Species status of Centrolene lema Duellman and Señaris, 2003 (Amphibia: Centrolenidae) revealed by Integrative Taxonomy Author Castroviejo-Fisher, Santiago Author Guayasamin, Juan M. Author Kok, Philippe J. R. text Zootaxa 2009 1980 16 28 journal article 10.5281/zenodo.185278 d3b44fed-b15d-4dec-8bc5-328900976607 1175-5326 185278 Centrolene gorzulae ( Ayarzagüena, 1992 ) Centrolenella gorzulae Ayarzagüena 1992 : 19 . Centrolene gorzulai Duellman 1993 : 35 . Centrolenella auyantepuiana Señaris and Ayarzagüena 1995 “1993”: 122. Centrolene gorzulae Ruiz-Carranza and Lynch 1995 : 2 . Hyalinobatrachium auyantepuiana Ayarzagüena and Señaris “1996” 1997: 13. Cochranella auyantepuiana Myers and Donnelly 1997 : 16 . Cochranella auyantepuiana Duellman 1999 : 300 . Centrolene papillahallicum Noonan and Harvey 2000 : 295 . Centrolene lema Duellman and Señaris 2003 : 247 , new synonym . Centrolene gorzulai Myers & Donnelly 2008 : 29 . Holotype . MHNLS 11221 (adult male), Venezuela , Estado Bolívar: Central sector of Auyan-tepui ( 05°56’ N , 62°34’ W ; 1850 m ). Diagnosis. This species is placed in the genus Centrolene because adult males have humeral spines (Ruiz- Carranza & Lynch 1991 ; Duellman & Señaris 2003 ). It differs from all other species of Centrolene by having a trilobate liver covered by a white peritoneum and a mostly transparent parietal peritoneum, with a small white anterior portion (the “bib-like white patch” of Duellman & Señaris 2003 ), which is only detectable in dissected specimens. FIGURE 1. Dorsal and ventral view of Centrolene gorzulae from Sierra de Lema ( A , B ; CVULA 7013; photos by C. L. Barrio-Amorós) and Auyan-tepui ( C , D ; MHNLS 17325; photos by S. Castroviejo-Fisher). Characterization. (1) dentigerous process on vomer and vomerine teeth absent; (2) snout subtruncate to truncate in dorsal view, truncate to slightly sloping in lateral profile; (3) tympanum distinct, small, oriented dorsolaterally with a slight posterior inclination; (4) dorsal surfaces shagreened; males with small spicules visible under magnification; (5) ventral skin strongly granular, cloacal ornamentation consisting of small enameled tubercles lateral to vent and enlarged paired round tubercles below vent; (6) parietal peritoneum mostly transparent, a small anterior portion white (the “bib-like white patch” of Duellman & Señaris 2003 ), only detectable in dissected specimens; pericardial peritoneum white; hepatic and visceral peritonea white, urinary bladder transparent; (7) liver trilobate; (8) humeral spine in adult males present; (9) finger webbing II 2 -– 3 III 2+–(2-–2) IV ; (10) toe webbing I (1–1½)–(2–2+) II 1–2 + III 1 –(2–2+) IV (2–2+)– 1 V ; (11) fringe on postaxial edge of Finger IV present, enameled, metacarpal fold present, weakly enameled, ulnar fold absent; fringe on postaxial edge of Toe V present, enameled, metatarsal fold present, weakly enameled, tarsal fold usually absent or very weak; (12) nuptial excrescences finely granular, white, consisting of a dense group of glands and situated in the medial, dorsolateral internal side of Finger I (Type-V); prepollex slightly enlarged; prepollical spine projecting (spine not exposed); (13) when appressed, Finger II = Finger I ; (14) diameter of eye about 2X width of disc on Finger III ; (15) in life, dorsum dark green with scattered paler flecks; (16) in preservative, dorsum lavender entirely covered with minute melanophores; (17) in life, iris metallic copper with dark brown reticulations; an incomplete pale yellow ring around pupil; (18) distribution of melanophores on digits variable, but always more numerous on Fingers III–IV than I and II ; Toes IV–V with many melanophores, rarely present on Toes I–III , if present, few and restricted to the two distal phalanges; in life, hands and feet bluish green to green, tips of fingers and toes yellowish green; (19) males call from the upper side of leaves; note single and pulsed (3–7 pulses per call), advertisement call of 0.02– 0.05 s duration, dominant frequency of 4416.97–5157.48 Hz; (20) combat behavior unknown; (21) clutches deposited on vegetation overhanging streams, in moss or between two leaves, clutch size 15– 22 eggs ( n = 4); none of the clutches observed were guarded; (22) tadpole unknown; (23) SVL in males 19.2–22.5 mm ( n = 11); in females 20.9–22.0 mm ( n = 4). FIGURE 2. Ventral view of Centrolene gorzulae showing the distribution of iridophores on viscera. ( A ) MHNLS 17326 from Auyan-tepui; ( B ) MHNLS 17267 from Sierra de Lema . Photos by S. Castroviejo-Fisher. FIGURE 3. Audiospectograms (top) and oscillograms of the advertisement calls of Centrolene gorzulae . ( A ) MHNLS 17326 from Auyan-tepui; ( B ) MHNLS 17142 from Sierra de Lema . Air temperature at recordings ~ 19°. FIGURE 4. Neighbor-joining tree based on a 16S rDNA fragment. Support is indicated at the node when bootstrap values are> 70%. Terminals are labeled according to their species identification, distribution, and museum number (Gen- Bank accession number in the case of Allophryne ruthveni ). The topology supports the hypothesis that Centrolene lema is a synonym of C. gorzulae . Natural history and distribution. The ecology of Centrolene gorzulae was recently discussed by Kok and Castroviejo-Fisher (2008) , on the basis of specimens from Kaieteur National Park, Guyana . The species was recently discovered at a new locality, the southeast slope of Mount Maringma on the border of Guyana and Brazil ( ca. 1400 m elevation), and additional observations were made on clutch deposition. FIGURE 5. Egg clutch of Centrolene gorzulae deposited on moss, on a branch overhanging water ( ca . 2.0 m high). Photograph by P. J. R. Kok. Four clutches were collected between 23–24 November 2007 ; three of them were found among moss on overhanging branches 1.5–2.0 m above water ( Fig. 5 ), the fourth clutch was found deposited between two leaves (attached to both), about 1.5 m above water. Many males were calling at night in the vicinity of the clutch deposition sites, but no male-male interaction was observed. Clutches contained 15– 22 eggs at various stages of embryogenesis, and no guarding males were detected. One female (IRSNB 14327, 20.9 mm SVL), collected on 22 November 2007 , contained 13 enlarged, partly pigmented, ovarian eggs, the largest with a diameter of 2.0 mm. Centrolene gorzulae was thought to be endemic to Auyan-tepui ( Ayarzagüena 1992 ; Señaris & Ayarzagüena 2005 ; Myers & Donnelly 2008 ). Kok and Castroviejo-Fisher (2008) extended its distribution to Guyana . Here we report C . gorzulae from three localities in Venezuela (Auyan-tepui, Atapare, and Sierra de Lema ) and Guyana (Peters Mountain, Kaieteur National Park, and Mount Maringma) ( Fig. 6 ). This species probably occurs in adjacent areas of Brazil . Centrolene gorzulae has a broad altitudinal distribution (~ 450–1850 m ) and has been found only on vegetation overhanging small streams.