MCZ LIBRARY OCCASIONAL PAPERS AUG 4 1989 r- r-i "A of the ■ ■ ' - MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 132, PAGES 1-14 JULY 20. 1989 NEW SPECIES OF FROGS, CENT ROLEN ELLA, FROM THE PACIFIC VERSANT OF ECUADOR AND SOUTHERN COLOMBIA By William E. Duellman' and Patricia A. Burrowes^ During the past two decades field parties from The University of Kansas have been surveying the anuran fauna of the Pacific slopes of the Andes in southern Colombia and Ecuador. Earlier collections of centrolenid frogs from Ecuador formed the basis for a synopsis of the members of that family in Ecuador by Lynch and Duellman (1973). Subsequently, four more new taxa were named by Duellman ( 1980, 1981), and one was named by Flores( 1985). Other unnamed taxa have accumulated during the past decade. Collections made by the junior author at La Planada in Colombia contained representa- tives of these new taxa. Her collections provided the impetus to prepare descriptions of the species named herein. In addition to describing three new species, we take this opportunity to present a review of the geographic and altitudinal distribution of the species of centrolenids on the Pacific versant of Ecuador and southern Colombia, north to Departamento Valle del Cauca. SYSTEMATICS At present, the recognition of species groups in Centrolenella (fide Savage, 1967) is strictly phenetic. As noted by Lynch and Duellman (1973) 'Curator, Division of Herpetology, Museum of Natural History, and Professor, Department of Systematics and Ecology, The University of Kansas, Lawrence, Kansas 66045-2454 U.S.A. ^Assistant Professor, Department of Biology, Colegio Universitario de Cayey, Cayey, Puerto Rico 00633 U.S.A. 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY and Cannatella and Duellman (1982), many South American species do not fit into Savage's groups, which were based on characters of Central American taxa. Consequently, we offer no suggestions about the relationships of the new species. The format for the following diagnoses and descriptions follows those of Lynch and Duellman (1973), Duellman (1981), and Cannatella and Duellman (1982). Centrolenella hallux new species (Fig. 1) Holotype.— KU 164725, an adult male, from 14 km (by road) west of Chiriboga (00° 1 8'S , 78^49^) , 1 960 m , Provincia de Pichincha, Ecuador, one of a series collected on 8 May 1975 by William E. Duellman, John E. Simmons, and Linda Trueb. Paratypes.— KU 164726-32 collected with the holotype; KU 164733 from QuebradaZapadores, 5 km (by road) east-southeast of Chiriboga, 2010 m, Provincia de Pichincha, obtained on 11 May 1975 by William E. Duell- man; IND-AN 1532, 1708, 17 17-26 from ReservaLaPlanada, 7 km (by road) south of Chucunes, 1780 m, Departamento de Narino, Colombia, collected April-June 1986 by Patricia A. Burrowes; KU 200272-77 from the same locality, collected by Patricia A. Burrowes and William E. Duellman on 28-30 March 1984. Diagnosis. — (1) Vomerine teeth absent; (2) bones green; (3) parietal peri- toneum white; visceral peritoneum clear; (4) color in life, dark Ume green with golden yellow flecks; in preservative, lavender with cream flecks; (5) modal webbing between outer fingers III2V2 — 21 V; (6) modal webbing on foot 12— 2ni— 21111— 2IV2V2— IV; (7) snout bluntly rounded in dorsal view and in profile; (8) dorsal skin smooth; (9) arms and legs lacking tubercles and dermal folds; (10) humeral spine present; (11) lower four-fifths of tympanum visible, oriented dorsolaterally with sUght posterior inclination. No other species of Centrolenella on the Pacific versant is green with golden yellow flecks and no black dots. The combination of pale flecks and dark dots occurs in (1) C. peris ticta, which has a row of tubercles along the ventrolateral edge of the forearm and tarsus, (2) C. lynchi, which has a low ulnar fold and finely spiculate dorsal skin, and (3) some individuals of C. prosoblepon, a larger species with vomerine teeth. Description. — Head slightly wider than body; width of head 34.4-39.6 (3c =37.0, n=28) percent of snout- vent length; snout moderately short, bluntly rounded in dorsal view and in profile; canthus rounded; loreal region barely concave; lips not flared; nostrils two-thirds of the distance from eyes to tip of snout, barely protuberant, directed anterolaterally; intemarial area flat. Eye moderately large, directed anterolaterally; supratympanic fold absent; lower NEW CENTROLENID FROGS X ^ i\ft Fig. 1. Top. Holotype of Centrolenella hallux, KU 164725, male, 21.3 mm SVL. Bottom. Paratype of Centrolenella orejuela, IND-AN 1520, female, 29.6 mm SVL. 4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY four- fifths of tympanum visible, oriented dorsolaterally with slight posterior inclination. Vomerine odontophores absent; vocal slits extending from mid- lateral base of tongue to angles of jaws. Humeral spine large, blunt, at 45° angle to humerus; ulnar fold and tubercles absent; first finger shorter than second; fourth finger longer than second; lateral fringes absent on fingers; webbing absent between first and second fingers, vestigial between second and third; webbing formula for outer fingers ni(2+-2V2) — (2-2*)IV; discs broadly elliptical; subarticular mbercles small, subconical, simple; palmar tubercle rectanglar, simple; nuptial excres- cences absent. Hind limbs slender; length of tibia 56.3-64.5 (:f=60.4, n=28) percent of snout-vent length; tarsal folds and tubercles absent; inner metatar- sal tubercle small, elliptical; outer metatarsal tubercle absent; subarticular tu- bercles small, subconical; supernumerary tubercles absent; feet about three- fifths webbed; webbing formulaI2— (2-2*)n( 1-1 V2)— (2-2^)10(1 -IV2)— (2-2*)IV(2-2V2) — (1-1*)V; discs on toes slightly smaller than those on fingers. Skin on dorsal surfaces smooth; skin on belly and proximal posteroventral surfaces of thighs granular; other ventral surfaces smooth; anal opening directed posteriorly at upper level of thighs; anal folds and tubercles absent. Color in preservative: Dorsal surfaces of head, body, forearms, thighs, shanks, and tarsi lavender with small cream flecks; hands, feet, margin of upper lip, and ventral surfaces cream. Color in life: Dorsum dark Ume green with minute pale gold-white flecks that are most numerous on the hind limbs; tips of digits yellowish green; venter translucent; parietal peritoneum creamy white; visceral peritoneum clear; heart not visible; bones green; iris bronze-copper with fine black reticu- lations. Measurements: Ranges of variation in mm (mean ± 1 standard deviation) for 25 males are: snout- vent length 19.2-22.2 mm (20.6 ± 0.9 11), tibia length 11.5-13.2 (12.4 ± 0.432), foot length 9.5-11.4 (10.3 ± 0.562), head width 7.2-8.3 (7.6 ± 0.264), head length 6.5-7.5 (6.9 ± 0.268). Ranges in mm (mean) for three females are: snout-vent length 21.0-23.3 (21.9), tibia length 12.8-14.1 (13.2),footlengthll.2-11.6(11.3),headwidth7.8-8.2(7.9),head length 7.0-7.7 (7.4). Distribution. — Centrolenella hallux is known from three localities in humid upper montane forest at elevations of 1700-2010 m on the Pacific versant of southern Colombia (Departamento de Nariiio) and northern Ecua- dor (Provincia de Pichincha). Ecology and behavior. — All individuals were found at night on the upper surfaces of leaves of bushes and small trees 0.5-2.0 m above small streams, or on ferns over roadside ditches. The call is a short, high-pitched trill. One male (IND-AN 1725) was on a leaf 5 cm below another leaf on which there was a clutch of 18 eggs having clear jelly. NEW CENTROLENID FROGS 5 Etymology. — The specific name is Latin; it means "gold dust" and is used in reference to the minute gold flecks on the dorsum. Centrolenella orejuela new species (Fig. 1) Holotype. — KU 145081,an adult female Jrom between EI Tambo and La Costa, 800 m, Departamento de Cauca, Colombia, obtained on 17 August 1937 by Kjell von Schneidem. Paratypes. — KU 145080, an adult female from the type locality; IND- AN 1520-21, adult males from Pialapi, 1250 m, Departamento de Narino, Co- lombia, collected on 20 May 1986, by Patricia A. Burrowes; LP248, also from Pialapi, obtained on 31 October 1984 by Jorge E. Orejuela. Diagnosis. — (1) Vomerine teeth present; (2) bones pale green; (3) parietal peritoneum white; visceral peritoneum white; (4) color in life, uniform dark green dorsally; in preservative, dull gray; (5) modal webbing on hand 11 1 V2 — 3mi V2— IIV; (6) modal webbing on footll— ini— lini— 1 VI— IV; (7) snout truncate in dorsal view and in profile; (8) dorsal skin smooth; (9) arms and legs lacking tubercles and dermal folds; (10) humeral spine absent; (11) lower four-fifths of tympanum visible, oriented dorsolaterally with slight posterior inclination. The only other large, uniformly green centrolenids on the Pacific versant in South America and in Central America are C. ilex and C. prasina. In the former, the visceral peritoneum is clear, the head is flat and broad, and the limbs are slender, as opposed to a deeper and proportionately narrower head and more robust Umbs in C. orejuela. Centrolenella prasina differs in being larger (males 33.0-34.5 mm) and in having much less webbing on the hands and feet. Description. — Head as broad as body, slightly wider than long; width of head 35.8-38.2 (3c =36.7, n=6) percent of snout- vent length; snout truncate in dorsal view and in profile; canthus rounded; loreal region concave; lips not flared; nostrils four-fifths of the distance from eyes to tip of snout, slightly protuberant, directed dorsolaterally; intemarial area slightly depressed. Eye moderately large, directed anterolaterally; supratympanic fold weak, curved posteroventrally from eye; lower four-fifths of tympanum visible, oriented dorsolaterally with slight posterior inclination. Dentigerous processes of vomers transverse between elliptical choanae, narrowly separated medially, each bearing 2-5 teeth; vocal slits extending from midlateral base of tongue to angles of jaws. Humeral spine absent; ulnar fold and tubercles absent; first finger barely shorter than second; fourth finger longer than second; lateral fringes present on fingers; webbing absent between first and second fingers; webbing formula for outer fingers n(r-lV2)— (3--3)III(r-lV2)— (l-l^IV; discs 6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY broad, truncate; subarticular tubercles moderately large, round, simple; supernumerary tubercles minute, conical, on basal segments of Digits II-IV; palmar tubercle bifid; nuptial excrescences absent. Hind limbs moderately robust; length of tibia 56.6-64.3 (3c =61 .2, n=6) percent of snout-vent length; tarsal folds and tubercles present; dermal fold present on outer edge of fifth toe; inner metatarsal tubercle small, elliptical; outer metatarsal tubercle absent; subarticular tubercles small, round; supernumerary tubercles minute, on proximal segments of all toes; toes webbed to bases of discs; discs slightly smaller than those on fingers. Skin on dorsal surfaces smooth; skin on belly and proximal posteroventral surfaces of thighs granular; other ventral surfaces smooth; anal opening directed posteriorly at upper level of thighs; anal folds and tubercles absent. Color in preservative: Dorsum, including hands, feet, and webbing dull gray; venter cream. Color in life: Dorsum uniform dark olive green; venter d-anslucent with greenish tint; parietal and visceral peritonea creamy white with golden suffusion (posterior part of visceral peritoneum clear in IND-AN 1520); heart not visible; bones green; iris dark gray with yellow ring around pupil. Measurements: Ranges of variation (mm) with means in parentheses for three males, followed by those of two females, are: snout-vent length 27.3-28.3 (27.9) 29.6-33.7 (31.6), tibia length 16.3-18.2 (17.5) 17.9-19.1 (18.5), foot length 14.2-15.9 (15.0) 15.6-17.2 (16.4), head width 9.9-10.8 (10.3) 10.6-12.3 (11.4), head length 9.4-10.1 (9.6) 9.7-10.7 (10.2). Distribution. — Centrolenella orejuela is known from two localities in lower humid montane forest at elevations of 800 and 1250 m on the Pacific versant of southern Colombia. Ecology and behavior. — At Pialapi males were found at night on rocks along a steep stream bank or on rocks within the stream; all were in the spray zone of cascades. Etymology. — The specific name is a noun in apposition for the Orejuela family (Jorge, Anamaria, and Tomas), who reside at, and administer, the Reserva La Planada, Colombia. Centrolenella scirtetes new species (Fig. 2) Holotype.— KU 202720, an adult male, from 1.4 km (by road) southwest of Tandayapa (00°07'S, 78°40'W), 1820 m, Provincia de Pichincha, Ecuador, obtained on 2 April 1984 by David M. Hillis. Paratypes. — IND-AN 1405 and 1533, adult females, from Reserva La Planada, 7 km (by road) south of Chucunes, 1780 m, Departamento de Narino, Colombia, collected 2 May and 3 June 1986 by Patricia A. Burrowes. Diagnosis. — (1) Vomerine odontophores absent; (2) bones green; (3) NEW CENTROLENID FROGS Fig. 2. Paratype of Centrolenella scirtetes, IND-AN 1533, female, 26.1 mm SVL. parietal peritoneum white; visceral peritoneum clear; (4) color in life, green with black dots; color in preservative, pale lavender with dark dots; (5) modal webbing on hand 112— 31113— 2IV; (6) modal webbing on foot II— 211 1— 2ini — 2IV2 — IV; (7) snout truncate in dorsal view, bluntly rounded in profile; (8) skin on dorsum finely spiculate in male, smooth in females; (9) arms and legs lacking tubercles and dermal folds; (10) humeral spine present in male; (11) lower four- fifths of tympanum visible, directed posterolaterally with shght dorsal inclination. The combination of the absence of vomerine teeth and presence of humeral spines and dark dots on the dorsum distinguish C. scirtetes from all other centrolenids on the Pacific versant of South America except C. peristicta. The latter is a smaller species (males 18.7-20.6, females 20.4-21.0 mm snout- vent length) with a rounded snout in dorsal view, shagreened dorsal skin, row of low tubercles on the ventrolateral edges of the forearm and tarsus, and the entire tympanum visible. Description. — Head as wide as body, broader than long; width of head 37.2-38.3 (Jc =37.9, n=3) percent of snout-vent length; snout short, truncate in dorsal view, bluntly rounded in profile; canthus rounded; loreal region barely concave; lips not flared; nostrils four-fifths distance from eyes to tip of snout, protuberant, directed dorsolaterally; intemarial area slightly de- pressed. Eye moderately large, directed anterolaterally; supratympanic fold absent; lower four-fifths of tympanum visible, directed posterolaterally with 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY slight dorsal inclination. Vomerine odontophores absent; vocal slits extend- ing from midlateral base of tongue to angles of jaws. Humeral spine moderately large, blunt, at 45° to humerus; ulnar fold and tubercles absent; first and second fingers equal in length; fourth finger much longer than second; lateral fringes absent on fingers; webbing absent between first and second fingers; webbing formula 112 — 31113 — 2IV; discs broad, truncate; subarticular tubercles, small, round, simple; palmar tubercle bifid; nuptial excrescenses absent. Hind limbs moderately slender; tibia length 55.6-60.1 (3(=58.0, n=3) percent of snout- vent length; tarsal folds and tu- bercles absent; inner metatarsal tubercle small, elliptical; outer metatarsal tubercle absent; subarticular tubercles small, round; supernumerary tubercles minute, subconical, on proximal segments of all toes; feet about two-thirds webbed; webbing formula II— 2II(l-r)— 2m(l-l*)— 2IV2— IV; discs on toes slightly smaller than those on fingers. Skin on dorsal surfaces in females smooth, finely spiculate (especially on upper lip and in tympanic region) in male; skin on belly and proximal postero ventral surfaces of thighs granular; other ventral surfaces smooth; anal opening directed posteriorly at upper level of thighs; two pairs of tubercles on proximal surfaces of thighs below anal opening. Color in preservative: Dorsal surfaces of head, body, forearms, thighs, shanks and tarsi pale lavender with black dots (and white spicules in male); hands, feet, and ventral surfaces cream. Color in life: Dorsum pale lime green with black flecks; digits green with orange-yellow discs; venter translucent with a greenish tint; parietal perito- neum creamy white; visceral peritoneum clear; heart not visible; bones pale green; iris creamy bronze with a copper suffusion and fine black reticulations. Measurements: The measurements (in mm) for the one male and two femalesare,respectively:snout-ventlength24.4,25.8,26.1;tibialengthl4.4, 14.5, 15.5; foot length 1 1.3, 11.6, 12.4; head width 9.3, 9.7, 9.9; head length 8.5, 8.6, 8.8. Distribution. — Centrolenella scirtetes is known only from two localities at elevations of 1780 and 1820 m in the humid upper montane forest on the Pacific versant of northern Ecuador and extreme southern Colombia. Ecology and behavior. — All individuals were found at night on large leaves of Araceae or palm fronds overhanging small streams. The two females contain dark green eggs. Etymology. — The specific name is derived from the Greek skirtetes and is used as a noun in apposition. The name means leaper and is used in reference to the jumping abilities of Centrolenella, and not coincidentally, to the same abilities of the collector of the holotype, David M. Hillis. NEW CENTROLENID FROGS 9 DISCUSSION We are aware of 13 other species of centrolenid frogs from the Pacific slopes of Ecuador and southern Colombia, northward to include the Depar- tamento Valle del Cauca. Of these, three species oiCentrolenella are known only from southern Colombia. The Colombian taxa include a species from the Anchicaya Valley in Departamento Valle del Cauca being named by Charles W. Myers and another species referred to C. ilex by Lynch and Duellman (1973). However, the latter species actually may represent another unnamed species (C. W. Myers, pers. comm.). Centrolenids usually are restricted to the vicinity of streams having at least moderate gradients. An exception is C. buckleyi, which at some high- Andean sites occurs in the vicinity of jwnds and drainage ditches. High-gradient streams are most numerous on the slopes of the Andes, where there is an altitudinal gradient in temperature and vegetation. Using the bioclimatic and ecological maps of Colombia (IGAC, 1977) and Ecuador (Canadas, 1983) based on the Holdridge (1967) system, we recognize four major vegetation types inhabited by centrolenids on the Pacific versant of Ecuador and southern Colombia. 1. Lowland rainforest (bosque muy hiimedo tropical and bosque muy humedo premontano) extending from sea level to elevations of about 600 m. 2. Humid lower montane forest (bosque muy humedo montano bajo) ranging from about 600 to 1500 m. 3. Humid upper montane forest (bosque muy humedo montano) extending from about 1500 to 2500 m. 4. Paramo and subparamo (p^amo subalpino), the supra-treeline vegeta- tion zones above 2500 m. The distribution of centrolenids with respect to these vegetation zones (Fig. 3) reveals that species richness is greatest in the humid montane forests — 12 species are in the upper montane forest and 14 species in the lower montane forest. Seven of these species occur in both vegetation zones. Five species occur in the lowland rainforest; of these, four also range into the lower montane forest and two into the upper montane forest. Only one species, C. buckleyi, occurs in the paramo and subparamo. The greatest diversity of centrolenids in the montane forests probably reflects an adundance of high-gradient streams there, in contrast to the lowland tropical rainforest and the paramo. Also, montane forests are charac- terized by high humidity throughout the year. This is important to the survival of centrolenid eggs, which are deposited on vegetation above the water [rocks in spray zones of waterfalls in the case of Centrolene geckoideum (Lynch et al, 1983)]. 10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY 4000 3000 2000 1000 M § ^ -aj ^ .c; 5 "33 S^ O .to C5, I $ 5 C3 ^ I I ' !lj