University of Kansas Publications MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly pub- lished in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Publica- tions, Museum of Natural History, began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970. Institutional libraries inetrested in exchanging publications may obtain the Occasional Papers and Miscellaneous Publica- tions by addressing the Exchange Librarian, University of Kan- sas Library, Lawrence, Kansas 66044. Individuals may pur- chase separate numbers of all series. Prices may be obtained upon request addressed to Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66044. Editor: Linda Trueb Managing, Editor: William E. Duellman PRINTED BY UNIVERSITY OF KANSAS PRINTING SERVICE LAWRENCE, KANSAS MUS. COMP. ZOOL. OCCASIONAL PAPERS LIBRARY of the 3^161972 MUSEUM OF NATURAL HISTORY The University of Kansas ^j^f^UYj Lawrence, Kansas NUMBER 11, PAGES 1-31 DECEMBER 6, 1972 A REVIEW OF THE NEOTROPICAL FROGS OF THE HYLA BOGOTENSIS GROUP By William E. Duellman^ By comparison with the highlands of Central America, which are inhabited by a diversity of stream-breeding hylid frogs, the Andes in northwestern South America are characterized by a paucity of stream-breeding hylids. Duellman (1970:327) pointed out that one stream-breeder, Hyla colijmba, in the highlands of Costa Rica and Panama seemed to be related to the Colombian Hyla bogotensis, and he tentatively referred specimens from Ecuador to Hyla colymba. Recent field work in northwestern South America has resulted in the accumulation of series of specimens from many localities, tadpoles, and recordings of mating calls. Thus, it is now possible to review systematically six species that seem to form a natural unit — the Hyla bogotensis group. The purposes of the present paper are to: 1) define the Hyla bogotensis group, 2) present the results of a multivariate analysis of characters, 3) diagnose the species, and 4) present the accumulated data on the biology of the species. ACKNOWLEDGMENTS For the loan of specimens or for provision of working space in their respective institutions, I am indebted to (abbreviations of collections given in parentheses; KU=Museum of Natural History, University of Kansas): Josef Eiselt, Naturhistorisches Museum, Wien (NHMW); Alice G. C. Grandison, British Museum (Natural History) (BMNH); Birgitta Hansson, Naturhistoriska Museet, Goteborg (NHMG); Alan E. Leviton, California Academy of Sci- ^ Curator, Division of Herpetology, Museum of Natural History, University of Kansas. 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY ences (former Stanford University collection) (CAS-SU); Hymen Marx, Field Museum of Natural History (FMNH), Eustorgio Mendez, Gorgas Memorial Laboratory, Panama (GML); Charles W. Myers, American Museum of Natural History (AMNH); Giinther Peters, Zoologisches Museum, Berlin (ZMB); James A. Peters, United States National Museum (USNM); Juan A. Rivero, University of Puerto Rico, Mayagiiez ( UPR-M ) ; Gharles F. Walker, University of Michigan Museum of Zoology (UMMZ); Ernest E. Williams, Museum of Comparative Zoology (MCZ); and John W. Wright, Los Angeles County Museum ( LACM ) . I am grateful to my field associates from the University of Kansas who collected much of the material; thus, I thank Joseph T. Collins, Stephen R. Edwards, John D. Lynch, Richard R. Montanucci, John E. Simmons, and Linda Trueb. Bruce MacBryde of the Universidad Catolica in Quito generously provided trans- portation and field companionship, and the Texaco Petroleum Com- pany of Ecuador provided helicopter flights to otherwise inacces- sible places. Computations were carried out at the University of Kansas Computation Center, and I thank Stephen R. Edwards for assistance in programming. MATERIALS AND METHODS This investigation was based on the study of 149 preserved frogs (including the type specimens of all nominal taxa), 17 lots of tadpoles, two skeletons, nine radiographs, and eight tape record- ings. For purposes of statistical analyses, 14 characters were re- corded for 114 adult frogs. Of these characters, three are mor- phometric, eight are structural, and three are coloration. These data were subjected to a stepwise discriminant analysis by use of the BMD07M Program (Dixon, 1971), which perfomis a multiple discriminant analysis in a stepwise manner, entering one variable at a time into a set of discriminating variables. The program com- putes canonical correlations and coefficients for canonical variables and plots the first two canonical variables in a two-dimensional matrix. Individuals were grouped on the basis of geography and sex, as follows: Central America 6$, 3$; Pacific slopes of Ecuador 13 S , 15 9 ; Amazonian slopes of Ecuador and Peru 13 ,5 , 3 $ ; Andes of central and southern Colombia 7^,4$; Andes of western Venezuela 37 5 , 11 9 . One male and one female from northern Colombia were each treated as a group. Thus, the matrix for the discriminant analysis consisted of 14 variables and 12 groups. Subsequently, the sexes were pooled, and each of the six groups was treated as an OTU in the NT-SYS Phenogram Program ( Rohlf and Kishpaugh, 1966); in this program unweighted arithmetic character states were analyzed and plotted as a distance phenogram. For purposes of the Phenogram Program, 11 coded characters were FROGS OF THE HYLA BOGOTENSIS GROUP 3 used (see following definition of characters). Five coded char- acters of tadpoles were analyzed by the Phenogram Program; again, groups (taxa) were treated as OTUs. Finally, the adult and tadpole characters were analyzed together by the Phenogram Pro- gram. All computations were done on a Honeywell 635 computer. Recordings of the mating calls of three species are available. The calls of eight individuals were analyzed for seven characters: 1 ) notes per call group, 2 ) note repetition rate, 3 ) duration, 4 ) pulse rate, 5) number of harmonics, 6) fundamental frequency, and 7) dominant frequency (see Duellman, 1970, for methodology and terminology ) . Definition of Characters In the list of characters, those marked with an asterisk were used in the Phenogram Program; all characters of adults were used in the BMD07M Program. Snout-vent Length (SVL)'*. — Measurement to nearest 0.1 mm of straight line distance from tip of snout to posterior edge of body; for NT-SYS coded: 1) 6 <44 mm, 9 <50 mm, 2) S >44 mm, 9 > 50 mm. Tibia Length (TL). — Measurement of length of tibia to nearest 0.1 mm; used as a ratio of snout-vent length. Head Width (HW). — Measurement of greatest width to nearest 0.1 mm; used as a ratio of snout-vent length. Webbing on Hand (WEB)''. — The extent of webbing on the inner edge of the fourth finger coded with respect to the point of termination of the webbing with respect to the distal subarticular tubercle: 1) proximal to tubercle, 2) at tubercle, 3) distal to tubercle. Subartictdar Tubercles (TUB)''.— The structure of the distal subarticular tubercles on the third and fourth fingers was coded: 1) conical, single, 2) conical, bifid, 3) flat, bifid. Uhiar Fold (ULN)''. — A dermal fold on the ventrolateral edge of the forearm was coded: 1) absent, 2) present. Tarsal Fold (TAR)*'. — A dermal fold on the ventrolateral sur- face of the foot was coded : 1 ) absent, 2 ) present. Calcar (CAL)**. — A triangular dermal appendage projecting posteriorly from the upper edge of the heel was coded: 1) absent, 2) small, 3) large. Snout Profile (SNO)*. — The shape of the snout in lateral view coded: 1) round, 2) truncate, 3) anteriorly inclined. Tympanum (TYM)'*. — The upper edge of the tympanum is covered by a supratympanic fold, but the tympanic ring below the fold coded: 1) distinct, 2) covered with undifferentiated skin. Mental Gland (MEN). — A round glandular structure at the apex of the jaw was coded: 1 ) absent, 2) present. 4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Dorsolateral Stripe (DST)*.— A pale stripe extending from the edge of the eyehd to a point on the side of the body was coded: 1) absent, 2) present. Tarsal Stripe (TST)*.— Longitudinal stripes on the outer edge of the foot were coded: 1) absent, 2) white, 3) white above and brown below. Anal Stripe ( AST )* .—Transverse stripes above the anus were coded: 1) absent, 2) white, 3) white above and brown below. Tooth Rows (TOO)*.— The number of upper/lower rows of teeth in tadpoles was coded: 1) 4/5, 2) 5/7, 3) 6/9, 4) 7-8/10. Labial Papillae (LAB)'*. — Labial papillae are continuous around the mouth in the tadpoles; the number of rows of papillae was coded: 1) one, 2) two. Serrations on Beak (S£R)*.— The serrations on the cutting edge of the upper beak in tadpoles were coded: 1) none, 2) small, 3) medium, 4) large. Shape of Body (SHAj'.— The shape of the body of tadpoles in dorsal view was coded: 1) narrowly ovoid, 2) broadly ovoid, 3) narrow anteriorly and widening abruptly just anterior to eyes. Caudal Muscidattire (CAU)".— The proportionate depth of the caudal musculature was coded: 1) <65%, 2) >i RESULTS For ease in discussion the nomenclature adopted in the follow- ing section on taxonomy is used throughout. The results of the discriminant analysis are discussed first, followed by the Phenogram Program and finally the mating calls. Multiple Discriminant Analysis Within-group Variation. — The amount of variation in snout-vent length is approximately the same in all species except H. bogotensis and H. platydactyla in which it is somewhat greater (Table 1). However, the amount of variation in the ratios of tibia length and head width to snout-vent length is no greater in these two species than in the others. Variation of all structural features, except presence or absence of ulnar and tarsal folds, occurs within groups; likewise, color pattern characters show within-group variation (Table 2). Tarsal and anal stripes are either present or absent within a group; if present, there may be one or two stripes (Hyla phyllognatha). The presence of a mental gland is the most variable structural feature, displaying variation in five of the 12 groups. The amount of webbing is the next most variable character; variation occurs in four groups. The variation within groups is not highly correlated with respect to the following characters — size, proportions, subarticular tubercles, snout shape, tympanum, and mental gland. Absolute positive cor- FROGS OF THE HYLA BOGOTENSIS GROUP Table 1. — Measurements and proportions of species in the Hyla hogotensis group. ( Mean and one standard deviation given below observed range. ) Sp ecies Sex N Snout-vent Length Tibia Length/ SVL Head Width/ SVL H. ahjtohjiax S 13 32.1-37.0 34. 85 ±1.51 0.443-0..525 0.474±0.024 0.298-0.331 0.316±0.010 9 15 37.2-43.9 40.38+2.23 0.462-0.532 0.496i!i0.021 0.296-0.331 0.310±0.010 H. hogotensis $ 7 29.4-43.0 35.20±5.58 0.491-0.514 0.502±0.010 0..329-0.365 0.345±0.012 9 4 29.4-48.1 41.43 0.449-0.506 0.483 0.313-0.380 0.347 H. cohjmba S 6 31.9-37.0 34.65±2.29 0.452-0.514 0.481±0.023 0..302-0.327 0.318±0.010 9 3 31.4-39.1 36.23 0.486-0.493 0.490 0.310-0..340 0.325 H. denticulenta 6 1 44.2 0.493 0..303 9 1 52.2 0.513 0.308 H. phyllognatha S 13 31.5-34.0 32.96+0.69 0.443-0.524 0.488±0.028 0.301-0. .348 0.319±0.013 9 3 33.0-39.3 36.87 0.488-0.506 0.495 0.326-0.331 0.329 H. platydactyla S 37 26.5-39.4 34.36±3.05 0.464-0..540 0.488±0.016 0.323-0.383 0.347±0.014 9 11 30.6-42.3 38.41±3.64 0.457-0.506 0.491±0.016 0.3.33-0.356 0.342±0.009 relation exists between presence or absence of ulnar and tarsal folds, between tarsal folds and tarsal stripes, and between tarsal and anal stripes. Webbing is the most extensive and calcars are the largest in the species, H. denticulenta. Sexual Dimorphism. — No sexual dimorphism is evident in pro- portions, amount of webbing, nature of folds or calcars, snout shape, or nature of tarsal and anal stripes. Minor sexual dimorphism occurs in the nature of the subarticular tubercles in H. platydactyla, in the tympanum in H. hogotensis, and in the dorsolateral stripe in H. colymba (Table 2). Snout- vent length is greater in females than in males; snout- vent lengths of 77 males is 26.5-44.2 (x=34.43) mm, and of 37 females is 29.4-52.2 (x=39.63) mm. As noted above, the nature of the mental gland is highly variable. The gland is present in 44 of 77 (57.1%) males and in nine of 37 (24.3%) females. The structure and function of the mental gland are unknown; its development ma>' be seasonal and associated with reproductive activity. Thus, its usefulness as a taxonomic character within the Hyla hogotensis group is questionable. Interpopulational Variation. — The combination of the sexes in 6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY C o CO c o a i-i a: > o -a o 13 01 .S c G O 'o o 13 c «2 D c o c C bc •S T3 O CJ n3 c C8 en fl O 2 -2 > J3 « o O U o a Q C/2 c^ CO t- CO in ^inoqc!,co en ^ ■' "•' V CO in J-. CO I— I a o (M 1—1 ^ ^ (c in ^ d ^ I ro <^ °0 I "^ •> ^ ' rH CO ^ (M (M I— I in 1 t- CO fvi ]^ in in ^' .-i d -H i — I c 3> a: o FROGS OF THE HYLA BOGOTENSIS GROUP 7 the geographic samples provides an assessment of populational dif- ferences for taxonomic distinction. Each of the 14 characters, in combination with others, serves to distinguish one population from another, but, with the exception of snout-vent length in H. denticulenta, no one character distinguishes one population from all of the others. The stepwise discriminant analysis provided a multivariate analysis of 14 characters in the 12 groups (males and females were treated separately in each of the geographic samples) and produced a two-dimensional plot of the first and second canonical variables ( Fig. 1 ) . The first canonical variable is weighted primarily on webbing, secondly on subarticular tubercles, and lastly on ulnar and tarsal folds, whereas the second canonical variable is weighted on snout-vent length, calcar, and dorsolateral stripe, in descending order. The plot clearly shows the clustering of sam- ples and the overlap ( in most cases ) of the sexes within the samples. Thus, on the basis of adult morphology, the samples can be treated as different taxa. Distance Phenograms In an attempt to determine the phenetic relationships of the six geographic samples, each of the six was treated as an OTU. Mean values for each of 11 characters (sexes pooled) were used. Body ratios and the nature of the mental gland were excluded from the analysis. The resultant distance phenogram (Fig. 2A) shows the separation of the phenetically similar H. bogotensis and H. platydactyla from the other taxa. Hyla denticulenta is separated from the remaining three species clumped in the middle of the phenogram. The arrangement of the taxa is compatible with the plot of the canonical variables (Fig. 1). Thus, the results of the Phenogram Program supports the results of the multiple discrim- inant analysis. The previous analyses were based on characters of the adult frogs. Tadpoles are available for all six geographic samples; these have been associated with adults on the basis of morphological char- acteristics of the metamorphosing young. The most obvious inter- populational differences are evident in the structures of the mouths (Fig. 3). The distribution of character states of five characters is shown in table 3. For purposes of analysis, the mean ratio of depth of caudal musculature to total caudal depth was used; the other characters are constant within populations and were coded. The phenogram of populations as OTUs differs only slighdy from the arrangement based on adults ( Fig. 2B ) . The major difference is that H. ahjtohjlax is clustered with H. denticulenta, whereas in the phenogram based on adults H. ahjtohjlax is grouped with H, cohjmba and H. phyllognatha. OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fig. 1. — Plot of first (horizontal axis) and second (vertical axis) canonical variables; solid dots are males and open circles are females. A. Hijla ahjtohjlax. B. H. cohjmha. C. H. phyllognatha. D. H. denticulenta. E. H. bogotensis. F. H. platydactyla. A final analysis of both adult and larval characters produced a phenogram somewhat intermediate between the first two (Fig. 2C). Here it can be seen that H. bogotensis and H. platydactyla are divergent from the other taxa and that H. colymba and H. phyllognatha are close to H. ahjtohjlax, whereas H. denticulenta is more distant. FROGS OF THE HYLA BOGOTENSIS GROUP 9 -i — I — I — I — I — I — r—i I I I I 1 r H. denticulenta H. alytolylax ' H. colymba H. phyllognatha H. platydactyla H. bogotensis B — H. denticulenta — H. alytolylax — H. colymba ^ — H. phyllognatha — H. platydactyla — H. bogotensis 3.0 2.0 1.0 DISTANCE H. denticulenta H. alytolylax H. colymba H. phyllognatha H. platydactyla H. bogotensis I I I I I I I I I I I I I I O.O Fig. 2. — Distance phenograni of Hyla bogotensis group based on un- weighted analysis of 11 adult and five larval characters. A. Adult characters. B. Larval characters. C. Adult and larval characters. 10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY ',",«."""'"•.„>..,„ ^/^"■:: -"n::-^^^.^ j§;^;:;;:;o ;; "-"'<^-k^. / / '':!T"""^«,„, Fig. 3. — Mouths of tadpoles of Htjia bogotensis group. A. H. phtijdactyla, KU 139521. B. H. ahjtohjlax, KU 112358. C. H. phyUognatha, KU 143542. D. H. denticulenta, KU 139531. X8. The mouth of the tadpole of H. bogotensis is like that of H. platijdactyla except that the former has much smaller serrations on the beaks; the mouth of the tadpole of H. colymba is like that of H. phyUognatha (see Duellman, 1970:330). Mating Calls The mating calls are known for all species, except H. bogotensis and H. denticulenta; the calls consist of a series of short, loud peeps. Analysis of tape recordings of calls of three species reveals notice- able differences in several parameters of the calls ( Table 4, Fig. 4 ) . Hyla colymba has the most distinctive call, differing from the others in notes per call group, duration of notes, pulse rate, and pitch; the second harmonic is dominant. The calls of H. phyUognatha and H. platydactyla are more nearly alike in having only one har- Table 3. — Characteristics of tadpoles of Hyla bogotensis group. Total Caudal Tooth Labial Serrations Length/ Musculature/ Species Rows Papillae on Beak Body length Caudal Depth H. alytolylax 5/7 2 small 30.2% 63.1% H. bogotensis 4/5 1 medium 32.3% 66.7% H. colymba 6/9 2 small 35.6% 61.1% H. denticulenta 7-8/10 2 none 35.3% 57.2% H. phyUognatha 6/9 2 small 35.2% 60.0% H. platydactyla 4/5 1 large 39.5% 80.0% FROGS OF THE HYLA BOGOTENSIS GROUP 11 monic and fewer, but longer, notes per call group. However, their calls differ in note repetition rate, pulse rate, and pitch. The ab- sence of analyzable recordings of the other three species precludes a complete biosonic analysis of the group. Nevertheless, the dif- ferences in the parameters of the calls of the three sepcies are of the magnitude separating both sympatric and allopatric species in other Neotropical hylids [see Duellman (1963), Duellman and Trueb (1966), Duellman and Fouquette (1968), and Duellman ( 1972 ) for examples] . Table 4. — Comparison of mating calls of species in the Hijla bogotensis group. Character iV" H. cohjmba 3/9 H. phijUognatha 4/13 H. platydactiila 1/16 Notes per Call Group 12-104 (40.3) 2-12 (5.5) 7-10 (8.8) Note Repetition Rate ( min ) 123-236 (179) 120-360 (210) 540 Duration (sec) 0.05 0.11-0.13 (0.12) 0.09-0.17 (0.14) Pulses per Second 120-160 (138) 260-280 (279) 170-180 (177) Hamionics 4 1 1 Fundamental Frequency ( Htz ) 1760-1820 (1796) 2550-2620 (2588) 2700-2860 (2790) Dominant Frequency ( Htz ) 3520-3640 (3592) 2550-2620 (2588) 2709-2860 (2790) " 2V=individuals/notes. DISCUSSION The grouping of specimens into geographic samples provided an a priori assumption that the groups represented different taxa. This assumption was tested by discriminant analysis of 14 charac- ters in adults. The results were re-tested by subsequent independent and combined analyses of 11 characters of adults and five of tad- poles by means of the distance phenogram program. The conclu- sions are supported by data on mating calls. Although it is possible that some of the taxa (H. bogotensis and H. platydactyla; H. colymba and H. ahjtolylax) might be geographical variants (sub- species), there exists no evidence for gene flow between popula- tions. Consequently, the six recognized taxa are treated as species. The distributions of the species are broadly allopatric (Fig. 5). Two species, H. bogotensis and H. platydactyla, occur in the Andes at elevations of 2500-2900 m and 1600-2700 m, respectively. Hyla colymba inhabits both Caribbean and Pacific slopes at elevations of 560-1410 m in Costa Rica and Panama. The other species occur on Andean slopes: H. alytolylax at 800-1460 m on Pacific slopes, 12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 6- 4- 0 6H 7? ^ .- 0^^f^ 2- 0- 8 4- 00 0.2 Time 0 4 in Seconds 0 6 Fig. 4. — Audiospectrograms of mating calls. A. Uyla platydacttjla, KU Tape 1029; 15-20 km NW Merida, Merida, Venezuela; temperature unknown B. H. phyllomatha, KU Tape 1166; Rio Azuela, Napo, Ecuador; 18° C. C. H. colymba, KU Tape 599; ridge between Rio Jaque and Rio Iniamado, Darien, Panama; 19° C. Band width 20 Htz. //. phijllognatha at 610-1740 m on Amazonian slopes, and H. denticiilenta at 1400-2400 m on Caribbean slopes. The principal habitat requirement seems to be cool, rocky streams for development of the tadpoles. Due to lack of substantive information on direction of evolu- tionary change in many of the characters utilized, no quantitative phyletic analysis (Kluge and Farris, 1969) was attempted. The FROGS OF THE HYLA BOGOTENSIS GROUP 13 80 75 70 J 1 I 1 / e> } , ^\^'^^~~\^ 10 10 5 i [^ )yv w 5 0 0 >^ Y ^1 5 \ } 5 0 5 1 1 1 . 1 — \ 00 \ \ ^ Kilometers \ V\ -^^/' , 80 75 70 Fig. 5. — Distributions of the species in the Hj/Zfl bogotensis group. A. H. ahjtolylax. B. H. cohjmha. C. H. phijUognatha. D. H. denticulenta. E. H. bogotensis. F. H. platydactyla. Lines enclose approximate knowii ranges, within which specific h)cahties are indicated by dots; the range of H. phyllognatha extends to southern Peru. presence of dermal folds and calcars in adults and proliferation of tooth rows in tadpoles are certainly derived states (Duellman, 1970 ) . The occurrence of extensive webbing and bifid subarticular 14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY tubercles are probably derived character states. Furthermore, it seems that the presence of two tarsal stripes may have evolved from a condition in which one stripe was present. Other characters of structure and coloration remain evolutionarily enigmatic. All evi- dence points to H. bogotensis and H. platydactyla being the most generalized species, that is, having fewest derived character states. Hyla denticulenta possesses an array of derived states in both larvae and adults and seems to be the most advanced species in the group. The remaining three species seem to be closely related and phylogenetically intermediate between the primitive H. bogotensis and H. platydactyla and the advanced H. denticulenta. Evidence from a variety of sources points to considerable Pleisto- cene climatic fluctuation and concomitant shifts in vegetation in the northern Andes (see Vuilleumier, 1971, for summaiy). This climatic fluctuation, together with orogenic changes in the Andes during the Pleistocene and volcanic activity to the Present, provided ample opportunity for isolation of populations on different slopes of the Andes (H. alytoh/lax, H. denticulenta, and H. phijUognatha) , in d'sjunct interandean basins (//. bogotensis and H. platydactyla), and dispersal across presently uninhabited lowlands (H. colymba). TAXONOMY The inclusion of six species in the Hyla bogotensis group neces- sitates a modification of the definition of the group as given by Duellman (1970:327). The group can now be defined: 1) moder- ate-size:l, stream-breeding frogs with males attaining snout-vent lengths of 45 mm and females 53 mm; 2) dorsum pale green or brown with or without dark flecks; 3) digits bearing small discs; 4) t )es about three-fourths webbed; 5) axillary membrane absent; 6) fjuadratojugal articulating with maxillary; 7) sphenethmoid broad, not ossified anteriorly; 8) nasals small, widely separated medially; 9) frontoparietal fontanelle large; 10) tadpoles having long muscular tails, low fins, ventral mouths completely bordered by papillae, and 4/5 — 8/10 tooth rows; 11) mating calls consisting of series of short, loud peeps. Distribution. — The combined distributions of the six species include the Andes and interandean valleys of western Venezuela and central and southern Colombia, Pacific and Amazonian slopes of Andes in Ecuador, Amazonian slopes in Peru, Caribbean slopes in northern Colombia, and Caribbean and Pacific slopes of high- lands in Costa Rica and Panama, with an elevational range of 560 to 2900 m (Fig. 5). Remarks. — Members of the Hyla bogotensis group superficially resemble another group of species inliabiting lower Andean slopes in Ecuador, Colombia, and Venezuela (H. palmeri Boulenger, H. albopunctata Boulenger, and H. lascinia Rivero). These frogs lack mental glands; the mating calls and tadpoles are unknown. FROGS OF THE HYLA BOGOTENSIS GROUP 15 Key to the Species in the Hyla bogotensis Group 1. Tarsal and ulnar folds and tarsal and anal stripes present; distal subarticular tubercle on fourth finger single, conical (if bifid, flat, not conical) 2 Tarsal and ulnar folds and tarsal and anal stripes absent; distal subarticular tubercle on fourth finger bifid, conical 5 2. Distal subarticular tubercle on fourth finger single, conical; calcar, if present, small; snout round or truncate; tarsal and anal stripes single or double 3 Distal subarticular tubercle on fourth finger bifid, flat; calcar large; snout anteriorly inclined; tarsal and anal stripes double, light above, dark below H. denticulenta 3. Dorsolateral stripe absent; webbing in males not extending to distal subarticular tubercle on fourth finger, reaching tubercle in females H. phyllognatha Dorsolateral stripe usually present; webbing in both sexes extending to or beyond subarticular tubercle on fourth finger 4 4. Dark pigment fine, scattered on dorsum; webbing usually extending beyond distal subarticular tubercle on fourth finger H. alytolylax Dark pigment coarse, dense on dorsum; webbing extending to distal subarticular tubercle on fourth finger H. colymba 5. Webbing extending to distal subarticular tubercle on fourth finger; dorsolateral stripe absent; dark pigment usually fine and scattered on dorsum H. bogotensis Webbing not extending to distal subarticular tubercle on fourth finger; dorsolateral stripe present or absent; dark pigment on dorsum forming discrete spots in adults H. platydactyla Hyla alytolylax new species Figure 6 A Holotype. — KU 111903 from Tandapi, Provincia Pichincha, Ecuador, 1460 m, obtained on 22 July 1967, by John D. Lynch. Paratopotypes.—KU 111892-6, 111898-902, 111906, John D. and Marsha Lynch, 15-24 July 1967; KU 117982-4, John D. Lynch, 1-3 March 1968; KU 120851-60, John D. Lynch and Gerald R. Smith, 27-28 July 1968; KU 132425-6, John D. Lynch, 6 August 1970. Diagnosis. — 1) Webbing extending to or beyond distal subar- ticular tubercle on fourth finger; 2) distal subarticular tubercles on fingers single, conical; 3) ulnar and tarsal folds present; 4) calcar absent in males, absent or small in females; 5) snout in profile round in males, truncate in females; 6) tympanum distinct; 7) 16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fig. 6. — Frogs of the Htjla hogotensis group. A. H. ahjtohjlax, KU 111900, 9, 39.6 mm SVL. B. H. denticulenta, KU 133451, $, 44.2 mm SVL. C. H. phyUognatha, KU 143197, 'pe); pertinent mea- surements and proportions given in table 1. Head as wide as body; snout moderately short, round in dorsal aspect, anteriorly inclined in profile (especially in female); nostrils three-fourths distance from eye to tip of snout, not protuberant; internarial area flat; canthus round; loreal region concave; lips thin, round; top of head flat; eyes not greatly protuberant; supratympanic fold weak, curved downward toward angle of jaw, obscuring upper part of tympa- num; tympanic ring distinct; tympanum one-half diameter of eye, separated from eye by distance equal to half again diameter of tympanum. Axillary membrane absent; forearm robust, bearing ulnar fold prepollical tubercle large, elliptical; palmar tubercle large, bifid distal subarticular tubercle on fourth finger flat, bifid; others round supernumerary tubercles small, indistinct; fingers short, bearing small discs (that on third finger slightly larger than tympanum); length of fingers from shortest to longest 1-2-4-3; webbing vestigial between first and second fingers, extending from base of penultimate phalange of second to distal end of antepenultimate phalange of third, from base of penultimate phalange of third to base (male) or middle ( female ) of penultimate phalange of fourth finger. Hind limb moderately robust; dermal fold on knee; calcar prominent, triangular; inner tarsal fold absent; outer tarsal fold extending length of tarsus; inner metatarsal tubercle elliptical, visible from above; outer tarsal tubercle absent; subarticular and supernumerary tubercles about equal in size, small, round; toes long; length of toes from shortest to longest 1-2-3-5-4; discs slightly smaller than those on fingers; webbing extending to base of penultimate phalange of fourth toe, to discs on other toes. Anal opening directed posteroventrally at midlevel of thighs ( male ) , posteriorly at upper level of thighs ( female ) ; skin on belly and proximal posteroventral surfaces of thighs weakly granular; skin on other surfaces smooth. Tongue broadly cordiform, shallowly notched posteriorly, barely free behind; dentigerous processes of prevomers slightly anteromedially inclined posterior to small ovoid choanae, bearing 21 (male), 20 (female) teeth; vocal slit extending from posterolateral base of tongue to angle of jaw; vocal sac sub- gular, possibly bilobate. Coloration. — In life, the male holotype had a pale green dorsum covered with many small white flecks; black flecks were present 24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY on the head and back. The flanks were creamy yellow; the throat, chest, groin, anterior surfaces of the thighs, and ventral surfaces of the shanks were bluish green. The belly was creamy yellow, and the iris was reddish gold. The female had a green dorsum with a white margin to the lips and white anal, ulnar, and tarsal stripes. The axilla, groin, and ventral surfaces of the thighs were blue-green; the flanks were yellow-orange. The throat and belly were greenish white, and the iris was dark gold with black reticulations. In preservative, all pigment is lost, except for many small brown flecks on the head and anterior two-thirds of the back in the male and a few scattered flecks on the eyelids, head, and middorsum of the female. Also, white flecks are apparent dorsolaterally in the male. In both specimens narrow white anal, ulnar, and tarsal stripes are bordered below by a narrow line of brown pigment, which is expanded on the palm and sole. Tadpoles. — Five tadpoles (KU 139531) are from the type locality. The smallest individual is in developmental stage 25 and has a body length of 17.5 mm and a total length of 51.5 mm; the largest is in stage 38 and has measurements of 22.0 and 65.5 mm. Body ovoid, four-fifths as deep as wide, widening and deepening abruptly just anterior to the eyes; nostrils midway between eyes and tip of snout; spiracle sinistral; spiracular opening directed pos- teriorly just below midline at about midlength of body; cloacal tube dextral; caudal fin not extending onto body; caudal muscula- ture robust, gradually tapering, extending nearly to tip of pointed tail; depth of musculature at one-third length of tail 57 percent of total depth of tail. Mouth large, ventral; lips deeply indented posteriolaterally, completely bordered by two rows of small papillae; seven or eight upper and ten lower rows of teeth; proximal upper row narrowly interrupted medially; upper beak slender with long, tapering lateral processes; lower beak broadly V-shaped; both beaks smooth or bearing minute serrations (Fig. 3D). In preservative, the body is dark brown above and dull gray below; the caudal musculature is cream with dense brown pigment laterally, forming distinct transverse bars, separated by narrow cream interspaces, dorsally. The fins are transparent, except for a few flecks on the dorsal fin. In life, the body is pale gray; the tail is distinctly mottled black, gray, and white, and the fins are transparent. Etymology. — The specific name is derived from the Latin den- ticuhis meaning with small teeth, and the Latin lentus meaning full of; the name refers to the many rows of teeth in the tadpoles. Distribution. — This species is known from only two localities: Charta at an elevation of 2400 m on the northern slope of the Cordillera Central, and Valdivia at an elevation of 1400 m on the northern slope of the Cordillera Central. These localities are sepa- rated by the low Rio Magdalena Valley. It is unlikely that gene FROGS OF THE HYLA BOGOTENSIS GROUP 25 flow exists between the populations; thus, it is possible that addi- tional material will reveal difi^erences between the populations. Remarks. — At Charta the frogs were found along a shallow river south of the village. The river, a tributary of the Rio Surata, flows through pasture land with a few scattered trees. Tadpoles and a metamorphosing young having a snout-vent length of 24.6 mm and a tail stub of 10.4 mm were found in pools in the river bed. The holotype was found by day under a large rock in a small stream cascading into the river bed. The adult female was found in the town of Valdivia, where it was perched on a leaf of a large herb 2 m above the ground on a steep bank at night. The mating call is unknown. Hyla phyllognatha Melin Figure 6C Hyla phyllopnatha Melin, 1941:30 [Holotype.— NHMG 474 from Roque, Departamento San Martin, Peru; Douglas Melin collector]. Diagnosis. — 1) Webbing not extending to, or just extending to, distal subarticular tubercle on fourth finger; 2) distal subarticular tubercles on fingers single, conical; 3) ulnar and tarsal folds present; 4) calcar absent or small; 5) snout in profile round or truncate; 6) tympanum distinct; 7) mental gland present in males, present or absent in females; 8) dorsolateral light stripe absent; 9) tarsal and anal stripes absent or white; 10) tadpoles having 6/9 tooth rows, two rows of labial papillae, and small serrations on beaks. Hijla phyUognatho is like H. cohjmha and H. ahjtohjlax but has less webbing and no dorsolateral stripe. In some specimens of H. phyllognatha a calcar is present, but this is smaller than the calcars in H. denticulenta, which further differs by having bifid subarticular tubercles, anteriorly inclined snout, and more webbing. The tadpoles of H. phyllognatha have 6/9 tooth rows, a character shared with H. cohjmha, the tadpoles of which differ from those of H. phyllognatha by not having transverse brown blotches on the dorsal caudal musculature. Coloration. — In life, the dorsum is olive-green anteriorly chang- ing to olive-tan posteriorly; the dorsal surfaces of the limbs are green. Minute black flecks are present on the head and middorsal part of the body, and small white flecks are scattered on the body and limbs. The margin of the lip is yellow, and the anal and tarsal stripes are pinkish white. The hidden surfaces of the limbs are dull green. The belly is creamy yellow, and the webbing is dull yellow. The vocal sac is green, and the lining of the mouth is bluish green. The iris is dull bronze with black reticulations. One female found by day was bright green with bluish white flecks. In preservative, the dorsum is cream to grayish brown with scattered dark flecks apparent in paler specimens. The venter and hidden surfaces of the thighs are cream. 26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Tadpoles. — A typical tadpole in developmental stage 27 from 16.5 km NNE of Santa Rosa, Ecuador, has a body length of 16.5 mm and a total length of 50.3 mm. Body ovoid, slightly wider than deep; nostrils slightly closer to tip of snout than to eyes; spiracle sinistral; spiracular opening directed posteriorly just below midline about two-thirds length of body; cloacal tube dextral; caudal fin not extending onto body; caudal musculature robust, tapering gradually, extending nearly to tip of pointed tail; depth of musculature at one-third length of tail 60 percent of total depth of tail. Mouth large, lips indented posterolaterally, completely bordered by two rows of small labial papillae; six upper and nine lower rows of teeth; sixth upper row narrowly interrupted medially; beaks moderately slender, bearing small, blunt serrations; upper beak arched with lateral processes barely expanded distally; lower beak broadly V-shaped ( Fig. 3C ) . In preservative, the body is dark brown above and gray below; the caudal musculature is cream with dense brown flecking laterally and dark brown transverse bars anterodorsally. The fins are trans- parent with brown flecks, except anteriorly on ventral fin. In life, the body is dark brown with small green, lichenous flecks. The caudal musculature is tan with brown blotches; the fins are clear with orange spots. The iris is dark bronze. Distributio7i. — Hyla phijllognatha is known from elevations of 610-1740 m on the Amazonian slopes of the Cordillera Oriental of the Andes and associated ranges, such as the Cordillera del Due and Cordillera Carabaya, from northern Ecuador to southern Peru. Remarks. — Males call from low bushes over torrential mountain streams; calling males have been found in April, July, August, and October. Gravid females have been found in October. The tadpoles live in gravel-bottomed pools in the streams. Recently metamor- phosed young have been found on vegetation along streams at night and in bromeliads by day. Four young have snout-vent lengths of 18.4-20.3 (x=19.5) mm; in life, the dorsum is green with black flecks middorsally. The mating call consists of a series of short, loud whistles, repeated at a rate of about 210 notes per minute. The notes have about 280 pulses per second; the fundamental frequency at about 2560 Hertz is dominant ( Fig. 4B ) . No information has been published on Hijla phtjllognatha since Melin's (1941) original description. The holotype (NHMG 474) is an adult male having a snout-vent length of 32.3 mm and a mental gland. The specimen compares favorably with series of fresh specimens from Equador and three individuals from southern and central Peru. FROGS OF THE HYLA BOGOTENSIS GROUP 27 Hyla platydactyla Boulenger Figure 60 Hyla platydactyla Boulenger, 1905:183 [Holotype.— BMNH 1904.6.30.17 (RR 1947.2.13.14) from Merida, Estado Merida, Venezuela; Briceno collector]. Hyla paramica Rivero, 1961:112 [Holotype. — UMMZ 59016 from Escorial, Estado Merida, Venezuela; received from W. F. H. Rosenberg]. New synonymy. Hyla jahni Rivero, 1961:113 [Holotype.— UMMZ 46465 from Escorial, Estado Merida, Venezuela; presented by A. G. Ruthven]. New synonymy. Diagnosis. — 1) Webbing not extending to distal subarticular tubercle on fourth finger; 2) distal subarticular tubercles on fingers single or bifid in males, bifid and conical in females; 3) ulnar and tarsal folds absent; 4) calcar absent; 5) snout in profile truncate; 6) tympanum distinct or covered; 7) mental gland present or absent in both sexes; 8) dorsolateral stripe present or absent; 9) tarsal and anal stripes absent; 10) tadpoles having 4/5 tooth rows, one row of labial papillae, and large serrations on beaks. Hyla platydactyla is like H. hogotensis in having bifid, conical subarticular tubercles, no tarsal or anal stripes, no ulnar or tarsal folds, and tadpoles with 4/5 tooth rows and one row of labial papillae; these characters distinguish these two species from other members of the group. Hyla platydactyla can be distinguished from H. hogotensis by having less webbing and, in large adults, discrete dark spots dorsally. Also, the tadpoles of H. platydactyla have larger serrations on the beaks than do those of H. hogotensis. Coloration. — In life, specimens from a locality 15-20 km NW of Merida, Venezuela, had reddish brown dorsal surfaces with many small black spots on the back. The throat was greenish tan, and the belly was white. The iris was copper with fine black reticulations. Through the courtesy of Juan A. Rivero, I have seen colored photographs of individuals having olive-tan dorsal colora- tion and creamy yellow dorsolateral stripes. Rivero stated (pers. com.) that some individuals are dark brown above and that in most specimens the anterior and posterior surfaces of the thighs are orange. In preservative, in adults the dorsal surfaces of the head, body, forearms, and shanks are tan to brown with small dark brown to black spots present in most specimens. The dorsolateral stripe, thighs, and ventral surfaces are cream. Tadpoles. — In a series from 15-20 km NW of Merida, Venezuela, the smallest tadpole is in developmental stage 25 and has a body length of 8.5 mm and a total length of 26.5 mm; the largest tadpole (stage 37) has measurements of 23.0 and 60.5 mm. Body ovoid, three-fourths as deep as wide; nostrils slightly closer to eyes than to tip of snout; spiracle sinistral; spiracular opening directed pos- teriorly on midline about midlength of body; cloacal tube dextral; caudal fin not extending onto body; caudal musculature robust, 28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY tapering gradually, extending nearly to tip of pointed tail; depth of caudal musculature at one-third length of tail 80 percent of total depth of tail. Mouth moderately large; lips deeply indented postero- laterally, completely bordered by one row of labial papillae; four upper and five lower rows of teeth; fourth upper row narrowly in- terrupted medially; fifth lower row short; upper beak moderately massive, arched, with robust lateral processes and large, pointed serrations; lower beak massive, broadly V-shaped, bearing large blunt serrations (Fig. 3A). In preservative, the body is dark brown above and paler brown below; the caudal musculature is creamy tan with a heavy suffusion of brown laterally and two or three distinct longitudinally rec- tangular blotches on the dorsum, separated by narrow cream inter- spaces. The dorsal fin and posterior one-third of ventral fin are heavily flecked with brown. Distribution. — This species is known only from the Merida Andes in western Venezuela, where it occurs at elevations from 1600 to at least 2500 m, inhabiting subtropical and temperate areas. Remarks, — Rivero (1961:115) diagnosed Hyla paramica from H. jahni by the former having a ". . . rounded snout, less defined canthus and in lacking supratympanic fold, canthal, palpebral, and supratympanic lines and closely set dark dots above." He (1961: 116) stated that Hyla phitydactyla differed from H. jahni ". . . in the shorter snout and less defined canthus, absence of a light canthal and supratympanic line, narrower interorbital space, more vertical loreal region and different coloration [no dorsal spots]." Rivero based his descriptions of H. paramica on five specimens and of H. jahni on seven, all of which are rather poorly preserved; he examined no specimens of H. platydactyla. I have examined all of the specimens studied by Rivero and the holotype of H. platydactyla, together with several series from western Venezuela (total of 59 adults and subadults). I find no consistent structural differences, such as snout shape, definition of canthal ridge, or inclination of loreal region, by which to distinguish more than one taxon in western Venezuela. The dorsolateral stripe ( canthal, palpebral, and supratympanic lines of Rivero ) is variable; a stripe is evident in the supratympanic region in slightly more than half of the specimens; in some of these it extends posteriorly to the axilla. Also, in some of these specimens no stripe is evident on the canthus. On the basis of preserved specimens alone, there seems to be a continuum from well-defined to no stripes. Thus, only one species, H. platydactyla, can be recognized. The mating call consists of a series of short notes repeated at a rate of 540 notes per minute; the pulse rate is about 177 pulses per second, and the fundamental frequency at about 2790 Hertz is dominant ( Fig. 4A ) . FROGS OF THE HYLA BOGOTENSIS GROUP 29 At a locality 15-20 km NW of Merida, Venezuela, adults were found at night on leaves of herbs on a steep bank over a roadside ditch. Tadpoles were found in quiet pools in a cascading stream. Juan A. Rivero stated (pers. com.) that adults are found in brome- liads by day and that males call from bushes over streams at night. He also noted that the frogs produce a smelly exudate when handled. Cochran and Coin (1970:256) proposed the name combination Hyhi lahkiVis pJatydactyla; the name was used for the population of frogs in the eastern Andes of Colombia that is referable to the highly variable Hyla labialis, a species quite distinct from H. platydoctyla. RESUMEN Las ranas del grupo HyJa hogotensis se crian en los arroyos de las vertientes de los Andes en el occidente de Venezuela, Colombia, Ecuador, y Peru, y en las sierras de Costa Rica y Panama. Las ranas de este grupo se caracterizan por: 1) su tamafio moderado (longitud de cabeza-cuerpo de los machos hasta 45 mm, en las hembras hasta 53 mm); 2) el dorso de color verde opaco o carmelita con manchas pequeiias oscuras espacidas en el; 3) los dedos con tres cuarta partes cubiertos por la membrana interdigital; 4) la glandula del menton en el apice de este; 5) membrana axilar ausente; 6) el cuadratoyugal articulado con el maxilar; 7) los nasales pequenos, separados ampliamente en el medio; 8) la fon- tanela frontoparietal grande; 9) renacuajos con colas musculares, largas, la boca ventral grande, completamente bordeada por papilas labiales, con 4/5 — 8/10 hileras de dientes. Se analizaron 14 caracteres de ranas adultas por medio del programa de computacion de analisis discriminante multiple; los seis grupos resultantes fueron tratados como OTUs en los programas de NT-SYS, en el cual 11 caracteres de adultos y cinco de los renacuajos constituyeron los datos basicos para la construccion de los fenogramas. Los resultados de estos analisis son sostenidos por el analisis de las llamadas de reclamo de tres especies. Se reconocen seis especies en el grupo: 1) H. alytolylax especie nueva, laderas del Pacifico de los Andes en Ecuador; 2) H. bogo- tensis (Peters, 1882), Andes del centro y sur de Colombia; 3) H. colymba Dunn, 1931, sierras de Costa Rica y Panama; 4) H. denticulenta especie nueva, laderas del Caribe de los Andes en el norte de Colombia; 5) H. phyllognatha Melin, 1941, laderas amazonicas de los Andes en Ecuador y Peru; 6) H. platydoctyla Boulenger, 1905, Andes del occidente de Venezuela. Hyla para mica Rivero, 1961, y Hyla jahni Rivero, 1961, son colocadas en sinonimia de Hyla platydactyla Boulenger, 1905. 30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY SPECIMENS EXAMINED Hyla aUjtohjhx.— COLOMBIA: Choco: Camino de Yupe, 420 m, LACM 73000-1. Valle: Anchicaya, KU 148703-4. ECUADOR: Bolivar: Balsabamba, 800 m, KU 132432, 132545 ( tadpoles ) . Cotopaxi: 20.3 km W Pilalo, 830 m, KU 142857 (tadpoles). Pichincha: 3.7 km E. Dos Rios, 1190 m, KU 142856 (tadpoles); Las Palmas, 920 m, KU 132427; Tandapi, KU 111892-903, 111904 (skeleton), 111905-10, 112357-8 (tadpoles), 112359 (young), 117982-4, 118123 (tadpoles), 120851-60, 132425-6. Hyla hogotensi.s.— COLOMBIA: Cauca: Moscopan, 2500 m, UMMZ 121032. Cundinamarca: Bogota, 2630 m, CAS-SU 22970 (tadpole), 22971 (young), KU 125363, UMMZ 123946-8, ZMB 10209; Boqueron, 2900 m, FMNH 81916; Paramo de Palacio, Siberia, UPR-M 1800. HuHa: east of Paramo de Purace, 2500 m, LACM 50562. Hyla cobjmba.— COSTA RICA: Cartago: Moravia, 1200 m, KU 30886, 31864, 31865 (skeleton). PANAMA: Bocas del Tow: La Loma, 610 m, MCZ 10234-5; Rio Changena, 650 m, KU 104237 (young); Rio Changena, 830 m, KU 104236 (tadpoles). Code: El Valle, 560 m, AMNH 59606. Darien: Cerro Citurio, Serrania de Pirre, 1100 m, KU 116357; Cerro Mali, 1410 m, GML 4-00426; Laguna, 820 m, KU 77414; ridge between Rio Jaque and Rio Imamado, 730 m, KU 116779 (tadpoles), 116780-1 (young). Panama: Altos de Pacora, 740 m, KU 95979. Hyla denticulenta. — COLOMBIA: Antioquia: Valdivia, 1400 m, KU 133452. Santander: Charta, 2400 m, KU 133450-1, 139531 (tadpoles). Hyla phyllognatha.— ECUADOR: Morona-Santiago: Macas, AMNH 33911. Napo: Bermejo No. 4 (oil well site), 15 km ENE Umbaqui, 720 m, KU 123130-2; Rio Azuela, 1740 m, KU 143199-204, 143543 (young), USNM 193349-50; Rio Salado, 1 km upstream from Rio Coca, 1410 m, KU 146794 (tadpoles); Salto de Agua, 2.5 km NNE Rio Reventador, 1660 m, KU 143205, 143544 (young), 146793 (tadpoles); San Jose, AMNH 22164; 16.5 km NNE Santa Rosa, 1700 m, KU 143197-8, 143542 (tadpoles). Pastaza: Mera, 1140 m, KU 121418 (tadpoles), 121419 (young); Puyo, 960 m, FMNH 172632; Rio Alpayacu, 1 km E Mera, 1100 m, KU 121040. Tungitrahtia: 18.5 km E Banos, 1600 m, KU 141596 (young). PERti: Jimin: Perene, AMNH 17260, 17277. Puno: La Union, Rio Huacamayo, Carabaya, 610 m, BMNH 1907.5.7.36. San Martin: Roque, NHMG 471. Hyla platydactyla.— VENEZUELA: Merida: Camino de La Culata, UPR-M 2763, 2765; Carretera La Azulita, PUR-M 2762; Escorial, 2500+ m, FMNH 3567-8, NHMW 6397 (4), 6398, UMMZ 46465, 59016, 105439, UPR-M 3158-62; La Culata, 2700 m, AMNH 10636-40, BMNH 1905.5.31.83-87, NHMW 6396 (3), UPR-M 3150-2, 3932-5, 4330; La Mucuy, 2500 m, UPR-M 4321-2; Merida, 1620 m, BMNH 1909.4.30.81-83, 1912.11.1.83, 1947.2.13.14, MCZ 2523; 15-20 km NW Merida, road to La Azulita, 1700 m, KU 133430-3, 139521 (tadpoles); 32 km NW Merida, road to La Azulita, 2010 m, KU 139522 (tadpoles); Rio Albarregas, 2400 m, UPR-M 2767-70; Rio Chama, Merida, 1620 m, UMMZ 95310 (tadpoles). Tachira: 15 km from Delicias, to Rubio, 1800 m, UPR-M 2249-50, 2766; Guacharaquita, near La Grita, UPR-M 4858. FROGS OF THE HYLA BOGOTENSIS GROUP 31 LITERATURE CITED BOULENGER, G. A. 1905. Descriptions of new tailless batrachians in the collection of the British Museum. Ann. Mag. Nat. Hist., (7)16:180-184. Cochran, D. M. and C. T- Goin 1970. Frogs of Colombia. Bull. U.S. Natl. Mus., 288:xii+655 pp. Dlxox, W. J. (Ed.) 1971. BMD Biomedical computer programs. Univ. California Publ. Auto. Comp., 2, ed. 2:x+600 pp. DUELLMAN, W. E. 1963. A review of the Middle American tree frogs of the genus Ptychohyla. Univ. Kansas Publ. Mus. Nat. Hist., 15:297-349. 1966. Taxonomic notes on some Mexican and Central American hylid frogs. Ibid., 17:263-279. 1970. The hylid frogs of Middle America. Monog. Mus. Nat. Hist. Univ. Kansas l:xi+753 pp. 1972. South American frogs of the Hyla rostrata group (Amphibia, Anura, Hylidae). Zool. Meded. Leiden, 47:177-192. DuELLMAN, W. E. and M.J. Fouquette, Jr. 1968. Middle American hylid frogs of the Hyla microcephala group. Univ. Kansas Publ. Mus. Nat. Hist., 17:517-577. DuELLMAN, W. E. and L. Trueb 1966. Neotropical hylid frogs, genus Smilisca. Ibid., 17:281-375. Dunn, E. R. 1931. New frogs from Panama and Costa Rica. Occas. Papers Boston Soc. Nat. Hist., 5:385-401. 1944. Herpetology of the Bogota area. Rev. Acad. Colombiana Cien. Exact. Fisicas y Nat., 6:68-81. Kluge, a. G. and J. S. Farris 1969. Quantitative phyletics and the evolution of Anurans. Syst. Zool., 18:1-32. Melin, D. 1941. Contribution to the knowledge of Amphibia of South America. Goteborgs Kungl. Vetensk.-och Vitterh.-Sam. Handl., Ser. B, 1(4):1-71. Peters, W. 1882a. Fine neue Gattung von Batrachiem, Hijlonomus, aus Bogota. Sitzber. Gesell. Naturfors. Fr. Berlin, 1882(7): 107-109. 1882b. Der Namen der Batrachiergattung Hijlonomus in Htjloscirtus zu andem und legte zwei neue Arten von Schlangen, Microsoma notatum und Liophis Ygraecum. Ibid., 1882(8) : 127-129. RiVERO, J. A. 1961. Salientia of Venezuela. Bull. Mus. Comp. Zool., 126:1-207. RoHLF, F. J. and J. Kishpaugh 1966. NT-SYS, a system of multivariate statistical programs for use in numerical taxonomy. Dept. Entomology, Univ. Kansas. Taylor, Edward H. 1952. A review of the frogs and toads of Costa Rica. Univ. Kansas Sci. Bull., 35(1) :577-942. Vuillermier, B. S. 1971. Pleistocene changes in the fauna and flora of South America. Sci- ence, 173(3999) :771-780.