, K.U. (1); 12 kilometers south of Tzitzio, U.M.M.Z. (1). Morelos: 3.5 kilometers west of Cuautli,\co, K.U. (3); Ocotlan del Rio, I.P.N. (1), Puente de I.xtla, I.P.N. (1); 1 kilometer northeast of Puente de I.xtla, K.U. (2); 20 kilometers south of Puente de Ixtla, F.M.N.H. (1), U.I.M.N.H. (1); Tequesquitengo, A.M.N.H. (4). Nayarit: 3 kilome- ters south of Acaponeta, U.M.M.Z. (4); 56 kilometers south of Esquinapa (Sinaloa), K.U. (1); Jesus Maria, A.M.N.H. (1); San Bias, K.U. (5), U.S.N. M. (1); 8.6 kilometers east of San Bias, U.M.M.Z, (1); Tepic, U.I.M.N.H. (4); 4 kilometers east of Tuxpan, K.U. (1); 11 kilometers southeast of Tuxpan, U.I.M.N.H. (28). Nuevo Leon: Galeana, F.M.N.H. (1); Salto Cola de Caballo, F.M.N.H. (63). Oaxaca: 11 kilo- meters .south of Candelaria, U.I.M.N.H. (4); Cerro San Pedro, 24 kilometers southwest of Tehuantepec, U.M.M.Z. (1); Chachalapa, K.U. (1); 8 kilometers south of Chiltepec, K.U. (1); 12 kilometers south of Chivela, U.M.M.Z. (1); 12 kilometers south of Chi- vela, U.M.M.Z. (1); Cocahuatepec, U.I.M.N.H. (1); Coyul, U.S.N.M. (1); 50 kilometers east-.southeast of Cuajinicuilapa, U.I.M.N.H. (1); Escurano, U.LM.N.H. (1); Garza Mora, U.I.M.N.H. (2); Jucha- tengo, K.U. ( 1 ); 17 kilometers northeast of Juchaten- go, K.U. (2 tadpoles); Juchitan, U.S.N.M. (1); La- gartero, U.I.M.N.H. (1); Matias Romero, U.I.M.N.H. (1); Mirador, AMN.H. (23); Mira Leon, 1.6 kilo- meters north of Huatulco, U.I.M.N.H. (2); Mixte- quilla, A.M.N.H. (1); Pochutla, K.U. (15), U.I.M.N.H. (9); 17.6 kilometers west-northwest of Puerto Escondido, U.MMZ, (1); Quiengola, A.M.N.H. (2); Rio del Corte, U.I.M.N.H. (1); Rio Mono Blanco, U.I.M.N.H. (1); Rio Sarabia, 5 kilo- meters north of Sarabia, U.M.M.Z. (4); 2.5 kilometers north of Salina Cruz, K.U. (2); San Antonio, U.I.M.N.H. (1); 5 kilometers north-northwest of San Gabriel Mixtepec, K.U. (1); San Pedro del Istmo, U.I.M.N.H. (1); Santo Domingo, U.S.N.M. (3); 3.7 kilometers north of Sarabia, U.M.M.Z. (3); Tapana- tepec, K.U. (1, 1 skeleton), U.I.M.N.H. (1), U.M.M.Z. (1); between Tapanatepec and Zanatepec, U.I.M,N,H, (2); Tecuane, U.M.M.Z. (3); Tehuante- pec, A.M.N.H. (3), U.M.M.Z. (38), U.S.N.M. (6); 4.5 kilometers west of Tehuantepec, K.U. ( 14 skele- tons); 10 kilometers south of Tehuantepec, K.LT. (2); Tehuantepec, K.U. (2); Temazcal, U.S.C. (3); 3 kilometers south of Tolocita, K.U. (4); Tolosa, A.M.N.H, (1); Tuxtepec, UM.MZ. (2), 13 kilome- ters south of Tuxtepec, U.I.M.N.H. ( 1 ); 27 kilometers south of Tuxtepec, U.I.M.N.H. (24); 2 kilometers south of Valle Nacional, K.U. (2); 11 kilometers north of Vista Hermosa, K.U. (1, 6 tadpoles); Yetla, K.U. (1); above Zanatepec, U.I.M.N.H. (1). Piiebla: 16 kilometers southwest of Mecatepec (Veracruz), U.I.M.N.H, (2); San Diego, A.M.N.H. (I), U.S.N.M. (1); Vegas de Suchil, A.M.N.H. (1); Villa Juarez, U.F. (1). Quintana Roo: Coba, F.M.N.H. (1); Es- meralda, U,M,M.Z, (1); 4 kilometers north-northeast of Felipe Carillo Puerto, K.U. (2); Pueblo Nuevo X- Can, K.U. (1); 4 kilometers west-southwest of Puerto Juarez, K.U. (5, 1 tadpoles); 12 kilometers west of Puerto Juarez, K.U. (5); San Miguel, Isla de Cozumel, U.M.M.Z, (18); 3.5 kilometers north of San Miguel, Isla de Cozumel, K.U. (4); 10 kilometers east of San Miguel, Isla de Cozumel, U.M,M,Z. (1); Telantrmich, F,M,N,H, (1). San Luis Potosi: Ciudad Valles, A.M.N.H. (12), F,M,N,H, (2), K.U. (1); 21 kilome- ters north of Ciudad Valles, U.M.M.Z. (1); 6 kilo- meters east of Ciudad Valles, U.F. (1); 24 kilometers east of Ciudad Valles, U.F. (5); 5 kilometers south of Ciudad Valle.s, U.I. M.N, H. (1); 16 kilometers south of Ciudad Valles, A,M,N,H. (1); 30 kilometers south of Ciudad Valles, F.M.N, H, (3), U,I,M.N.H. (2); 63 kilometers south of Ciudad Valles, U.I.M.N.H. (2); Pujal, U.M.M.Z. (2); Rio Axtla, near A.xtla, A.M.N.H. (6), K.U. (1); Tamazunchale, A.M.N.H. (1), F,M,N,H. (4), U.F. (2), U.I,M,N.H. (1), U,M,M,Z. (11), U.S.N.H. (1); 17 kilometers north of Tama- zunchale, U.I.M.N.H, (1); 2.4 kilometers south of Tamazunchale, A.M.N.H. (1); 17 kilometers east of Tamuin, U.F. (2); Xilitla, U.I.M.N.H. (2), Sinaloa: 8 kilometers north of Carrizalejo, K.U. (1); 4 kilome- ters northeast of Concordia, K.U. (1); 5 kilometers southwest of Concordia, K.U, (2); 6 kilometers east of Cosala, K.U. (1); Costa Rica, 16 kilometers .south of Culiacan, U.I.M.N.H. (3); 51 kilometers south- southeast of Culiacan, K.U. ( 1 ); El Dorado, K.U. ( 1 ); 1.6 kilometers northeast of El Fuerte, F.M.N.H. (1); Lsla Palmito del \'erde, middle, K.U. (2); 21 kilo- meters north-northeast of Los Mochis, U.I.M.N.H. (2); Matatan, K.U. (1); 7.3 kilometers southwest of Matatiin, K.U. (6); Mazatlan, A.M.N.H. (1), U.M.M.Z. (3); 57 kilometers north of Mazatlan, 1970 DUELLMAN: HYLID FROGS 725 U.I.M.N.H. (1); Plomosas, U.S.N.M. (2); Presi- dio, U.I.M.N.H. (1), U.S.N.M. (1); Rosario, K.U. (2), U.I.M.N.H. (1); 5 kilometers east of Ro.sario, U.I.M.N.H. (17); 8 kilometers east of Rosario, U.I.M.N.H. (17); 8 kilometers south-south- east of Rosario, K.U. (1); 5 kilometers southwest of San Ignacio, K.U. (I); 1.6 kilometers east-northeast of San Lorenzo, K.U. (8); Teacapan, Isla Palmito del \'erde, K.U. (I); 9.6 kilometers north-northwest of Teacapan, K.U. (1); Villa Union, K.U. (I); 9 kilo- meters northeast of Villa Union, K.U. (4); 1 kilometer west of Villa Union, A.M.N.H. (I). Sonora: Guira- coba, A.M.N.H. (25). Tabasco: 4 kilometers north- east of Colmalcaico, A.M.N.H. (1); 10 kilometers south of Huimanguillo, U.M.M.Z. (1); Teapa, U.M.M.Z. ( 1 ); 5 kilometers north of Teapa, U.M.M.Z. (4); 10 kilometers north of Teapa, U.M.M.Z. (3); 13 kilometers north of Teapa, U.M.M.Z. (7); 21 kilome- ters north of Teapa, U.M.M.Z. (2); 29 kilometers north of Teapa, U.M.M.Z. (11); Tenosique, U.S.N.M. (3). Tamaulipas: Acuna, U.M.M.Z. (1); 5 kilome- ters south of Acuna, U.M.M.Z. (1); 13 kilometers north of Antiguo Morelos, U.I.M.N.H. (4); 3 kilome- ters south of Antiguo Morelos, U.F. (1); 3 kilometers northeast of Chamal, U.M.M.Z. (1); 1.6 kilometers east of Chamal, U.M.M.Z. (1); Ciudad Mante, U.M.M.Z, (7); 16 kilometers north of Ciudad Vic- toria, F.M.N.H. (1); 34 kilometers north of Ciudad Victoria, K.U. (17); 8.8 kilometers south of Ciudad Victoria, U.I.M.N.H. (3); 11 kilometers west of Ciu- dad Victoria, U.I.M.N.H. (1); 16 kilometers west of Ciudad Victoria, U.I.M.N.H. (1); 3 kilometers west of El Carizo, U.M.M.Z. ( 1 ); Gomez Farias, U.M.M.Z. (2); 8 kilometers northeast of Gomez Farias, U.M.M.Z. (18); 8 kilometers northwest of Gomez Farias, U.M.M.Z. (2); 16 kilometers west of Gon- zales, K.U. (2); Jimenez, K.U. (1); 1.2 kilometers south of La Castilla, U.I.M.N.H. (7); 5 kilometers south of La Castilla, U.I.M.N.H. (23); La Clemen- tina, 6 kilometers west of Forlon, U.S.N.M. (1); Limon, U.I.M.N.H. (1); Llera, U.S.N.M. (4); 3 kilo- meters east of Llera, U.I.M.N.H. (1); 21 kilometers south of Llera, U.I.M.N.H. (2); 23 kilometers south of Llera, U.I.M.N.H. (1); 11 kilometers southwest of Ocampo, U.M.M.Z. (1); 22 kilometers west, 5 kilo- meters south of Piedra, K.U. (4); Rio Sabinas, U.M.M.Z. (1); 5 kilometers west of San Gerardo, U.M.M.Z. (2); Santa Barbara, U.M.M.Z. (2); Villa- gran, F.M.N.H. (4), U.I.M.N.H. (2); 1.7 kilometers west of Xicotencatl, U.M.M.Z. (1). Veracruz: 1.6 kilometers northwest of Acayucan, U.M.M.Z. (1); 28.5 kilometers southeast of Alvarado, U.M.M.Z. (1); 2.4 kilometers south-southwest of Amatitlan, U.M.M.Z. ( 1 ); Azveta, I.P.N. ( 1 ); Barranca Metlac, U.I.M.N.H. (1); Boca del Rio, U.I.M.N.H. (12), U.M.M.Z. (9); 16 kilometers south of Boca del Rio, U.I.M.N.H. (1); between Boca del Rio and Anton Lizardo, U.I.M.N.H. (1); Canada, F.M.N.H. (1); Catemaco, U.M.M.Z. (4); Cerro Chicahuastle, U.I.M.N.H. (1); Ciudad Aleman, U.M.M.Z. (3); Cordoba, F.M.N.H. (3), U.S.N.M. (4); 5.2 kilometers east-southeast of Cor- doba, U.M.M.Z. (4); Cosamaloapan, U.M.M.Z. (2); Coyame, U.I.M.N.H. (5), U.M.M.Z. (5); 1 kilometer southeast of Coyame, U.M.M.Z. (3); Cuatotolapam, U.M.M.Z. (15); Cuautlapan, F.M.N.H. (13), K.U. (22, 9 skeletons), U.I.M.N.H. (78), U.M.M.Z. (38), U.S.N.M. (25); Dos Rios, F.M.N.H. (1); 5 kilome- ters east-northeast of El Jobo, K.U. (3); 6.2 kilome- ters east of Encero, U.I.M.N.H. (I); Escamillo, F.M.N.H. (1), U.I.M.N.H. (I); 1 kilometer north of Fortin de las Flores, U.F. (1); 1 kilometer southwest of Huatusco, L'.M.M.Z. (1); 4 kilometers southwest of Huatusco, U.M.M.Z. (1); 10 kilometers southeast of Hueyapan, U.M.M.Z. (1); 20 kilometers south of Jesus Carranza, K.U. (3); 38 kilometers southeast of Jesus Carranza, K.U. (I); Laguna Catemaco, U.M.M.Z. (62); 1.6 kilometers north of La Laja, U.I.M.N.H. (1); La Oaxaqueiia, A.M.N.H. (2); 16 kilometers west-southwest of Las Conejos, K.U. (4); 17 kilometers east of Martinez de la Torre, U.I.M.N.H. (3); 6.2 kilometers west of Martinez de la Torre, U.I.M.N.H. (3); 2 kilometers east- northeast of Mata Oscura, K.U. (7); Minatitlan, A.M.N.H. (2); Mirador, U.S.N.M. 6 kilometers south of Monte Blanco, U.F. (4); 21 kilometers east of Nanchital, U.M.M.Z. (1); 2 kilometers south of Naranja, U.M.M.Z. (3); 1.6 kilometers northeast of Novillero, U.M.M.Z. (2); 3 kilometers northeast of Novillero, U.M.M.Z. (1); 5.2 kilometers northeast of Novillero, U.M.M.Z. (4); 6 kilometers northeast of Novillero, L'.M.M.Z. (1); 5 kilometers north of Nueva Colonia, U.M.M.Z. (1); Orizaba, U.S.N.M, (2); 4 kilometers northeast of Orizaba, U.M.M.Z. (2); Otatilan, U.I.M.N.H. (17); 10 kilo- meters west-southwest of Pachuquillo, K.U. (5); Panuco, U,M,M.Z. (1); Paraje Nuevo, U.M.M.Z. (73); Paso del Macho, U.I.M.N.H. (1); Paso de Talayo, Jicaltepec, U.S.N.M. (2); Perez, F.M.N.H. ( 5 ) ; 20 kilometers north of Piedras Negras, Rio Blanco, K.U. (1); Plan del Rio, K.U. (6), U.M.M.Z. (6), Potrero, U.I.M.N.H. (4), U.M.M.Z. (4), U.S.N.M. (5); Potrero Viejo, F.M.N.H. (1), K.U. (40, 4 skeletons), U.I.M.N.H. (1), U.M.M.Z. (27), U.S.N.M. (10), 5 kilometers south of Potrero Viejo, K.U. (3); Puente Nacional, U.I.M.N.H. (6); 3 kilo- meters north of Rinconada, U.M.M.Z. (5); Rio de las Playas, U,S,N,M, (2); Rio Seco, U,M.M.Z. (9); Rod- riguez Clara, F.M.N.H. (1); San Andres Tuxtla, F.M.N.H. (4), U.I,M,N,H. (3); 5 kilometers north of San Andres Tuxtla, U.I.M.N.H. (1); 10 kilometers northwest of San Andres Tuxtla, U.M.M.Z. (1); 13.4 kilometers northwest of San Andres Tu.xtla, U,M.M.Z. (2); 19.8 kilometers northwest of San Andres Tuxtla, U.M.M.Z. (1); 27.2 kilometers northwest of San Andres Tuxtla, U.M.M.Z. (I); 4 kilometers west of San Andres Tuxtla, U.M.M.Z. (1); 37.4 kilometers south of San Andres Tu.xtla, U.M.M.Z. (12); 15 kilo- meters east-southeast of San Juan de la Punta, K.U. (1); San Lorenzo, U.S.N.M. (5); 3 kilometers south- west of San Marcias, K.U. (1); 1.5 kilometers south of Santa Rosa, U.I.M.N.H. (1); 2 kilometers south of Santiago Tuxtla U.M.M.Z. (4); Suazel, U.M.M.Z. (1); 14 kilometers east of Suchil, U.I.M.N.H. (1); 15 kilo- meters south of Tampico (Tamaulipas), U.M.M.Z. (4); 4 kilometers north of Tapalapan, U.M.M.Z. (2); Tecolutia, U.I.M.N.H. (24); 16 kilometers northwest 726 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 of Tehuatlan, U.I.M.N.H. (4); 5 kilometers south of Tehuatlan, K.U. (1); Teoce'o, K.U. (1); Tierra Colo- rada, F.M.N.H. (3), U.I.M.N.H. (3); Veracruz, .\.M.N.H. (9), I.P.N. (1), U.I.M.N.H. (1), U.M.M.Z. (3); 24 kilometers west of Veracruz, F.M.N.H. (3). Yucatan: No .specific locality, F.M.N.H. (2), U.S.N.M. (1); Chichen-Itza, F.M.N.H. (17), U.I.M.N.H. (5), U.M.M.Z. (73), U.S.N.M. (I); 9 kilometers east of Cliichen-Itza, K.U. (2); 12 kilome- ters east of Chichen-Itza, K.U. ( 1 ); Merida, F.M.N.H. (8), U.I.M.N.H. (2), U.M.M.Z. (1); 6 kilometers south of Merida, K.U. (1); 8.8 kilometers southeast of Ticul, U.M.M.Z. (1); Valladolid, F.M.N.H. (3); Xcalah-op, F.M.N.H. (9); 3.5 kilometers east of Yokdzonot, K.U. (3, 1 tadpoles). BRITISH HO\DUR.\S: Belize: Belize, F.M.N.H. (4), U.M.M.Z. (1), U.S.N.M. (I); Manatee, F.M.N.H. (4). Cayo: Cayo, U.M.M.Z. (1); 6 kilo- meters south of Cayo, M.C.Z. (1); 2.5 kilometers southwest of Cayo, U.M.M.Z. (1); Cocquericot, U.M.M.Z. (2); Cohune Ridge, U.M.M.Z. (15); Doub'e Falls, F.M.N.H. (1); Mountain Pine Ridge, M.C.Z. (2); Pine Ridge Road, U.M.M.Z. (18); San .\ugustin, U.M.M.Z. (1); Valentin, U.M.M.Z. (8). Orange Walk: Gallon Jug, M.C.Z. (8). S/aiin Creek: Bokowina, F.M.N.H. (2); Hummingbird Highway between Roaring Creek and Stann Creek, U.M.M.Z. (1); 16 kilometers from Hummingbird Highway on road to Monkey River, U.M.M.Z. (1); 5 kilometers south of Waha Loaf Creek, M.C.Z. (1). Toledo: San Pedro Colombia, M.C.Z. (3). GUATEMAL.A.: Alia Verapaz: 5.1 kilometers northeast of Campur, K.U. (2 tadpoles); 28.3 kilo- meters northeast of Campur, K.U. (20, 2 skeletons); Chama, M.C Z. (2), U.M.M.Z. (56); Chinaja, K.U. (8, 1 eggs, 3 tadpoles); Coban, F.M.N.H. (1); Cubil- quitz, U.M.M.Z. (10); Finca Canihor, U.M.M.Z. (1); Finca Chicoyou, K.U. (4, 3 tadpoles); Finca Los .\'pes, K.U. (5, 1 tadpoles); Finca Los Pinales, U.M.M.Z. (2); Finca Tinajas, B.Y.U. (1); Finca Vol- can, U.M.M.Z. (6); Panzos, M.N. H.N. (I), U.M.M.Z. (1); Samac, U.M.M.Z. (1); Samanzana, U.M.M.Z. (6). Baia Verapaz: Chejel, U.M.M.Z. (10); San Geronimo, U.M.M.Z. (16). Chiquiinida: 1.6 kilo- meters southeast of Chiquimula, U.M.M.Z. (1); Es- quipulas, U.M.M.Z. (28). El Peten: Asseradero Machaquila, U.M.M.Z. (1); 20 kilometers north- northwest of Chinaja (Alta Verapaz), K.U. (42); Flores, U.M.M.Z. (1); La Libertad, K.U. (1 tad- poles); U.M.M.Z. (57); 3 kilometers southeast of La Libertad, K.U. (2); 13 kilometers south of La Liber- tad, M.C.Z. (2); Pacomon, U.S.N.M. (1); Piedras Negras, U.S.N.M. (3); Popti.m, U.M.M.Z. (3); Pozo de la Jicotea, U.S.N.M. (1); Ramate-Ya.\ha trail, U.M.M.Z. (1); Rio de la Pasion between Sayaxche and Subin, K.U. (1); Rio San Roman, 16 kilometers north-northwest of Chinaja (Alta Verapaz), K.U. (5); Sacluc, U.S.N.NL (1); Sayaxche, K.U. (2); Tikal, U.M.M.Z. (22); Toocog, K.U. (2, 2 tadpoles); Uaxac- tiin, U.M.M.Z. (3); Yaxha. U.M.M.Z. (1); 19 kilome- ters east of Yaxha, U.M.M.Z. (4). El Quiche: Finca Tesoro, U.M.M.Z. (3, I tadpoles). Escuintla: Rio Guacalate, Ma.sagua, L'.S.N.NL (1); Tiquisate, U.M.M.Z. (7). Guatemala: 16 kilometers northeast of Guatemala, K.U. (9). lluehuetenango: Barillas, U.M.M.Z. (2); Cuilco, U.NLM.Z. (12); Finca San Rafael, 16 kilometers southeast of Barillas, F.M.N.H. (5); 45 kilometers west-northwest of Huehuetenango, K.U. (2); Jacaltenango, U.M.M.Z. (33); La Demo- cracia, U.M.M.Z. (8). Izabal: 2 kilometers southwest of Puerto Matias de Galvez, K.U. (2 tadpoles); Quiri- gua, F.NLN.H. (1), U.M.M.Z. (I). Jalapa: Jalapa, U.M.M.Z. (12). Juiiapa: Finca La Trinidad, U.M.M.Z. (10); Jutiapa, U.M.M.Z. (I); 1.6 kilome- ters southeast of Mongoy, K.U. ( 1 ) ; Santa Catarina Mita, U.M.M.Z. (1). Progreso: Finca Los Leones, U.M.M.Z. (1). Qiictzaltcnango: Coatepeque, A.M.N.H. (1). Retalhucleu: Casa Blanca, U.M.M.Z. (18); Champerico, U.M..M.Z. (3). San Marcos: Puente Talisman, U.S.N.M. (2). Santa Rosa: Finca La Guardiana, U.M.M.Z. (6); Finca La Gloria, U.M.M.Z. (6); 1.6 kilometers west-soutliwest of El Molino, K.U. (4). Suchitepequez: Mazatenango, U.I.M.N.H. (1). EL SALVADOR: La Libertad: 16 kilometers northwest of Santa Tecla, K.U. (3). Morazdn: Divisadero, Lf.S.N.M. (I). San Salvador: San Sal- vador, F.M.N.H. (13), K.U. (34, 2 skeletons, 1 eggs, 7 tadpoles), U.M.M.Z. (6), U.S.N.M. (1). HONDURAS: Atlantidad: Isla de Roatan, F.M.N.H. (4); La Ceiba, U.S.N.\L (4); Lancetilla, M.C.Z. (5); Tela, M.C.Z. (3), U.M.M.Z. (1), U.S.N.M. (2). Choluleca: 1.5 kilometers northwest of Choluteca, K.U. (5); 10 kilometers northwest of Choluteca, K.U. (1); 10 kilometers east of Choluteca, K.U. (2); 12 kilometers east of Choluteca, K.U. (1); 5 kilometers south of Choluteca, U.S.C. (2). Colon: Ba'fate, A.M.N.H. (4); Bambii, U.F. (1); Patuca, U.S.N.M. (I); Rio Segovia, M.C.Z. (I). Comatjagua: La Mision, 3.5 leagues north of Siguatepeque, M.C.Z. (2). CopdtJ: Copan, U.M.M.Z. (2). Cortes: Cofra- dia, A.M.N.H. (2); Hacienda Santa Ana, F.M.N.H. (8); Lago de Yojoa, M.C.Z. (2); Rio Lindo, A.M.N.H. (1); San Pedro Su'a, K.U. (1). El Paraiso: El Vol- can, M.C.Z. (1). Francisco Morazdn: Guaimaca, U.M.M.Z. (1); Tegucigalpa, B.Y.U. (9), M.C.Z. (3), U.S.N.M. (1). Santa Barbara: Santa Barbara, U.S.N.M. (4) . NICARAGUA: Carazo: 3 kilometers north, 4 kilo- meters west of Ciriamba, K.U. (5). Chinandega: 4 kilometers north, 2 kilometers west of Chichigalpa, K.U. (1); Chinandega, M.C.Z. (1); Hacienda Bella- vista, Volcan Casita, K.U. (4); Rio Tama, U.S.N.M. (1); San .Antonio, K.U. (20, 6 skeletons). Chontales: I kilometer northeast of ."Kcoyapa, K.U. (1); 1 kilo- meter north, 2.5 kilometers west of \'illa Somoza, K.U. (1). Esteli: Finca Daraili, 5 kilometers north, 15 kilometers east of Condega, K.U. (5); Finca \'enecia, 7 kilometers north, 16 kilometers east of Condega, K.U. (1). Leon: 1.6 kilometers east-northeast of Poneloya, K.U. (2). Managua: Managua, U.S.N.NL (2); 8 kilometers northwest of Managua, K.U. (17); 20 kilometers northeast of Managua, K.U. (3); 5 kilometers southwest of Managua, K.U. (8); 1-3 kilo- 1970 DUELLMAN: HYLID FROGS 727 meters north of Sabana Grande, K.U. (14); 20 kilo- meters sonth of Tipitapa, K.U. (1). Matapalpa: Guasqnalie, U.M.M.Z. (1); Matagalpa, U.M.M.Z, (1); 19 kilometers north of Matagalpa, U.M.M.Z. (1); Sebaco, K.U. (1). Nueva Segovia: 1.5 kilometers north, 1 kilometer east of Jalapa, K.U. (8); 5 kilome- ters north, 2.5 kilometers east of Jalapa, K.U. (6). Rio San Juau: Greytown, U.S.N.M. (4). Rivas: Javillo, U.M.M.Z. (1); Moyogalpa, Isla Ometepe, K.U. (10, 1 tadpoles); Pefias Blancas, K.U. (1); Rio Jaxillo, 3 kilometers north, 4 kilometers west of Sapoa, K.l'. (3, 1 skeleton); 13.1 kilometers southeast of Rivas, K.U. (1); 14.8 kilometers southeast of Rivas, K.U. (3); 11 kilometers south, 3 kilometers east of Rivas, K.U. (1); 16 kilometers south of Rivas, M.C.Z. (2); 7.7 kilometers northeast of San Juan del Sur, K.U. (2); 16.5 kilometers northeast of San Juan del Sur, K.U. (2, 1 tadpoles); 5 kilometers southeast of San Pablo, K.U. (5). Zelaya: Bonanza, K.U. (2); Cooley, .^.M.N.H. (11); Cukra, A.M.N. H. (2); El Recreo K.U. (10); Masahuas, Rio Huaspuc, A.M.N.H. (4); 11 kilometers northwest of Rama, Rio Siquia, U.M.M.Z. (8); Rio Escondido, U.S.N.M. (2); Rio Siquia at Rio Mico, U.M.M.Z. (10); Sioux Plantation, A.M.N.R (15). COSTA RICA: Alajuda: Los Chiles, A.M.N.H. (1); Orotina, xM.C.Z. (2); San Carlos, U.S.N.M. (1). Guanacastc: Finca Taboga, K.U. (5); La Cruz, U.S.Z. (3); 4.3 kilometers northeast of La Cruz, U.S.N.M. (1); 18.4 kilometers south of La Cruz, U.S.C. (1); 23.5 kilometers south of La Cruz, U.S.C. (4); 3 kilo- meters west of La Cruz, U.S.C. (4); 2 kilometers northeast of Las Caiias, K.U. (3); Las Huecas, U.M.M.Z. (2); Liberia, K.U. (1), U.S.C. (1); 11.5 kilometers north of Liberia, U.S.C. (1); 13 kilometers north of Liberia, U.S.C. (1); 22.4 kilometers north of Liberia, U.S.C. (1); 8 kilometers north-northwest of Liberia, K.U. (1); Penas Blancas, K.U. (2); 8.6 kilometers east-southeast of Playa del Coco, U.S.C. ( 1 ) ; 2.8 kilometers east-southeast of Playa del Coco, U.S.C. (1); Rio Piedra, 1.6 kilometers west of Bagaces, U.S.C. (1); Rio Bebedero, 5 kilometers south of Bebedero, K.U. (1); 5 kilometers northeast of Tilaran, K.U. (5). Heredia: Puerto Viejo, K.U. (4); 13 kilometers southwest of Puerto Viejo, K.U. (5). Limon: Batan, K.U. ( 1 ); Guacimo, U.S.C. (1); Los Diamantes, K.U. (1); Pandora, U.S.C. (3); Suretka, K.U. (2); Tortugero, U.K. (4). Puntarenas: Barranca, F.M.N.H. (3); 15 kilometers west-north- west of Barranca, K.U. (3), U.M.M.Z. (1); 18 kilo- meters west-northwest of Barranca, U.M.M.Z. (4); 4 kilometers west-northwest of Esparta, K.U. (38, 5 skeletons); 19 kilometers northwest of Esparta, K.U. (8). Smilisca cyanosticta MEXICO: Chiapas: Monte Libano, M.C.Z. (9); 8 kilometers north of San Fernando, 24 kilometers east of Tuxtla Gutierrez, U.I.M.N.H. (1). Oaxaca: 11 kilometers north of Campamento Vista Hennosa, K.U. (15, 3 skeletons, 1 eggs, 6 tadpoles), U.I.M.N.H. (3), 8 kilometers south of Yetla, K.U. (1); U.M.M.Z. (8). Veracruz: Coyame, U.M.M.Z. (2); Ijetween Coyame and Tebanco, U.M.M.Z. (1); Dos Amates, U.M.M.Z. (1); between Laguna de Catemaco and Volcan San Martin, LI.M.M.Z. (1); Volcan San Mar- tin, U.I.M.N.H. (7), U.M.M.Z. (6). GUATEMALA: Alta Verapaz: Chinaj;'i, K.U. (3, 1 skeleton). El Peten: 10 kilometers north-northwest of Chinaja (Alta Verapaz), K.U. ( 1 ); Piedras Negras, F.M.N.H. (3), U.I.M.N.H. (1), U.S.N.M. (8); 8 kilometers south of Piedras Negras, F.M.N.H. (1); Semicoch, U.S.N.M. (1). Smilisca phaeota NICARAGUA: Matagalpa: Finca Tepeyac, 10 kilometers north, 9 kilometers east of Matagalpa, K.U. (1, 1 tadpoles); Matagalpa, M.C.Z. (2), U.M.M.Z. ( 1 ); 19 kilometers north of Matagalpa, U.M.M.Z. (2); Santa Maria de Ostuma, K.U. (1). Zelaya: Bonanza, K.U. (31, 3 skeletons, 2 tadpoles); Cukra, A.M.N.H. (1); El Recreo, K.U. (62); Rio Mico, 16 kilometers east of Recreo, U.M.M.Z. (10); junction of Rio Mico and Siguia, U.M.M.Z. (10); Rio Siguia, 11 kilometers northwest of Rama, U.M.M.Z. (49). COSTA RICA: Alajuela: Cinchona, K.U. (4); 5 kilometers south of Ciudad Quesada, U.S.C. (1); Laguna Monte Alegre, K.U. (2); Las Playuelas, 11 kilometers south of Los Chiles, U.S.C. (1); San Car- los, LI.S.N.M. (1). Cartago: Moravia de Turrialba, K.U. (41, 1 skeleton), U.S.C. (3); Peralta, K.U. (2); Rio Chitaria, 3 kilometers north-northeast of Pavones, K.U. (7, 1 eggs, 7 tadpoles); Rio Reventazon, M.C.Z. (8), U.M.M.Z. (9); Turrialba, K.U. (46, 3 skele- tons), M.C.Z. (3, 1 tadpoles), U.S.N.M. (1). Guana- caste: Tilaran, K.U. (3); 8 kilometers northeast of Ti'ariin, K.U. (2). Heredia: Barranca del Rio Sara- piqui below Isla Bonita, K.LT. (2); Cariblanco, K.U. (6, 1 skeleton), M.C.Z. (1); Isla Bonita, K.U. (5); Puerto Viejo, K.U. (1); 4.2 kilometers west of Puerto Viejo, K.LI. (2); 7.5 kilometers west of Puerto Viejo, K.U. (1). Limon: Bambu U.S.C. (4); Batan, U.M.M.Z. (1); Coen, M.C.Z. (1); La Lola, K.U. (3), U.F. (1), M.C.Z. (3); Los Diamantes, F.M.N.H. (4), K.U. (6); Pandora, U.M.M.Z. (2); U.S.C. (4); Puerto Limon, K.U. (1); Rio Lari at Rio Dipari, 21 kilometers southwest of Amubre, U.S.C. (1); Rio Toro Amarillo, 7 kilometers west of Gualipes, K.U. (1, 1 tadpoles); Suretka, K.U. (4). Puntarenas: Agua Buena, K.U. (1); 1.6 kilometers east of Buenos Aires, U.M.M.Z. (1); 3 kilometers northwest of Buenos Aires, K.U. (1); 4 kilometers north, 15 kilo- meters west of Dominical, K.U. (2 tadpoles); Esparta, M.C.Z. (3); Golfito, K.U. (1); 6 kilometers east of Golfito, K.U. (2 skeletons); Gromaco, U.M.M.Z. (4); Palmar, K.U. (1); 4 kilometers east-southeast of Pal- mar Sur, K.U. (2); 5.6 kilometers southeast of Palmar Sur, K.U. (1 tadpoles); 7.0 kilometers southeast of Palmar Sur, K.U. (1 tadpoles); 8.5 kilometers south- east of Piedras Blancas, K.U. (12); Quebrada Boruca, 22 kilometers east of Palmar Norte, K.U. (1); Rin- c6n de Osa, K.U. (15, 2 tadpoles), U.M.M.Z. (3), L'.S.C. (1); Rio Ferruviosa, 7 kilometers south of Rincon de Osa, U.S.C. (1); 1.6 kilometers west- 728 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 northwest of Villa Neily, K.U. (2 tadpoles). San Jose: San Isidro el General, K.U. ( 1 ), U.M.M.Z. ( 1 ); 10 kilometers north of San Isidro el General, M.C.Z. (5); 13 kilometers west-southwest of San Isidro el General, K.U. (1); 15 kilometers west-southwest of San Isidro el General, K.U. (3 tadpoles); 20 kilo- meters west-southwest of San Isidro el General, K.U. (1). PANAMA: Bocas del Tow: Alniirante, K.U. (3); 1.6 kilometers west of Alniirante, K.U. (I); 3 kilo- meters west of Alniirante (10, 1 skeleton), 11 kilo- meters northwest of Alniirante, F.M.N.H. (9); 13 kilometers west of Alniirante, K.U. (6, 3 skeletons); Fish Creek, K.U. (1); Isla Popa, K.U. (2); mouth of Rio Cahuita, K.U. (1); Rio Changena, 650 meters, K.U. (2); Rio Changena, 830 meters, K.U. (8). Canal Zone: Barro Colorado Island, F.M.N.H. (9), M.C.Z. (5), U.F. (1), U.M.M.Z. (18); 3.7 kilometers west of Cocoli, K.U. (1); Fort Sherman, M.C.Z. (1); Gatun, M.C.Z. (1); junction roads C25B and C16, T.N.H.C. (1); Madden Forest, K.U. (1), T.N.H.G. (2); Rio Agua Salud, 13 kilometers northwest of Ganiboa, K.U. (5). Chiriqui: 2 kilometers west of Concepcion, A.M.N.H. (1); Progreso, U.M.M.Z. (5). Code: El Valle, K.U. (4, 1 tadpoles), T.N.H.C. (1). Colon: Achiote, K.U. (5, 1 tadpoles); Quipo, A.M.N.H. (2); Rio Candelaria, F.M.N.H. (2). Darien: Camp Creek, Camp Townsend, A.M.N.H. (3); Cana, K.U. (1), U.S.N. M. (1); Rio Chucuna- que, 7 kilometers above Rio Morti, K.U. (2); Rio Esnape, Sambu Valley, M.C.Z. (1); Rio Subcuti, Chaiichiman's Creek, A.M.N.H. (I); Rio Tuira at Rio Mono, K.U. (1). Panama: northwest slope of Cerro Prominente, K.U. (1); Finca La Sumbadora, K.U. (1 skeleton). Son Bias: Arniila, U.S.N. M. (1); Sasardi, K.U. (4). Veraguas: mouth of Rio Concep- cion, K.U. (1). Smilisca puma NICARAGUA: No specific locality, U.S.N.M. (1). COSTA RICA: Alajuela: Jabillos, 5 kilometers north of Santa Clara, U.S.C. (6); 5 kilometers west of La Fortuna, U.S.C. (2); Rio La Fortima at La Fortuna, L'.S.C. (3). Cariago: Lagima Bonilla, tun- nel camp near Peralta, K.U. (1). Heredia: Puerto Viejo, K.U. (2), M.C.Z. (2); 5.9 kilometers west of Puerto Viejo, K.U. (1); 7.5 kilometers west of Puerto Viejo, K.U. (18, 7 skeletons, 2 tadpoles). Limon: Batan, K.U. (3); La Losa, K.U. (1), U.S.C. (3); Los Diamantes, K.U. (1), U.M.M.Z. (6), U.S.C. (1); 2.4 kilometers east of Los Diamantes, U.S.C. (5). Smilisca sila COSTA RICA: Puntarenas: 6 kilometers east of Golfito, K.U. (1); Quebrada Boruca, 22 kilometers east of Palmar Norte, K.U. (2); Rio Zapote, 8 kilo- meters east of Palmar Norte, U.S.C. (2). San Jose: San Isidro el General, K.U. (1); 14 kilometers north- west of San Isidro el General, U.S.C. (2); 15 kilo- meters west-southwest of San Isidro el General, U.S.C. (I). PANAMA: Canal Zone: Barro Colorado Island, A.M.N.H. (4), F.M.N.H. (10), K.U. (7, 1 skeleton, 1 tadpoles), U.M.M.Z. (5), U.S.C. (1). Chiriqui: Boquete, A.M.N.H. (1), U.M.M.Z. (5); El Volc;in, K.L^ (24, 4 skeletons, 1 eggs, 1 tadpoles); 6 kilome- ters south of El Volcan, F.M.N.H. (1); 16 kilometers north-northv\est of El Volcan, K.U. (12); Finca Palo- santo, 6 kilometers v\est-northwest of El Volcan, K.U. (12, 1 skeleton); Finca Santa Clara, K.U. (1); Rio Colorado, 17 kilometers north-northwest of El Volcjin, K.U. (2); Valle Hornito, 19 kilometers northeast of Gua'aca, K.U. (13). Code: El Valle, A.M.N.H. (21), F.M.N.H. (20), K.U. (3, 1 tadpoles), T.N.H.C. (2), U.S.N.M. (1). Darien: Camp Creek, Camp Townsend, A.M.N.H. (7); Rio Chico, A.M.N.H. (3); Rio Pita. F.M.N.H. (3); Tacarcuna, U.S.N.M. (7); Three Falls Creek, A.M.N.H. (2). Los Santos: Cerro Cambutal, K.U. (9); Cerro Hoya, K.U. (5, 2 tad- poles), U.S.N.M. (2); Guanico Arriba, Rio Guanico, K.U. (3, 1 tadpoles); Lajamina, Rio Puira, K.U. (1). Panama: Altos de Pacora, K.U. (1); Cerro Jefe, K.U. (2); Cerro La Campana, F.M.N.H. (1), K.U. (5), U.S.N.M. (1); Finca La Sumbadora, K.U. (15, 2 skeletons, 1 eggs, 4 tadpoles); Rio Calobra, U.S.N.M. ( 1 ) ; Rio Pacora, 9 kilometers north-northeast of Pacora, K.U. (I). San Bias: Sasardi, K.U. (17, 5 skeletons). Veraguas: Cerro Carbunco, U.S.N.M. (1); Cerro Tute, F.M.N.H. (5); Isla Cebaco, Rio Platanal, K.U. (3). Smilisca sordida NICARAGUA: Zelaya: Rio Grande, M.C.Z. ( 1 ) COST.iV RICA: Alajuela: between Atena and Sa'to de San Mateo, U.S.C. (1); 8 kilometers north of Ciudad Quesada, U.S.C. (4); La Fortuna, U.S.C. (20); 3 kilometers east of La Fortuna, U.S.C. (1); San Carlos, U.S.N.M. (1); Sarchi, K.U. (12). Car- iago: Cartago, B.M.N.H. (I); headwaters of Rio Pacuare, U.S.C. (1); 10 kilometers north of Rio Reventazon bridge, Lf.S.C. (1); 5 kilometers south- west of Rio Reventazon bridge on Paraiso-Orosi road, U.S.C. (1); Turrialba, K.U. (1), M.C.Z. (1), U.M.M.Z. (1), U.S.C. (3), U.S.N.M. (4). Heredia: Puerto Viejo, K.U. (1); 5 kilometers west of Puerto Viejo, T.C.W.C. (15). Guanacaste: Las Cafias, U.S.C. (1); Santa Cecilia, M.C.Z. (2); Tilaran, use. (5). Limon: Bambii, U.S.C. (15); La Lola, U.S.C. (10); Pandora, U.S.C. (16); Pico Blanco, U.S.N.M. (2); Rio Lari, 14-16 kilometers southwest of Amubre, U.S.C. (11); Sipurio, U.S.N. NL (2); Suretka, K.ll. (14, 1 skeleton). Puntarenas: 6 kilo- meters north of Dominical, K.U. (2, 2 tadpoles); Esparta, M.C.Z. (1); 6 kilometers east of Golfito, K.U. (24, 4 skeletons, 2 tadpoles), U.S.C. (23); Quebrada Auga Buena, 3 kilometers southwest of Rincon de Osa, U.S.C. (6); Quebrada Boruca, 22 kilometers east of Palmar Norte, K.U. (1); Rincon de Osa, K.U. (44, 3 tadpoles), U.M.M.Z. (6, 1 skele- ton), U.S.C. (7); Rio Barranca, U.S.C. (2); Rio Ceiba, 6 kilometers northwest of Buenos Aires, K.U. (2), U.S.C. (7); Rio Ciruelitas, 16 kilometers north- west of Esparta, U.S.C. (3); Rio Claro, 14.2 kilo- 1970 DUELLMAN: HYLID FROGS 729 meters northwest of Villa Neily, U.S.C. (4); Rio Ferriuiosa, 7 kilometers south of Rincon de Osa, U.S.C. (4); Rio Lagarto at Pan-American Highway (Guanacaste border), U.S.C. (4); Rio La Vieja, 30 kilometers east of Palmar Norte, K.U. (4, 1 tadpoles), U.S.C. (2); Rio Oro, 28.5 kilometers northwest of Villa Neily, K.U. (1); Rio Volcan, 10 kilometers west of Buenos Aires, U.S.C. (1); Rio Zapote, 8 kilometers east of Palmar Norte, U.S.C. (4); 3-5 kilometers west of Palmar, U.S.C. (18); 7 kilometers southeast of Palmar Sur, K.U. (3); 1-5 kilometers northwest of Villa Neily, U.S.C. (23). San Jose: Bajos de Jorco, K.U. (1 tadpoles); Escazu, K.U. (8), U.S.C. (1); Paso Ancho, Rio Jorco, U.M.M.Z. (6), U.S.C. (3); Rio Jorco, near Desamparados, K.U. (11, 4 skele- tons), U.S.C. (9); Rio Peje, 10 kilometers south- southeast of San Isidro el General, U.S.C. 7115, (3); Rio Tirivi, M.C.Z. (1); San Isidro el General, F.M.N.H. (1), K.U. (2), U.M.M.Z. (1); 15 kilo- meters west-southwest of San Isidro el General, K.U. (19, 3 tadpoles), U.S.C. (6); 17.1 kilometers west- southwest of San Isidro el General, U.S.C. (1); 18 kilometers west-southwest of San Isidro el General, U.S.C. (1); 20 kilometers west-southwest of San Isidro el General, K.U. (6, 3 skeletons, 6 tadpoles); San Jose, A.M.N.H. (4), U.S.C. (1); Santa Rosa, U.S.C. (3). P.A.NAMA: Chiriqui: Rio Jacu, 5.8 kilometers east-southeast of Paso Canoas, K.U. (1). Veraguas: No specific locality, Z.M.B. (2). Triprion petasatus MEXICO: Campeche: 5 kilometers south of Champoton, K.U. (1); Dzibalchen, K.U. (14); 7.5 kilometers west of Escarcega, K.U. (4). Quintana Roo: 6.5 kilometers south of Las Palmas, 57 kilome- ters south of Felipe Carrillo Puerto, U.I. M.N. H. (4). Yucatan: Cenote Tamanche, U.S.N.M. (1); Chichen Itza, F.M.N.H. (3), U.M.M.Z. (76); 2.5 kilometers east of Chichen Itza, K.U. (11, 1 skeleton); 9 kilo- meters east of Chichen Itza, K.U. (17, 2 tadpoles); 12 kilometers east of Chichen Itza, K.U. ( 19, 3 skele- tons, 1 tadpoles); Dzibichaltun, K.U. (4); 6 kilome- ters south of Merida, K.U. (1 tadpoles); 7 kilometers north of Muna, K.U. (1); Piste, (2, 2 skeletons); 3.5 kilometers north of Piste, K.U. (1 tadpoles); Santa Elena, 4.8 kilometers south of Talcha, U.M.M.Z. (1); Tekom, F.M.N.H. (14); 8.8 kilometers southeast of Ticul, U.M.M.Z. (1); 3.5 kilometers east of Yokdzo- not, K.U. (35, 2 skeletons, 4 eggs, 2 tadpoles). GUATEMALA: El Peten: La Libertad, F.M.N.H. (2), S.U. (1), U.M.M.Z. (34, 1 skeleton); Tikal, U.F. (8). Triprion spatulatus reticulatus MEXICO: Colima: 10.5 kilometers south of Colima, U.I.M.N.H. (67); 19.5 kilometers south of Colima, U.I.M.N.H. (3); 21 kilometers south of Colima, K.U. (3), T.N.H.C. (5); 5.6 kilometers north of Los Asmoles, A.M.N.H. (2); 22.2 kilometers east of Manzanillo, U.I.M.N.H. (27); Santiaguito, 11 kilometers north of Colima, U.I.M.N.H. (196); 10.9 kilometers north of Santiaguito, U.I.M.N.H. (12); Tecolapa, U.I.M.N.H. (10). Guerrero: 5 kilometers east of El Zapote, K.U. (31, 6 skeletons). Michoacdn: Ostula, U.M.M.Z. (8); between Rio Marquez and Cuatro Caminos, K.U. (3). Oaxaca: Cerro Arenal, U.S.N.M. (1); Cerro San Pedro, U.I.M.N.H. (1); Chivela, A.M.N.H. (1), M.C.Z. (1); Garza Mora, U.I.M.N.H. (1); 2.4 kilometers north of Salina Cruz, K.U. (1, 1 skeleton), U.M.M.Z. (26, 2 skeletons, 2 tadpoles); San Antonio, near Tehuantepec, C.A.S. (1), T.C.W.C. (1), U.I.M.N.H. (3); Tehuantepec, U.I.M.N.H. (2); 8.6 kilometers west of Tehuantepec, K.U. (1), U.M.M.Z. (11, 2 skeletons). Triprion spatulatus spatulatus MEXICO: Sinaloa: 4 kilometers northeast of Concordia, K.U. (2); 3 kilometers east of Concordia, L.B.S.C. (7, 1 skeleton), 5 kilometers southwest of Concordia, K.U. (8); 8 kilometers southwest of Con- cordia, K.U. (4); 88 kilometers south of Culiacan, K.U. (4, 3 skeletons); 36.5 kilometers south of El Salado, U.I.M.N.H. (1); 10 kilometers northeast of La Cruz, L.A.C.M. (51); Mazatlan, K.U. (1), M.C.Z. (1, 1 skeleton); 6.6 kilometers north of Mazatlan, L.B.S.C. (1); 11 kilometers north of Mazatlan, L.B.S.C. (1 skeleton); 13 kilometers north of Mazat- lan, L.B.S.C. (1); 14.4 kilometers north of Mazatlan, A.M.N.H. (1); 25 kilometers north of Mazatlan, L.B.S.C. (2); 31 kilometers north-northwest of Mazadan, A.M.N.H. (1 skeleton), K.U. (1 skeleton), U.M.M.Z. (15); road to San Ignacio, L.A.C.M. (6); Venadillo, U.S.N.M. (1); 9.1 kilometers northeast of Villa Union, K.U. (9); 15.4 kilometers northeast of Villa Union, L.A.C.M. (1); 21 kilometers southeast of Villa Union, L.A.C.M. (1); 26 kilometers south- east of Villa Union, L.A.C.M. (1). APPENDIX 2 The data for the specimens, recordings, and photographs comprising the illustrations on plates 1-72 are given below. PLATE 1. 1. Htjla zeteki, K.U. No. 36481, La. Palma, San Jose Province, Costa Rica. 2. Hyla mixe, K.U. No. 87100, 4.2 kilometers south of Campamento Vista Hermosa, Oaxaca, Me>uco. 3. Hyla regilla curta, K.U. No. 78370, Todos Santos, Baja California Sur, Mexico. PLATE 2. 1. Pternohyla dentata, K.U. No. 60083, 15 Idlonie- ters east of Aguascalientes, Aguascalientes, Mexico. 2. Hyla cchinata, U.M.M.Z. No. 123987, Campamento Vista Hermosa, Oaxaca, Mexico. 3. Hyla valancifcr, K.U. No. 95416, Volcan San Martin, Veracruz, Me.xico. PLATE 3. 1. Hyla fimbrimembra, R.C.T. No. 761, Cinchona, Alajuela, Costa Rica. 2. Hyla thysanota, U.S.N.M. No. 151080, Cerro Mali, Darien Province, Panama. PLATE 4. 1. Hyla pachyderma, U.S.N.M. No. 115026, Pan de Olla, Veracruz, Mexico. 2. Hijla crassa, U.l.M.N.H. No. 25050, Cerro San Felipe, Oaxaca, Mexico. 3. Plec- trohyla lacertosa, U.LM.N.H. No. 33693, "Region de Soconusco," Chiapas, Mexico. PLATE 5. 1. Plectrohyla pycnochila, T.C.W.C. No. 21459, 5 Idlonieters north-northwest of san Cristobal de las Casas, Chiapas, Mexico. 2. Plecirohyla hartwegi, U.M.M.Z. No. 94428, Barrejonel, Chiapas, Me.xico. PLATE 6. Hyla miliaria, K.U. No. 101610 Finca Santa Clara, Chiriqul Province, Panama; gliding pose, from a field sketch by Linda Trueb. PLATE 7. Hemipractus panamcnsis, B.Y.U. No. 19142, Rio Changena, Darien Province, Panama; female carrying young, each attached to dorsum by a pair of double- stranded cords. PLATE 8. Egg clutches: 1. Hyla pseudopuma pscudopuma in shallow pond at Tapanti, Cartago Province, Costa Rica. 2. Hyla lancasteri on herb above stream on the north slope of Cerro Pando, Bocas del Toro Province, Panama. 3. Hyla thorectes on fern above stream at 37 kilometers north of San Gabriel Mixtepec, Oaxaca, Mexico. 4. Agalychnis annae, on branch above pond at La Palma, San Jose Province, Costa Rica. PLATE 9. 1. Nests of Hyla boans in creek at the Rio Sasardi, Camp Sasardi, San Bias Province, Panama; note machete for scale. 2. Close-up of nest of Hyla boans containing small tadpoles. Both photographs by Charles W. Myers. PLATE 10. 1. Pond at Puerto Viejo, Heredia Province, Costa Rica. Ten species of hylids (Hyla boulengeri, elaeoch- roa, loquax, phlebodes, ebraccata, Smilisca baudinii, phaeota, puma, Agalychnis calUdryas, and A. saltator) are known to breed in the pond; photographed on June 21, 1966. 2. Pond at 4 kilometers west-north- west of Esparta, Puntarenas Province, Costa Rica. Four species of hylids (Hyla staufferi staiiffcri, H. microccphala underwoodi, Smilisca baudiriii, and Phrynohyas venulosa) are known to breed in the pond; photographed on June 22, 1961. PLATE 11. Stream in cloud forest at 3 kilometers southwest of Huatusco, Veracruz, Mexico, elevation 1325 meters. Hylids found along this stream include Hyla mio- tympanum, mixomaculata, mibicola, and taeniopus. Hyla dendroscarta, picta, Smilisca baudinii, Phryno- hyas venulosa, and Aga/yc/iriis moreletii occur in the cloud forest. PLATE 12. Mating calls: 1. Hyla regilla curta, K.U. Tape No. 271; Todos Santos, Baia California Sur, Me.xico; July 9, 1963; 28°C. 2. Hyla cadaverina, A.M.N.H. Tape No. 7-6; Sentenac Canon, San Diego County, Cali- fornia; March 24, 1956; 15.5°C. 3. Hyla arenicolor, A.M.N.H. Tape No. 76-1; Chiricahua Mountains, Cochise County, Arizona; June 28, 1958; 23.9°C. PLATE 13. Mating calls of Hyla cximia. 1. K.U. Tape No. 596; 8 kilometers northwest of Queretaro, Queretaro, Mexico; June 15, 1966; 21.6"'C. 2. A.M.N.H. Tape No. 60-2; 16 kilometers southwest of Huachinango, Puebla, Mexico; August 9, 1956; 17.6°C. 3. A.M.N.H. Tape No. 129-1; 1.6 kilometers east of Buenos Aires, Durango, Mexico; July 3, 1963; 15°C.; second in- dividual in background. PLATE 14. Mating calls: 1. Hyla euphorbiacea, K.U. Tape No. 589; 6 kilometers southeast of Oaxaca, Oaxaca, Mexico; August 6, 1966; 21.6°C. 2. Hyla walkeri. University of Texas Tape No. 179; 1.6 kilometers northwest of Pueblo Nuevo Solistahuacan, Chiapas, Mexico; June 12, 1958; 19°C. 3. Hyla plicata, K.U. Tape No. 597; El Chico Parque Nacional, Hidalgo, Mexico; June 16, 1966; 19°C. PLATE 15. Mating calls: 1. Hyla miotympanum, K.U. Tape No. 188; south slope of Volcan San Martin Tuxtla, Veracruz, Mexico; August 11, 1960; 30°C. 2. Hyla arborescandens, K.U. Tape No. 198; 6.5 kilometers south of Campamento Vista Hermosa, Oaxaca, Mex- ico; June 27, 1962; 19.7°C. 3. Hyla eryihromma, K.U. Tape 350; 8 kilometers south of Yetla, Oaxaca, Mex- ico; June 16, 1964; 21.6''C., stream in background. PLATE 16. Mating calls: 1. Hyla thorectes, K.U. Tape No. 575; 37 kilometers north of San Cabriel Mixtepec, Oaxaca, Me.xico; August 2, 1966; 17.3°C.; stream in background. 2. Hyla hazelae, K.U. Tape No. 579; 2 kilometers south of El Punto, Oaxaca, Me.xico; August 8, 1966; 18.3°C.; stream in background. 3. Hyla loquax, University of Texas Tape No. 283; 1 kilometer east of Rio Tonola, Tabasco, Mexico; June 30, 1959; 25.5°C.; insects in background. 730 1970 DUELLMAN: HYLID FROGS 731 PLATE 17. Mating calls: 1. Hyla godmani. University of Texas Tape No. 280; 5 Idlonieters east-southeast of Cordoba, Veracraz, Me.xico; June 23, 1959; 28.5°C. 2. Hyla melanomma melanomma, K.U. Tape No. 340; 12 kilometers north-northwest of San Gabriel Mix- tepec, Oaxaca, Me.xico; June 20, 1964; 18.8°C; long note, short note, and section of short note. 3. Hijla melanomma bivocata, K.U. Tape No. 160; 6.2 kilome- ters south of Rayon Mescalapa, Chiapas, Me.xico; August 5, 1960; 18.3°C.; long note, short note, and section of short note. PLATE 18. Mating calls: 1. Htjla bwmeliacia, K.U. Tape No. 162; Finca Chicoyou, near Coban, Departamento Alta Verapaz, Guatemala; July 17, 1960; 20.7°C. 2. Hyla sumichrasti, K.U. Tape No. 581; Portillo Nejapa, 14 kilometers east of El Camaron, Oaxaca, Mexico; August 9, 1966; 20.7°C. 3. Htjla chaneque, K.U. Tape No. 356; 4.2 kilometers south of Canipa- mento Vista Hemiosa, Oaxaca, Mexico; June 17, 1964; 21.7°C. PLATE 19. Mating calls: 1. Hyla picta, Univ. Texas Tape No. 279; Villa Juarez, Puebla, Mexico; June 18, 1959; 19°C. 2. Hyla ^nthii, K.U. Tape No. 343; 17 kilo- meters north-northwest of San Gabriel Mixtepec, Oaxaca, Mexico; June 20, 1964; 19°C. 3. Hyla rivularis, K.U. Tape No. 550; Rama Sur Rio Las Vueltas, Heredia Province, Costa Rica; March 29, 1966; 18.4''C. PLATE 20. Mating calls: 1. Hyla pseudopuma pseudopuma, K.U. Tape No. 564; Cinchona, Alajuela Province, Costa Rica; April 5, 1966; 17.8°C. 2. Htjla angus- lilineata, K.U. Tape No. 562; Rama Sur Rio Las Vueltas, Heredia Pro\ince, Panama; March 31, 1966; 12.8°C. 3. Hyla boans, K.U. Tape No. 601; Camp Sasardi San Bias Province, Panama; January 12, 1967; 26°C. PLATE 21. Mating calls: 1. Hyla tica, K.U. Tape No. 547; Tapanti, Cartago Province, Costa Rica; March 26, 1966; 21.0°C. 2. Hyla debilis, K.U. Tape No. 559; north slope of Cerro Pando, Bocas del Toro Province, Panama; May 29, 1966; 18°C. 3. Hyla uranochroa, K.U. Tape No. 208; Cinchona, Alajuela Province, Panama; June 12, 1961; 18.6''C. PLATE 22. Mating calls: 1. Hyla rufioculus, K.U. Tape No. 337; 14 kilometers north of San Isidro el General, San Jose Province, Costa Rica; July 19, 1964; 15.6°C; stream in background. 2. Hyla salvadorensis, K.U. Tape No. 569; west slope of Cerro Uyuca, Departa- mento Francisco Morazan, Honduras; July 5, 1966; 19.3°C. 3. Hyla legleri, K.U. Tape No. 210; 15 kilometers southwest of San Isidro el General, San Jose Province, Costa Rica; April 6, 1961; 23.5°C. PLATE 23. Mating calls: 1. Hyla pictipes, K.U. Tape No. 147; Rio Poasito, 1 kilometer west of Poasito, Alajuela Province, Costa Rica; June 28, 1961; 15°C. 2. Hijla colymba, K.U. Tape No. 290; Laguna, Darien Prov- ince, Panama; July 4, 1963; 24°C. 3. Hyla rufitela. K.U. Tape No. 526; Barro Colorado Island, Canal Zone, Panama; June 24, 1965; 27°C.; insects in back- ground. PLATE 24. Mating calls: 1. Htjla lancasteri, K.U. Tape No. 202; 3 kilometers south of Pavones, Cartaeo Province, Costa Rica; June 7, 1961; 21.5°C. 2. Hyla lancasteri, K.U. Tape No. 513; north slope of Cerro Pando, Bocas del Toro Province, Panama; May 28, 1966; 16.8°C.; stream in background. 3. Anotheca spinosa, K.U. Tape No. 357; 11 kilometers north of Campamento Vista Hermosa, Oaxaca, Mexico; June 28, 1964; 21.5°C. PLATE 25. Mating calls: 1. Htjla crepitans, K.U. Tape No. 273; Camp Chagres, Canal Zone, Panama; June 18, 1963; 25°C.; Engijstomops ptisttilosus in background. 2. Possible natural hybrid between Htjla crepitans and Hijla rosenbergi, A.M.N.H. Tape No. 122; Rio Bejuco, Panama Province, Panama; June 1, 1962; 26°C. 3. Hyla rosenbergi, K.U. No. 529; Rio Tuira at Rio Mono, Darien Province, Panama; July 25, 1965; 26.4°C. PLATE 26. Mating calls: 1. Hijla elaeochroa, K.U. Tape No. 98; Turrialba, Cartago Province, Costa Rica; June 9, 1961; 21°C. 2. Hijla statifferi staufferi, K.U. Tape No. 93; San Salvador, Departamento San Salvador, El Salvador; July 10, 1960; 24°C. 3. Hyla statifferi altae, K.U. Tape No. 502; 2 kilometers west-southwest of Chepo, Panama Province, Panama; June 5, 1966; 25.6°C. PLATE 27. Mating calls: 1. Hyla rubra, K.U. Tape No. 612; Santa Cecilia, Napo Province, Ecuador; March 10, 1967; 23.5°C. 2. Hijla boiilengeri, K.U. Tape No. 511; Summit Gardens, Canal Zone, Panama; June 8, 1966; 28.5°C. 3. Htjla rostrata, K.U. Tape No. 288; 3 kilometers west-southwest of Chepo, Panama Prov- ince, Panama; June 28, 1963; 27°C. PLATE 28. Mating calls: 1. Htjla microcephala microcephala, K.U. Tape No. 19; Palmar Sur, Puntarenas Province, Panama; April 18, 1961; 28''C. 2. Hyla phlebodes, K.U. Tape No. 6; Puerto Viejo, Heredia Province, Costa Rica; June 16, 1961; 29.3°C.; second individual in background. 3. Htjla robertmcrtensi, K.U. Tape No. 41; 32 kilometers north of Arriaga, Chiapas, Mex- ico; August 4, 1960; 28.5°C. PLATE 29. Mating calls: 1. Htjla ebraccata, K.U. Tape No. 129; Moravia de Turrialba, Cartago Province, Costa Rica; July 7, 1961; 19°C. 2. Htjla sttbocularis, K.U. Tape No. 285; Laguna, Darien Province, Panama; July 6, 1963; 23.4°C. 3. Htjla sartori, Univ. Texas Tape No. 293; 6 kilometers east of Tecpan de Gale- ana, Guerrero, Mexico; July 25, 1959; 26°C. PLATE 30. Mating calls: 1. Ptychohyla schmidtorum chamti- lae, K.U. Tape No. 52; 6.2 kilometers .south of Rayon Mescalapa, Chiapas, Mexico; August 5, 1960; 19°G. 2. Ptychohyla ignicolor, K.U. Tape 399; 4.2 kilometers south of Rayon Mescalapa, Chiapas, Me.xico; June 17, 1964; 20.7°C. 3. Rain call of Agalychnis callidrtjas. 732 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 K.U. Tape No. 61; 3 kilometers southeast of La Libertad, Departamento El Peten, Guatemala; July 1, 1960; 30°C. PLATE 31. Mating calls: 1. Plychohyla cutliysanota macro- tympanum, K.U. No. 48; Rio Hondo, 9.5 kilometers south of Pueblo Nuevo Solistahuacan, Chiapas, Me.x- ico; June 16, 1960; 18°C. 2. Ptychohyla leonhard- schultzei, U.M.M.Z. Tape No. 525; Campamento Vista Hermosa, Oaxaca, Mexico; March 30, 1959; 19.5°C. 3. Ptychohyla spinipoUex, K.U. Tape No. 53; Finca Los Alpes, Departamento Alta Verapaz, Guate- mala; July 31, 1961; 22.8°C.; stream in background. PLATE 32. Mating calls: 1. Smilisca baudinii, K.U. Tape No. 74, 1.5 kilometers northwest of Coluteca, Departa- mento Choluteca, Honduras; July 25, 1961; 19°C. 2. Smilisca cyanosticta, K.U. Tape No. 373; 11 kilome- ters north of Campamento Vista Hermosa, Oaxaca, Mexico; June 28, 1964; 21.3°C. 3. Smilisca phaeota, K.U. Tape No. 79; 7 kilometers southeast of Piedras Negras, Puntarenas Province, Costa Rica; April 10, 1961; 25°C. PLATE 33. Mating calls: 1. Smilisca puma, K.U. Tape No. 382; 7.5 kilometers west of Puerto Viejo, Heredia Province, Costa Rica; February 19, 1965; 23.5°C. 2. Smilisca sila, K.U. Tape No. 385; El Volcan, Chiri- qui Province, Panama; February 5, 1965; 18°C. 3. Smilisca sordida, K.U. Tape No. 398; Rio Jorco, 2 kilometers south of Desamparados, San Jose Province, Costa Rica, February 18, 1965; 20.5°C. PLATE 34. Mating calls: 1. Pternohyla fodicns, K.U. Tape No. 584; 16 kilometers north of Mazatlan, Sinaloa, Mexico; August 24, 1967; 26.3°C. 2. Triprion spatu- laius reticulatus, L.A.C.M. Tape (specimen L.A.C.M. No. 36815); 12 kilometers southwest of Colima, Co- lima, Me.xico; July 12, 1967; 26.2°C. 3. Triprion petasatus, K.U. Tape No. 218; 2.5 kilometers east of Chichcn Itza; Yucatan, Mexico; July 23, 1962; 26.5°C. PLATE 35. Mating calls: 1. Acris crepitans, K.U. Tape No. 331; Rockefeller Experimental Tract, Douglas County, Kansas; May 8, 1964; 15.5°C.; other individuals in background. 2. Phyllomedusa lemur, K.U. Tape No. 67; La Palnia, San Jose, Province, Costa Rica; May 8, 1961; 17.7°C.; band of insect noises at about 3.500 cycles per second in background. 3. Plectrohyla ixil. K.U. Tape No. 543; 6.2 kilometers south of Rayon Mescalapa, Chiapas, Me.\ico; February 24, 1966; 15°C.; stream in background. PLATE 36. Mating calls: 1. Gastrotheca ceratophrys, K.U. Tape No. 595; Rio Claro near junction witli Rio Changena, Bocas del Toro Province, Panama; May 23, 1966; 18.5°C.; river in background. 2. Gastrotheca nicefori, K.U. Tape 600; South ridge of Cerro Citurio, Serrania de Pirre, Darien Province, Panani;i; January 24, 1966; 20°C.; band of insect noises in background. 3. Phrynohyas venulosa, K.U. Tape No. 593; Palmar Norte, Puntarenas Province, Costa Rica; April 8, 1966; 24.5°C. PLATE 37. Mating calls: 1. Pseudacris clarkii, A.M.N.H. Tape No. 147; 5 miles north of Independence, Mont- gomery County, Kansas; June 20, 1965; 21.5°C. 2. Pachymedusa dacnicolor, A.M.N.H. Tape No. 27; near Tepic, Nayarit, Mexico; August 16, 1956; 23°C. 3. Release call of Phyllomedusa venusta, K.U. Tape No. 527; Rio Tuira at Rio Mono, Darien Province, Panama; July 14, 1965; 25.5°C. PLATE 38. Mating calls: 1. Agalychnis saltator, K.U. Tape No. 487; 2 kilometers east of Tilaran, Guanacaste Province, Costa Rica; August 21, 1964; 22.3°C. 2. Agalychnis callidryas, K.U. Tape No. 61; 3 kilometers southeast of La Libertad, Departamento El Peten, Guatemala; July 1, 1960; 30°C. 3. Agalychnis more- letti, K.U. Tape No. 64; Finca Chicoyou, near Coban, Departamento Alta Verapaz, Guatemala; August 7, 1961; 21.5°C.; band of insect noises at about 2000 cycles per second in background. PLATE 39. Mating calls: 1. Agalychnis annae, K.U. Tape No. 55; Tapanti, Cartago Province, Costa Rica; April 19, 1961; 20°C. 2. Agalychnis spurrelli, K.U. Tape No. 295; Tacarcuna, Darien Province, Panama, July 16, 1963; 22°C. 3. Agalychnis litodryas, K.U. Tape No. 486; Rio Tuira at Rio Mono, Darien Province, Pana- ma; July 27, 1965; 16.3°C. PLATE 40. Release calls: 1. Pachymedusa dacnicolor, A.M. N.H. Tape No. 92; 32 Idlonieters east of Manzanillo, Colima, Mexico; July 22, 1959; 25°C. 2. Smilisca baudinii, A.M.N.H. Tape No. 94; 3 kilometers south of the Rio Cihuatlan, Colima, Mexico; July 16, 1959; 28.5°C. 3. Pternohyla fodiens, A.M.N.H. Tape No. 92; Chapala, Jalisco, Me.xico; July 14, 1959. PLATE 41. 1. Pachymedusa dacnicolor, U.M.M.Z. No. 115308, 1.6 kilometers northwest of Cuautlixco, Morelos, Me.x- ico. 2. Phyllomedusa venusta, K.U. No. 96150, Rio Tuira at Rio Mono, Darien Province, Panama. PLATE 42. 1. Agalychnis saltator, K.U. No. 86512 (night), 2 kilometers east of Tilaran, Guanacaste Province, Costa Rica. 2. Agalychnis callidryas, K.U. No. 96131, Rio Tuira at Rio Mono, Darien Province, Panama. 3. Agalychnis calcarifer, K.U. No. 77451, Laguna, Darien, Panama. 4. Agalychnis saltator, K.U. No. 86512 (day), 2 kilometers east of Tilaran, Guanacaste Province, Costa Rica. 5. Agalychnis callidryas, K.U. No. 63935, 4.2 kilometers west of Puerto Viejo, Heredia Province, Costa Rica. PLATE 43. 1. Agalychnis /noreletii, K.U. No. 57954, Finca Chicoyou, near Coban, Departamento Alta Verapaz, Guatemala. 2. Plnjllomcdusa lemur, K.U. No. 63940 (night), Tapanti, Cartago Province, Costa Rica. 3. Agalychnis annae, K.U. No. 64020, Tapanti Cartago Province, Costa Rica. 4. Agalychnis litodryas, K.U. No. 96149, Rio Tuira, at Rio Mono, Darien Province, Panama. 5. Phyllomedusa lemur, K.U. No. 63940 (day). Tapanti, Cartago Province, Costa Rica. 6. Agalychnis spurrelli, K.U. No. 77499, Barro Colorado Island, Canal Zone, Panama. 1970 DUELLMAN: HYLID FROGS 733 PLATE 44. 1. Hyla miliaria, K.U. No. 101610, Finca Santa Clara, Chiriqui Province, Panama. 2. Hemipliractus panamensis, K.U. No. 107422, south ridge of Ccrro Citurio, Serrania de Pirre, Darien Province, Panama. 3. Atwthcca spinosa, U.M.M.Z. No. 118173, south slope of N'olcan San Martin, Veracruz, Mexico. PLATE 45. 1. Gastrothcca nicefori, K.U. No. 111991, ridge between Rio Jaque and Rio Iniamado, Darien Prov- ince, Panama. 2. Gastrolheca ceratophrys, K.U. No. 77016, Laguna, Darien, Panama. PLATE 46. 1-4. Vhnjnohyas venulosa, respectively: U.M.N.Z. No. 104814, Barranca Bejuco, Michoacan, Me.\ico. U.M.M.Z. No. 115237, 5.3 kilometers east-northeast of Encinal, Veracruz, Mexico. U.M.M.Z. No. 119158, 3.5 kilometers south of Villahermosa, Tabasco, Me.x- ico. K.U. No. 64068, 4 kilometers west-northwest of Esparta, Puntarenas Province, Costa Rica. PLATE 47. 1. Hyla staufferi staufferi, U.M.M.Z. No. 115198, 7 kilometers east-southeast of Cordoba, Veracruz, Me.xico. 2. Hyla staufferi staufferi, U.M.M.Z. No. 119196, 6.1 kilometers west of Tuxt'a Gutierrez, Chiapas, Mexico. 3. Hyla staufferi altae, K.U. No. 116861, 6 kilometers soutli-southwest of Penonome, Code Pro\ince, Panama. 4. Hyla rubra, K.U. No. 109472, Santa Cecilia, Napo Province, Ecuador. 5. Hyla claeochroa, K.U. No. 64414, Turrialba, Cartago Province. Costa Rica. 6. Hyla elaeochroa, K.U. No. 64499, Laguna Monte Alegre, Alajuela Province, Costa Rica. PLATE 48. 1. Hyla boulengeri, K.U. No. 64321, 10.5 kilome- ters west-northwest of Villa Neily, Puntarenas Prov- ince, Costa Rica. 2. Htjla boulengeri, K.U. No. 95978, 3.2 Idlonieters west of Almirante, Bocas del Toro Province, Panama. 3. Hyla rostrata. No. 77164, 3 kilometers west of Chepo, Panama Province, Panama. PLATE 49. 1. Hyla nticrocephala microcephala, K.U. No. 64593, Palmar Sur, Puntarenas Province, Costa Rica. 2. Hijla microcephala underwoodi, K.U. No. 64565, Laeo de Yojoa, Departamento Cortes, Honduras. 3. Hyla robertmcrteusi, U.M.M.Z. No. 115243, 7 kilome- ters west-northwest of Tapanatepec, Oaxaca, Mexico. 4. Hyla phlebodes, K.U. No. 64798, Turrialba, Car- tago Province, Costa Rica. 5. Hyla sartori, U.M.M.Z. No. 119225, 12 kilometers west-southwest of Tierra Colorado, Guerrero, Mexico. 6. Hyla suboctdaris, K.U. No. 77348, Laguna, Darien Province, Panama. 7. Hyla ebraccata, K.U. No. 96005, 3.2 kilometers west of Almirante, Bocas del Toro Province, Panama. 8. Hyla ebraccata, K.U. No. 65120, 10.5 kilometers west- northwest of Villa Neilly, Puntarenas Province, Costa Rica. PLATE 50. 1. Hyla rufitela, K.U. No. 77307, Barro Colorado Island, Canal Zone, Panama. 2. Hyla crepitans, K.U. No. 80457, Finca La Sumbadora, Panama Province, Panama. 3. Hyla boans, K.U. 96013, juvenile, Rio Tuira at Rio Mono, Darien Province, Panama. 4. Hyla rosenbergi, K.U .No. 65015, 10.5 kilometers west-northwest of Villa Neily, Puntarenas Province, Costa Rica. PLATE 51. 1 and 2. Hyla boans, K.U. Nos. 108834 and 108835, respectively. Camp Sasardi, San Bias, San Bias Province, Panama. PLATE 52. 1. Hyla picadoi, K.U. No. 64872, Rio Poasito, 1 kilometer west of Poasito, Alajuela Province, Costa Rica. 2. Hyla colymba, K.U. 95979, Altos de Pacora, Panama Province, Panama. 3 and 4. Hyla angustilie- ata, K.U. Nos. 103590 and 103576, respectively, Rama Sur Rio las Vueltas, south slope of Volcan Barba, Heredia Province, Costa Rica. 5. Hyla pseudo- puma pseudopuma, K.U. No. 103715, Cinchona, Ala- juela Province, Costa Rica. 6. Hijla pseudopuma in- fucata, K.U. No. 101770, Rio Changena, 830 meters, Bocas del Toro Province, Panama. PL.4TE 53. 1. Hyla tica, K.U. No. 65132, Rio Maria-Aguilar, 3 kilometers west of Cariblanco, Alajuela Province, Costa Rica. 2. Hyla ricularis, K.U. No. 64986, La Concordia, Heredia Province, Costa Rica. 3. Hijla debilis, K.U. No. 101568, north slope Cerro Pando, 1450 meters, Bocas del Toro Province, Panama. 4. Hyla xanthosticta, K.U. No. 103772, Rama Sur Rio las Vueltas, south slope of Volcan Barba, Heredia Province, Costa Rica. 5. Hyla pictipes, K.U. No. 64646, c? , Rio Poasito, 1 kilometer west of Poasito, Alajuela Province, Costa Rica. 6. Hyla pictipes, K.U. No. 103605, 9 , Rama Sur Rio las Vueltas, south slope of Volcan Barba, Heredia Province, Costa Rica. PLATE 54. 1. Hyla rufioculus, K.U. No. 65140, 15 kilometers southwest of San Isidro el General, San Jose Province, Costa Rica. 2. Hyla lancasteri, K.U. No. 64729, 3 kilometers south of Pavones, Cartago Province, Costa Rica. 3. Hijla uranochroa, K.U. No. 65110, 3 kilome- ters south of Pavones, Cartago Province, Costa Rica. 4. Hyla lancasteri, K.U. No. 101736, north slope of Cerro Pando, 1920 meters, Bocas del Toro Province, Panama. 5. Hyla legleri, K.U. No. 64529, 15 kilome- ters southwest of San Isidro el General, San Jose Prov- ince, Costa Rica. PLATE 55. 1. Hyla smithii, U.M.M.Z. No. 115224, 1.6 kilo- meters south of Temixco, Morelos, Me.xico. 2. Hyla smithii, K.U. No. 57678, 3 kilometers north of Po- chutla, Oaxaca, Me.xico. 3. Hijla godmani, U.M.M.Z. No. 115171, 7 kilometers east-southeast of Cordoba, Veracruz, Mexico. 4. Hyla picta, U.M.M.Z. No. 115260, 7 kilometers east-southeast of Cordoba, Vera- cruz, Me.xico. 5. Hyla loquax, K.U. No. 103686, Puerto Veijo, Heredia Province, Costa Rica. PLATE 56. 1. Hyla miotympanurn, U.M.M.Z. No. 118155, 9 , Salto Cola de Caballo, Nuevo Leon, Me.xico. 2-4. Hyla miotympanurn, U.M.M.Z. No. 115290, 7 kilometers northeast of Huatusco, Veracruz, Mexico, showing metachrosis. 5. Hyla erythromma, K.U. No. 87104, 8 kilometers south of Yetla, Oaxaca, Mexico. PLATE 57. 1. Hyla hazelae, K.U. No. 100968, 2 kilometers south of El Punto, Oaxaca, Mexico. 2. Hyla thorectes. 734 MONOGRAPH MUSEUM OF NATURAL HISTORY NO. 1 K.U. No. 100947, 37 kilometers north of San Gabriel Mixtepec, Oaxaca, Mexico. 3. Hyla arborescandens, K.U. No. 64316 from Rio Octapa, 8 kilometers north- east of Tezuitlan, Puebla, Me.xico. 4. Hyla arbore- scandens, K.U. No. 61216 from 4.2 kilometers south of Campamento Vista Hermosa, Oaxaca, Me.xico. 5. Hijla valancifer, U.M.M.Z. No. 118157, south slope of Volcan San Martin, Veracruz, Me.xico. PLATE 58. 1. Hyla melanonmia melanomma, K.U. No. 86954, 12 kilometers north-northeast of San Gabriel Mix- tepec, Oaxaca, Me.xico. 2. Hyla melanomma bivocata, K.U. No. 58446, 6.2 kilometers south of Rayon Mes- calapa, Chiapas, Mexico. 3 and 4. Hyla pinorum, U.M.M.Z. Nos. 125371 and 125367, respectively, from 1.6 kilometers east of San Andreas della Cruz, Guer- rero, Me.xico. 5. Hyla mixomaculata, U.M.M.Z. No. 118171, 7.5 kilometers Huatusco, Veracruz, Mexico. 6. Hyla pellita, K.U. No. 100970, 33 kilometers north of San Gabriel Mixtepec, Oa.xaca, Mexico. 7. Hyla pinorum, K.U. 87612, juvenile, 3.3 kilometers north of San Vincente, Guerrero, Mexico. 8. Hyla nubicola, U.M.M.Z. No. 118160, 3 kilometers southwest of Huatusco, Veracruz, Mexico. PLATE 59. 1. Hyla sumichrasti, K.U. No. 100935, Portillo Nejapa, 14 kilometers east of El Camaron, Oaxaca, Mexico. 2. Hyla sumichrasti, K.U. No. 57851, 2 kilo- meters northwest of Pueblo Nuevo Solistahuacan, Chiapas, Me.xico. 3. Hyla smaragdina, K.U. No. 75301, Santa Lucia, Sinaloa, Mexico. 4. Hyla salva- dorensis, K.U. No. 103256. west slope of Cerro Uyuca, Departamento Francisco-Morazan, Honduras. 5. Hyla bromclacia, K.U. No. .57249, Finca Chicoyou, near Cohan, Departamento Alta Verapaz, Guatemala. 6. Hyla dendroscarta, U.M.M.Z. No. 118167, Mirador, Veracruz, Mexico. PLATE 60. 1. Hyla altipotens, K.U. No. 101009, 37 ki'ometers north of San Gabriel Mixtepec, Oaxaca, Me.xico. 2 and 3. Hyla chaneque, K.U. Nos. 58439, S , and 58442, 9 , respectively, 6.2 kilometers north of Rayon Mescalapa, Chiapas, Mexico. PLATE 61. 1 and 2. Hyla iaeniopus, U.M.M.Z. Nos. 118170, <5 , 7.5 kilometers southwest of Huatusco, and 118169, 9 , 3 kilometers southwest of Huatusco, Veracruz, Mexico, respectively. 3. Hyla altipotens, K.U. No. 101001, 37 kilometers north of Rayon Mescalapa, Oaxaca, Me.xico. PLATE 62. 1. Hyla histincta, U.M.M.Z. No. 119193, 12.5 kilometers east-northeast of Dos Aguas, Michoacan, Mexico. 2. Hyla bistincta, A.M.N. H. No. 76422, 1 kilometer northeast of Avutla, Oaxaca, Mexico. 3 and 4. Htjla pcntheter, K.U. Nos. 100932, i, and 100931, 9 , respectively, 37 kilometers north of San Gabriel Mixtepec, Oaxaca, Mexico. PLATE 63. 1. Hyla charodricola, U.M.M.Z. No. 118166, Rio Totolapa, 14.4 kilometers west of Huachinango, Pueb- la, Mexico. 2. Hyla chryses, U.M.M.Z. No. 125373, between Puerto Chico and Asoleadero, Guerrero, Mexico. 3. Hyla robertsorum, K.U. No. 57655, El Chico Parque Nacional, Hidalgo, Mexico. 4 and 5. Htjla siopela, K.U. Nos. 100990, juvenile, and 100977, (5 , west slope Cofre de Perote, Veracruz, Mexico. PLATE 64. 1. Hyla cadaverina, K.U. No. 109866, Palm Can- yon, Borrego, San Diego County, California. 2. Hyla arenicolor, U.M.M.Z. No. 115170, Agua del Obispo, Guerrero, Mexico. 3. Hyla arenicolor, U.M.M.Z. No. 119204 from Chinapa, Michoacan, Mexico. 4. Pseuda- cris clarkii, K.U. No. 1102.32, 2 miles south of Wau- netka, Jefferson County, Oklahoma. 5. Acris crepitans, K.U. No. 116930, Lawrence, Douglas County, Kansas. PLATE 65. 1-3. Hyla regilla hypochondriaca, K.V. Nos. 109875, 109876, and 109872, respectively, Ramona, San Diego County, California. 4. Hyla regilla hypo- clwndriaca, K.U. No. 109871, Borrego, San Diego County, California. 5. Hyla exitnia, U.N.M. No. 5637, 16 miles south of Springerville, Apache County, Ari- zona. PLATE 66. 1 and 3. Hyla eximia, K.U. Nos. 86993 and 86991, respectively, 3.5 kilometers west of Cuautlixco, Morelos, Me.xico. 2. Hyla plicata, K.U. No. 57389, El Chico Parque Nacional, Hidalgo, Mexico. 4 and 5. Hyla walkcri, K.U. Nos. 57832 and 57836,^ 18 kilo- meters northwest of Comitan, Chiapas, Me.xico. 6. Hijla euphorbiacea, K.U. No. 57346, 8 kilometers southeast of Oaxaca, Oaxaca, Mexico. PLATE 67. 1. Ptychohyla schmidtorum schmidtorum, K.U. No. 58035, Finca La Paz, near La Reforma, Departa- mento San Marcos, Guatemala. 2. Ptychohyla schmidtorum chamulae, K.U. No. 58069, 6.2 kilome- ters south of Rayon Mescalapa, Chiapas, Mexico. 3. Ptychohyla ignicolor, U.M.M.Z. 119062, Campa- mento Vista Hermosa, Oaxaca, Mexico. 4. Ptychohyla eulhysanota euthysanota, K.U. No. 58001, Finca La Paz, near La Reforma, Departamento San Marcos, Guatemala. 5. Ptychohyla euthysanota macrotym- paniim, K.U. No. 58047, Rio Hondo, 9.5 kilometers south of Pueblo Nuevo Solistahuacan, Chiapas, Mex- ico. 6. Ptychohyla Iconliardschultzei, U.M.M.Z. 115514, 8 kilometers south of Yetla, Oaxaca, Mexico. 7. Ptychohyla spinipollex, K.U. No. 58054, Finca Los Alpes, Departamento Alta Verapaz, Guatemala. PLATE 68. 1. Plectrohyla matudai. K.U. No. 58869, Finca La Paz, near La Reforma, Departamento San Marcos, Guatemala. 2. Plectrohyla ixil, K.U. No. 58853, 5.6 kilometers south of Rayon Mescalapa, Chiapas, Mex- ico. 3. Plectrohyla sagorum, K.U. No. 103164, Granja Lorena, 1.3 kilometers north-northeast of Colomba, Departamento Quetzaltenango, Guatemala. 4. Plec- trohyla quccchi, K.U. 64107, Finca Los Alpes, De- partamento Alta Verapaz, Guatemala. PLATE 69. 1 and 2. Plectrohyla glandutosa, K.U. 58703, - o o _) I - 3- ■■"■"-' ■;. , ,-'-i-!ati^ -, r-rii-:.i:i.A".— — I — I — I I 02 0.4 0.6 0.8 TIME IN SECONDS SECTION 1. Hyla regilla curta. 2. Hijla cadaverina. 3. Hyh arenicolor. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 13 4- l^-^ O 4 o to a. CO 2 o S I o .-a5jc*s;*r$s»;i , £ * Sif II ^ 5 ^5 5 3£ 2- ;H|ipi>»iilta)ii*nilffli»^ ^.a:ei. 06 T 1.0 02 04 06 0.8 TIME IN SECONDS SECTION 1. Hyla euphorbiacea. 2. Hijla walkeri. 3. Hijla plicata. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 15 *■- r 3- 2- hfc H Jhu- .^-^'^.*A..4»«v««-~^-,...<...^».^;.^....-, ....■.....■, »^-:^...>»->-,.^^,^ ^:..^.>,..^^.>> O O \ii (n a: 3 2- o >- o o ,. ,'IL* ^^A^,j^ »^o.»4WW^v.w^^^^^^.\^^VMaai>aafej^->«.^'>w>^ 3- 2- 'A- .-^•»-x..?e>- iUui^^^^MMkikM^^^Mk^a^ ■^^>'^- i2J^ 0.2 04 06 08 TIME IN SECONDS SECTION 1. Hyla miotympanum. 2. Hyla arborescandens. 3. Hyla erythromma. NO. 1, PLATE 16 MONOGRAPH MUS. NAT. HIST. 3- O 4 o LlI Q. CO 2 UJ _l o o I o \-^:^^T^J^^,S^:^>rS' ^- :^- -j a"""';! n'>. '% 6- 4- ^,r 1»5 -j*^^jv^^-_ _^i;.^=^ ^-t 2 J^.?*^ :^^-i>>^Ae^r>fc«3.a*^=-^:-iirg-?^t^|r^ -^ ^J^. -^-?^^Ma^ 0 1 0.2 0.3 0 4 TIME IN SECONDS 05 SECTION 1. Hyla thorectes. 2. Hylu hazelae. 3. Hylu loquax. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 17 V i t V i i k i » I O O 6- ir lij CO 4 o o 8 - 2- -.r-^..<^--\ — •■— ,-^^.*^— ,....^Jri-<-. ^•-tj-,--i-l --..i^vJ 0.1 02 03 04 TIME IN SECONDS 05 SECTION 1. Hxjla godvaani. 2. Hijla melanomma melanomma. 3. Hyla melanomma bivocata. NO. 1, PLATE 18 MONOGRAPH MUS. NAT. HIST. 4- ilBij^jjBji^dtj^tliMAlId ,irs3s^ O o llJ w 01 u a. o V o o ^^»e£?5r n3^ I - ^aSa. ..-^— ,-i^^.,rJ;--.*-— .,-1 -■■-.^- . -,-,j*in. -.--,, 3- ;k.; Hif I- :.;^i:^^3^S454fi^, i>». .,-„„-. ■.„^.,^.,^^,^^ Y-.- «K.-->.-^ Min. ^-^^'^^ k.-'-^>.*-»^->.^^^-,'>t^f.^-,^-^- 1.0 0.2 04 06 08 TIME IN SECONDS SECTION 1. Hyla bromeliacia. 2. Hyla sumichrasti. 3. Hyla chaneque. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 19 12 10 8 6 4 2 12- o cn 10 a: LlI Q- 8 (/5 e o O o , , t 1 . I » t 1 • • < ' f ' ' * » * ' t ( ' ^ ^ ■> ■> 11^ ■ ' - - " ' • ^ A ■ . -^ -^ ^. .-» . i t V I I- \ » ( » » s > t". i^=>*-*^ .^:..^,|i4,:^:fe^,^»r#f-^. 12 10 8 6 4 2 *=^ •»^ !■*'■ —I 1 1 — 01 0.2 0.3 TIME IN SECONDS SECTION 1. Hijla picta. 2. Hyla smithii. 3. Hijh rivularis. NO. 1, PLATE 20 MONOGRAPH MUS. NAT. HIST. 3- I- j»ii.>»->irtyJife>ato*j- o o 4- -iiii b^ JJr?* •i^-j.^ii^'^ >,^^jf\^--i-i.i>^.-.'^ .^ iT-1-.iTi I ^m 0.4 0.6 0.8 TIME IN SECONDS SECTION 1. Hyla pseudopuma pseudopuma. 2. Hyla angustilineata. 3. Htjla boons. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 21 '.# >-si ►*t t*r o o Ixl CO 6 or. CL CO 4 LlJ _J O ti 2 o |->-^^'>, 2- F^ F^ 02 TIME 1 1 — 03 04 IN SECONDS 0.5 SECTION 1. Hijla tica. 2. Hyla debilis. 3. Hyla uranochroa. NO. 1, PLATE 22 MONOGRAPH MUS. NAT. HIST. 2- o o 3- 2- o > o o t v1 -1 4- 2- --^1^- -I. 1^ -.I-. ^.1^12., j.^-^^^^-^:^ 0.2 0.4 06 0.8 TIME IN SECONDS 1.0 SECTION 1. Hyla rufioculis. 2. Hijla salvadorensis. 3. Hyla leglen. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 23 2- O o UJ q: a. •>^ 4- o o o 10^ 9 . ^ ^m^: ^^^r V^ji&^^. - ft:.J^^>^naoi^fefei»-^.:taJ^-^-v. ^-ik-^Tii^^tj 4- 3- 2- tVjVi^-*^ - -i^ir^i^^ .; v>p ^^^ . 0 ^ : i^' ^fi •^r'T. '■■u^^-.i' ■"'?^'"rr!'.r7t"^,iji 01 0.2 03 0.4 0.5 TIME IN SECONDS SECTION 1. Hyjla pictipes. 2. Hyla colymha. 3. Hyla rufitela. NO. 1, PLATE 24 MONOGRAPH MUS. NAT. HIST. o 8 o LU 0. 6- (/] 4 > o 2 O v*. I- > -Hvi...l^|>^^^y^^^i^^;^^.;«;;^^;-•':'|>>f 0.2 0.3 0.4 TIME IN SECONDS 0.5 SECTION 1 and 2. Hyla lancasteri. 3. Anotheca spinosa. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 25 O o UJ ^ m^^--^ ^^^^w^ I liiT mil m \m1itt - - .jr ■ -r^- 0.4 06 0.8 TIME IN SECONDS SECTION 1. Hyla crepitans. 2. Hyla crepitans X rosenhergi. 3. Hij]a rosenbergi. NO. 1, PLATE 26 MONOGRAPH MUS. NAT. HIST. au nm if i '>ffft '^5^t o o LU '^ 3 (r ui Q. CO 2 _l o s I o 'ss^^f -±t!>i. :--- : :^ -7-;. -«^- .^---I.- ^..^.- T^^^^rPz- — 1 — \ 1 -—t— 0.2 0.4 06 08 TIME IN SECONDS 1.0 SECTION 1. Uyla elaeochroa. 2. Hyla staufferi staufferi. 3. Hyla staufferi altae. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 27 Ll) CO LlI Q. O >- o .5^ 4- 3- 2- — \ 1 — r 1 — 0.2 04 0.6 08 TIME IN SECONDS 1.0 SECTION 1. Hyh. rubra. 2. Hyla houlengeri. 3. Hyla rostrata. NO. 1, PLATE 28 MONOGRAPH MUS. NAT. HIST. 3% ^^ 0^ Si -^■r:'-'^^' i 8 o o UJ ■^ 6 a: UJ CL o ^ 2 t; "i Y ^- ^ i ^ o o lif * # r-l .;ar;;^-:;-a-:xr-;--;-:T^ ' 8- 4- 3 :ft;?^' # Ji^ jSk: — r" 01 0 2 0 3 0 4 0 5 TIME IN SECONDS SECTION 1. Hyla microcephala microcephala. 2. Hyla phlehodes. 3. Hijla robertmertensi. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 29 4- o 8 o lU - 82 * *- i^ . i. ,v S » :'? 4^ %, •U. '^ !;x *# ' f ' f^ _:» ' — ■* •'' - -^ 0.1 0.2 0.3 0.4 0.5 TIME IN SECONDS SECTION 1. Hj/Za ehraccata. 2. Hyla subocularis. 3. Hyla sartori. NO. 1, PLATE 30 MONOGRAPH MUS. NAT. HIST. 6 tf! ^ *"' ■^- a-r.i^- .^•■?. -^^'^ o o LlI 6 (T UJ CL o o '- o mm- -■■^^f "'^^-^ 2 \D=- *• «&•- - 1^- ih Ji- T- T 0.1 0.2 0.3 04 TIME IN SECONDS 05 06 07 SECTION 1. Ptychohyla schmidtorum cliamulae. 2. Ptijchohyla ignicolor. 3. Agalychnis callidryas (rain call). MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 31 4- : I cc tt SIi^««iSj=;5yl^*«t5^?sS5*te*i<«Tj• 0 6-\ o UJ (/) UJ a. en /, o §2 .>v?iK '3?^,:^ -^iS^^^^^sfeas^&a^^gJ 01 02 03 04 05 TIME IN SECONDS 1. Ptychohyla euthysanota macrotympanum. 2. Ptycholiyla leonhardschtdtzei. 3. Ptychohijla spinipollex. NO. 1, PLATE 32 MONOGRAPH MUS. NAT. HIST. o o u CO a w 4 o >- o 2 O 6- 0.1 02 03 TIME IN SECONDS SECTION 1. Smilisca haudinii. 2. Smilisca cijanosticta. 3. Smilisca phaeota. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 33 a z o o LlI t/) 8- 6- ir UJ to 4 o >- o 2 o ^^ :,i^ss^^(fe*N^ »^i$|^r'^-..?^S^^-^-^ "-^-''*^"-^- -'--^»' ya^ 0.1 0 2 0 3 0 4 TIME IN SECONDS 0.5 SECTION 1. Smilisca puma. 2. Smilisca sila. 3. Smilisca sordida. NO. 1, PLATE 34 MONOGRAPH MUS. NAT. HIST. Q 4 z O o (n 3 cr LlI a. 2 o o ittUUttJIU >,,>' .Jj'j c- V ■ L . ^ ■- ^ 0.2 0.3 0.4 TIME IN SECONDS 0.5 SECTION 1. Pternohyla fodiens. 2. Triprion spatulatus reticulatus. 3. Triprion petasatus. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 35 64^ m 2- ^^•■i^-^: o 8 o a. 6H Q. CO 4 _l O 5 2 o sg ????iM -^°^*^^"~''^" '■''^- ■'^ -^r^ -^y^ ' .-^"^^ ^ r- OUi! 3- 2- Mt 01 02 03 0.4 TIME IN SECONDS -m 0.5 SECTION 1. Acris crepitans. 2. Phyllomedusa lemur. 3. Plectrohyla ixil. NO. 1, PLATE 36 MONOGRAPH MUS. NAT. HIST. o o 4- "^^ vy*"- 3- iLl a. o ^ I o 3- 0.2 0.4 0.6 0.8 TIME IN SECONDS SECTION 1. Gastrot}ieca ceratophrtjs. 2. Gastrotheca nicefori. 3. Phnjnohyas venulosa. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 37 :m I- ■■'iift i tBjiii;- , _ ^ », . . . ■ ■ .-.-.^^j ■-,-■ ...r,. . ^ I Til nil- fr 1 !■ o O Ul ^T>t> T ■r 0.1 0.2 03 0.4 TIME IN SECONDS 0.5 SECTION 1. Agalychnis annae. 2. Agalychnis spurrelli. 3. Agalychnis litodryas. NO. 1, PLATE 40 MONOGRAPH MUS. NAT. HIST. 8- 4 2- ( I )i •>> ^ ^ t. ? f •' f t I o o >- 2 .^-wr-jj^'-a-> .a-^- iJcgW-v^>- ^\w>ff;^ 07 0.2 0.3 04 0.5 TIME IN SECONDS 06 1. Pachymedusa dacnicolor. 2. Smilisca baudinii. 3. PternohyJa fodiens (all release calls) MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 41 1. Pachymedusa dacnicolor. 2. Phyllomedusa venusta. X 1-5. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 42 IP^ 1. Agaltjchnis saltator (night). 2. Agahjchnis calUdryas. ^. Agahjchnis calcarifer. 4. Agalychnis saltator (day). 5. Agalychnis calUdryas. X 1-5. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 43 ^ «> w 1. Agahjchnis moreletii. 2. PhtjUomedusa lemur (night). .3. Agalychnis annae. 4. Agaltjchnis litodryas. 5. PJiijllomedusa lemur (day). 6. Agalychnis spurrelli. X 1- MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 44 1. Hyla miliaria. 2. Hemiphractus panamensis. 3. Anotheca spinosa. X 1-5- MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 45 ,,,^..««s«^^-* .*s»r«***rsr?-'--" 1. Gastrotheca nicefori. 2. Gastrotheca ceratophrys. X 15. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 46 1-4. Phrynohyas venulosa. X !■ MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 47 1, 2. Hijla staufferi staufferi. 3. Hyla staufferi altae. 4. Hyla rubra. 5, 6. Hyla elaeochroa. X 2. MONOGRAPH MUS. NAT. HIST. NO. 1. PLATE 48 1, 2. Hi/la ])ouleng,eri. 3. Hyla rostrata. X 2. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 49 1. Hijla microcephala micwcephala. 2. Hijla niicroceplwla imdcrwoodi. 3. ihjla rohertmertensi. 4. Hyla phlebodes. 5. HyJa sartori. 6. Htjla stihociilaris. 7, 8. Hyla ehraccata. X 2. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 50 1. Hijla rufiteJa. 2. HyJa crepitans. 3. Hyla hoans juvenile. 4. Hyla rosenbergi. X 1-5- MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 51 1, 2. Hyla boans. X 1- MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 52 1. Hyla picadoi. 2. Hyla colyrnba. 3. Hyla angttstilineata, adult. 4. Hyla angustilineata, juvenile. 5. Hyla pseudopuma pseudopuma. 6. Hyla pseudopuma infucata. X 2. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 53 \ > >^' 4^ 1. Hyla tica. 2. Hyla rivularis. 3. Hyla debilis. 4. Hyla xanthosticta. 5. Hyla pictipes, $ 6. Hyla pictipes, 2 . X 2. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 54 1. Hijla rufiocuUs. 2. HyJa lancasteri, lowlands. 3. Htjia uranochroa. 4. Hijla lancasteri, Cerro Pando. 5. Hijla legleri. X 2. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 55 "»i>-5f 1, 2. Hyla smithii. .3. Hijla godmani. 4. Hijla picta. 5. Htjla loquax. X 2. MONOGRAPH MUS. NAT. HIST. NO. 1. PLATE 56 ^■:%-.. 1-4. Htjla miotijmpanum. o. Hijla erythromma. X 2. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 57 1. Hyla hazelae. 2. HyJa thorectes. 3, 4. Htjla arborescandens. 5. Hyla valancifer, juvenile. X 2. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 58 1. Hyla melanomma mehnomma. 2. Hijla melanomma hivocata. 3, 4. Hyla pinorum. 5. Hyla mixomaculata. 6. Hyla pellita. 7. Hyla pinorum. juvenile. 8. Hyla nubicola. X 2. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 59 r 1, 2. //y/a sumichrasti. 3. //y/a smaragdina. 4. //y/o salvadorensis. 5. Hy/a bromeUacia. 6. Hy/a dendroscarta. X 2. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 60 1. Hyla altipotens. 2. Htjla chaneque, i . 3. Hyla chaneqiie, 2 . X 1-5. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 61 1. Hyla taeniopus, $ . 2. Hyla taeniopiis, 9 . 3. Htjia altipotens. X l-o. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 62 ''^^ .j»- 1, 2. Hijla histincta. 3, 4. Hijla pentheter. X 1-5- MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 63 1. Hijla charadricola. 2. Hyla chryses. 3. Hyla robertsorum. 4. Hyla siopela, juvenile. 5. Hyla siopela, £ . X 1.5- MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 64 .--^ffla^ H 1. Hyla cadaverina. 2, 3. Hyla arenicolor. 4. Pseudacris clarkii. 5. Acris crepitans. X 2. MONOGRAPH MUS. NAT. HIST. NO. 1, PLATE 65 S...-*^^ |^