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1071 JOURNAL OF HERPETOLOGY 5(2.4): 97-101 A New Striped Treefrog from Central Colombia William F, Pyburn and M. J, Fouquette, Jr. Depertment of Bicloay, University of Texas at Arlington 76010; and Depariment of Zoology, Arizona State University, Tempe 85281 ABSTRACT — Hyla wandae is described from the western edge of the Colombian Ilanos. It seems most closely related to Myla staufferi, H. parkeri, and H. squalirosiris, from which it differs in structural features and color pattern. The mating call of H. wandée is a distinctive series of loud, harsh, regularly tepeated notes, Mr. Jerry Glidewell and the senior author collected ten specimens of an apparently unnamed hylié frog in June, July, and August 1969, in Central Colombia. The collecting locality, near Villavicencio, lies at the base of the eastern Andes in what once may have been piedmont forest (Bates, 1948), but is now cleared pasture-land, The frogs were calling from grass clumps and low shrubs around temporary pasture pools. Specimens are deposited in the U, S. National Museum (USNM), The University of Texas at Arlington (UTA), Arizona State University (ASU), University of Kansas Museum of Natural History (KU), and American Museum of Netural History (AMNH). Hyle wandae, sp. nov. (Fig. 1) Holotype. — USNM 192305; 12 km NNE of Villavicencio, Meta, Colombia; elevation about 580 m; 30 July 1969; Jerry R. Glidewell and William F. Pyburn, collectors. Paratypes. ~ USNM 192306-192310 (5 specimens), same data as holotype; UTA 2819, ASU 13129, 13 August 1969, same locelity; UTA 2821, 21 June 1969, same locality; KU 131717, 18 km NNE of Villavicencio, 620 m, Diagnosis. — Small hylids (males to 26.9 mm snout-vent) of the rubra complex, with 4 dark, dorsolateral stripes; indistinct transverse bars on shanks, Snout pointed; head about as long as wide or slightly longer: tympanum about half of eve diameter; no ulnar or tarsal folds. Digital discs subtruncate, those of finger 3 and 4 and toe 4 about as wide as tympanum; finger webbing virtually lacking; toes about two-thirds webbed, reduced to vestige between toes 1 and 2. Mating call a sequence of short, harsh, regularly repeated buzzes; note duration somewhat greater than interval between notes. Description of Holotype. — Adult male, snout-vent length 23.4mm. Head flat, about as wide (7.5mm) as long (7.7mm); dorsal espect of snout pointed, lateral aspect of snout acutely rounded and projecting beyond lower jaw; snout rather elongate (3.6mm, anterior corner of eye to tip of snout}; canthus rounded; loreal region nearly flat (barely concave); nostrils somewhat protuberant, the internarial distance (1.6mm) less than width of upper eyelid (1.9mm); eves somewhat protuberant, length (2.5mm) almost equal to interorbital distance (2.6mm) and greater than eye (anterior corner) to nostril (posterior edge) (2.2mm); tympanum distinct, small (1.3mm|, its diameter about half of eye length; a weak supratympanic fold covers the dorsal edge of the tympanum, Tongue rounded, notched posteriorly, edges not adherent, posterior edge about 1/4 free: vocal slits large, lateral and posterior to tongue; vomerine odontoids in two short diagonal groups, their medial ends deflected posteriorly, Iving between the posterior level of the small, elliptical choanae; vocal sac lerge, single, median subgular, extending laterally above base of arm when deflated. 97 NEXT PREV 98 WILLIAM F, PYBURN AND M. J. FOUQUETTE, JR. FIGURE 1, Hyla wandie, 12 km NNE Vilavicenci, Colombe. Body rather long; skin smooth dorsally, venter coarsely granular, chest and throat finely ‘granular; thoracic fold present; a very small axillary membrane; anal flep simple. Forearm without folds or tubercles; a weak dermal fold on wrist; finger webbing rudi mentary, essentially lacking; finger discs subtruncate, width of discs of third and fourth fingers equal to diameter of tympanum; subarticular tubercles small, round to subconical; superumerary tubercles very small, few, indistinct; palmar tubercle large, bifid; thenar tubercle poorly defined; thumb shorter than second finger: prepollex not enlarged; no prepollical spine or nuptial excre- sence. Thigh length 10.0mm, shank length 11,4mm; no tarsal fold or tubercles: no calcar; meta- tarsal tubercles distinct, inner elliptical, outer smaller and round; subarticular tubercles small, round; few, very small, indistinct supernumerary tubercles; tor discs subtruncate, somewhat more rounded than those of fingers; toes about 2/3 webbed, web vestigial between toes 1 and 2, extending trom proximal end of penultimate phalanx of toe 2 to proximal end of antepenultimate of toe 3, from middle of penultimate phalanx of toe 3 to proximal end of antepenultimate of toe 4, from proximal end of antepenultimate phalanx of toe 4 to distal end of penultimate of toe 5; heel of adpressed leg reaches posterior edge of eve. Color (alcohol after formalin) of dorsum ray-brown with four dark brown longitudinal stripes, the dark stripes on each side separated from each other by anarrow postorbital light strine: the more lateral dark stripe extending from posterior comer of eye, along supratympanic fold to about midway back on each side; the more medial dark stripe extending from upper eyelid to near base of thigh, not so dark as lateral stripe; loreal region dark brown, fading somewhat on upper lip; ‘an indistinct, dark interocular bar with light gray line along its anterior edge; upper surfaces of arm, hand, and fingers medium gray-brovin; upper surface of thigh uniform medium gray-brown: upper surface of shank with about § indistinct, dark cross-bars; upper surfaces of tarsus, foot and toes medium gray-brown; palmar surface, lower sides of thigh, shank and foot pale gray-brown; belly and lower sides of body cream white. Color in lie similar to color in preservative except light parts of dorsum more olive or yellowish-gray, dark dorsal stripes and bars on shanks more distinctive than in preservative. NEXT PREV NEW TREEFROG FROM COLOMBIA 99 Paratype Variation. — The chest fold is incomplete medially in USNM 192308 and 192309. Tho thenar tubercle is distinct in USNM 192306, 192307, 192309, ASU 13129, and KU 131717, but poorly defined in the others. USNM 192310, UTA 2819, and KU 131717 have a very shallow tongue notch, and the vomerine odontoids are poorly developed in USNM 192307 and 192310 {absent on right); the number of odontoids is very difficult to discern, but appears to be primarily 5 on each side. In life the skin had scattered tubercles. The dorsal coloration of the preserved specimens varies from dark brown (USNM 192310) to light gravish tan (USNM 192306): dorsal stripes and limb bars also vary in intensity but are discernable in all members of the series. In KU 131717 the dark markings on the shank are spots rather than bars. USNM 192307, 192309, and KU 131717 have a dark spot in the center of the rostrum, and some specimens (USNM 182309) have a dark interocular triangle rather than a bar. UTA 2821 is a juvenile; all other specimens in the type series are adult males, Variation in some mensural characters is indicated in Table 1. TABLE 1. Means (ranges in parenthesis) of several measurements of Hyla wandae (in mmr). Type series from near Villavicencio, July-August 1989; Hutchison collection from near Villavicencio and Puerto Lopez, October 1965-January 1966. ADULT MALES TOWENICE TYPE SERIES. HUTCHISON COLL. PARATYPE Number ° 13 1 ‘Snout-vent fength 24.9 (23.4269) 21.1 (19.5-23.0) 174 Heed length 79 (7.68.4) 7.216.579) 61 Head width 777.48.4) 65 (6.0.7.3) 57 Eye length 26 |233.0) 23.(2.1-25) 24 ‘Tympanum tength 13 11.2-41.4) 1.0 (0.9-1.1) 09 Shout lenath (from anterior corner of eye) 3.6 (9.4-4.0) = 34 Eye to nostrit 2.4 (222.6) 24 (1.82.3) = ‘Shank length 120 (11.4-12.8) 10.7 (9.7-11.5) 94 Thigh length 99 9.311.1) = 85 Width of disco: Finger 3 1.3(1.14.5) 08 (0.80.9) 09 Fingor 4 1.2 10.91'5) = 09 Too 131.245) 08 (0.809) 09 Referred Specimens. — A collection made in Colombia by Victor Hutchison in 1965-1966, and deposited in the American Museum of Natural History and the U. S. National Museum, contains 13 specimens (all males) referrable to H. wandae. The following were examined: AMNH 75688, 82128-82131, 18 January 1966, Metz, Puerto Lépe2, cast side of Rio Meta; AMNH 75691-75697, 13 November 1965, 7 km $ of Villavicencio; USNM 158028, 12 October 1968, Puerto Lopez. They ate smaller (Table 1) but agree well with the type series. Those from near the type locality as well as the Puerto Lépez specimens are significantly smaller. Perhaps this is correlated with the time of the collections; adults of the type series were collected in late July and ‘August, whereas Hutchinson's specimens were taken between October and January. Mating Call and Behavior. ~ Recordings were made of mating calls of Hyla wandze at the type locality on the evening of 30 July 1969, at an air temperature of 21.9 C, using a Uher 4000 Report-L recorder. The calls were later analyzed in the laboratory using a Kay Electric Sonagraph, Model 6061B. The call (Fig. 2) consisted of a series of single notes, repeated regularly. There were two to 18 notes ina call sequence: the interval between notes in a series was about 0.5 second, with a mean note-repetition rate of 47.7 per minute. Each note had a fairly uniform series of about 90 pulses, producing a buzzing sound, An analysis of 12 calls of 3 individuals in the type series revealed a mean note duration of 0.673 (0.653-0.696) second; mean fundamental frequency was 131.3 (121,7-140,0) hertz (Hz); there was a dominant frequency band of about 400 Hz width centered at about 4893 (4800-5050) Hz. The fundamental and its first harmonic (about 260 Hz) NEXT PREV 100 WILLIAM F. PYBURN AND M. J. FOUQUETTE, JR. SECONDS FIGURE 2 Audiospectrogram of mating call of Myla wandae, 12 km NNE Villavicencio, Colombia, 30 July 1969 (air 21.9 C). Effective bandwidth of analyzing titer 90 Hz (narrow ‘band, tape speed one-half original, scale magnified approx. 2 X). were weakly emphasized; there is another emphasized band at about 3000 Hz. Individuals calling near each other gave the impression of intentionally alternating their calls. Victor Hutchison recorded calls of some of the individuals he collected at Puerto Lépez. Several of these were analyzed and agree well with the calls recorded at the type locality. On the evening of 29 July 1969, males at the type locality called from the ground in dense ‘grass and from perches low in grass clumps and shrubs around temporary pasture pools (air 20.3 C). Chorusing was occasionally interrupted by silent intervals of several minutes, then suddenly resumed, The calls were loud and unlike those of any other species at the pools, but the frogs called from concealed perches and were very difficult to capture. When illuminated with a hand light they almost immediately jumped out of the lighted area or scurried away in the dense vegetation, a behavior often observed in Panamanian Hyla stavfferi Calling continued the following night (air 21.9, water 25.7 C). On 13 August, a calling male (UTA 2819) was taken from a low herb in 2 pool at 2045 hours (local), at which time the air was 21.0 C. An immature specimen (UTA 2821) was found on a grass blade well away from the water on 21 June. Relationships. — Hyla wandze is a member of the Hyla rubra complex. Specifically, its relationships lie with that group including Hyla staufferi of Central America, H. parkeri of Bolivia, and H. squalirostris of southern South America. All three of the latter and H, wandae are small, striped frogs with protruding, pointed snout, single median subgular vocal sac, reduced webs (relative to other members of the rubra complex), and no tarsal fold. Hyla wandee differs from the nearest population of Hyla staufferi, H. «. altae of Panam, in color pattern and mating call. H. s a/tae always have a single, lengthwise stripe on the shank (Leén, 1969), whereas the shank of H. wandae is marked with several transverse bars or spots, as in the more northern H. s. staufferi. The latter differs from H. wandae in generally lacking complete dorsal stripes and in breeding call (sce Leén, 1969: 534, Pl. 4). Leén reported a note duration of 0.15 (0.14-0.18) second in Panamanian H. s. altae, as compared to about 0.67 reported above for wandae; and the interval between notes in a/tae was reported as about 2.5 times the note duration, as compared with slightly less than the note duration in wandae. The dominant frequency bands reported for altae lie about 1000 Hz below those for wandae. Hyla wandae differs from both H. parkeri and H. squalirostris in having somewhat more protuberant nostrils and eyes, and in the size of its finger and toe discs. The width of the fourth finger disc of wandae is about 16 per cent of the head width, whereas this value is about 11 per ‘cent in parker! and 9 per cent in squalirostris. The width of the fourth toe disc is about 17 per cent of the head width in wandae, approximately 12 per cent in parkeri, and 9 per cent in squalirostris. NEXT PREV NEW TREEFROG FROM COLOMBIA 101 Hyla parkeri differs from H. wandae in having, on each side, a single broad, dorsolateral gray stripe, usually dark-edged medially, and a distinctive white lip band beginning just anterior to the eye (Gaige, 1929). Some specimens of H. parkeri also have a fine, dark vertebral line. The call of parkeri has not been described. Hyla squatirostris has a much more elongated snout than any of the other species in the group. Head width is about 81 per cent of head length in squalirostris, compared to about 98 per cent in wandae. It also differs from wandae in having a single lengthwise stripe on the shank as in Panamanian staufferi. Width of the disc of toe 4 is about equal to tympanum diameter in wandae; in squalirostris it is about 55 per cent of the tympanum diameter. Barrio (1963) analyzed the call of squalirostris at different temperatures. At an air temperature comparable to that of our wandae sample (22 C), the note duration is much shorter (0.24-0.37 sec), with an interval about twice the note duration, Two dominant frequency bands ate centered at about 6000 and 4000 Hz, significantly higher than in wandae, Each note in squalirostris has a trilled sound, being composed of about seven pulses with a repetition rate of about 23-25 per second, as compared to the buzzing sound of wandae, with pulses repeated at about 130 per second. Comparative Material. — \n addition to the material of Hyla wandae already cited, the following comparative material was examined: Hyla parkeri Gaige. Bolivia: Santa Cruz— Buenavista (Univ. Michigan Mus. Zool. 67461 [ten paratypes] , USNM 84359-84360, 101440-101441 [four paratypes] }. Hyla staufferi Cope. Panama: Chiriqui—David (Univ. Texas Memorial Mus, [TNHC] 32013); 4 mi N David (TNHC 32014), Panamé— 2 km WSW Chepo (KU 101680-101688); 2 km N Toucmén (KU 101689-101695). Canal Zone: (TNHC 24406). México: Veracruz— El Encinal {UTA 2044, 2084, 2135, 2168-2171, 2174, 2356-2357, 2378, 2521, 2823-2834). Hyla squalirostris A. Lutz. Brazil: Sio Paulo— Bonito, Serra da Bocaina (USNM 96719, 96720 [syntypes] ; USNM.96609, 96611, 96613 [paratypes] ; USNM 96721). Argentina: Buenos Aires (Mus. Compar. Zool., Harvard Univ. 35421-35422). ACKNOWLEDGMENTS It is a pleasure to name this species in honor of Wanda Carl Pyburn for her valuable contributions to our work in Colombia and elsewhere in Latin America, We are indebted to Mr. and Mrs, Tom Kirby for permitting access to their property near Villavicencio, and for many other courtesies; and also in Colombia, to Sr. Eduardo Valdarrama, Dr. José M. Idrobo, and Dr. Federico Medem, for helpful suggestions concerning travel to certain remote areas. Special thanks are due W. Frank Blair, whose efforts made possible the work in Colombia, The following individuals and institutions kindly made available specimens in their care: William E. Duellman, Museum of Natural History, Univ, of Kansas; Robert F. Martin, Memorial Museum, The Univ. of Texas (Austin); James A. Peters, U. S, National Museum; Charles F. Walker, Univ. of Michigan Museum of Zoology; Ernest E. Williams, Museum of Comparative Zoology, Harvard Univ.; and Richard G. Zweifel, American Museum of Natural History. In addition, W. E. Duellman examined specimens of the type series and furnished helpful advice. Victor Hutchison loaned tapes of mating calls and agreed to our description of the new form separate from the report of his Colombian collection. J. R. Glidewell assisted in field work. A. J. Delahoussaye assisted in call analysis. Field work by Pyburn was supported by NSF grant GB-8265 to W. F. Blair; call analysis was supported by NSF grant GB-12560 to Fouquette. This paper is a contribution to the Evolutionary and Ecological Diversity section of the Integrated Research Program on Origin and Structure of Ecosystems (formerly Convergent and Divergent Evolution IRP) of the International Biological Program. LITERATURE CITED Barrio, Avelino, 1963, Influencia de la temperatura sobre el canto nupcial de Myla squalirostris A. Lutz (Anura, Hylidae). Physis 24: 137-142, Bates, Marston. 1948, Climate and vegetation in the Villavicencio region of eastern Colombia. Geogr. Rev. 38: 555-574, Gaige, Helen T. 1923, Three new tree-frogs from Panama and Bolivia. Occas. Pap. Mus. Zool. Univ. of Michigan (207): 1-6. LLedn, Juan R. 1969, The systerratics of the frogs of the Hy/a rubra group in Middle America. Univ. Kansas Publ. Mus. Nat 18: 505-545.

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