1 AMPHIBIA: ANURA: CENTROLENIDAE Centrolene savagei Catalogue of American Amphibians and Reptiles 912 Vargas-Salinas, F., A. M. Ospina-L., J. A. Rios-Soto, and M. Rivera-Correa. 2017. Centrolene savagei. Centrolene savagei (Ruiz-Carranza and Lynch) Savage’s Cochran Frog Cochranella savagei Ruiz-Carranza and Lynch 1991:8. Type locality: [Colombia,] "De- partamento del Quindío, municipio de Filandia, vereda El Roble, bosque Reser- Figure 1. Male of Centrolene savagei from La Reserva Nacional Forestal Bosque de Yotoco, in va Bremen, vertiente occidental, Cordil- the municipality of Yotoco, department of Valle ° ° lera Central, 4 39' Latitud N, 75 40' W del Cauca, Colombia. Photo by Fernando Var- de Greenwich, 2050 m.” Holotype, Museo gas-Salinas. de Historia Natural, Universidad Nacion- al de Colombia, Bogotá, ICN 9769, adult female, collected by P. M. Ruiz-Carranza cal external apertures; no depression present on 3 June 1981. between nostrils. Canthus rostralis undefined Centrolene savagei: Guayasamin, Castrovie- and concave; small eyes directed anterolater- jo-Fisher, Trueb, Ayarzagüena, Rada, and ally with an angle of 45 degrees. Eyes do not Vilà 2009:26. protrude to the labial edges from a ventral perspective. Anteroposterior diameter of the CONTENT. No subspecies are recognized. eye is equivalent to 26–36% of the cephalic length (mean: 31%, SD: 2.272); width of the DEFINITION. Individuals of this small glass superior eyelids 33–50% of the distance be- frog species exhibit a body size (snout-vent tween the eyes (mean: 45%, SD: 4.969). The length, SVL) between 19.0 and 24.2 mm with tympanic membrane is large and conspicu- adult males smaller than adult females (mean ous, with the supratympanic fold hiding the SVL ± standard deviation [SD] for 90 males: posterior margin of the tympanic ring; the 21.6 ± 1.2 mm, range: 19.0–24.2 mm; mean diameter of tympanic ring is 54–76% of the SVL ± SD for six females: 24.09 ± 1.11 mm, ocular diameter (mean: 60%, SD: 6.337). The range: 23.3–26.3 mm) (Ruiz-Carranza and medial vocal sac is external and subgular. Lynch 1991; Vargas-Salinas et al. 2007; Var- A humeral spine is absent; the forearm is gas-Salinas et al. 2014; unpublished data). The thin with a dermal fold and row of tubercles following description is based on 17 individ- extending until the outer margin of Finger IV. uals (two males, 15 females): head wider than The distal edge of the finger discs are trun- larger and as wide as the body; head width cate. Fingers I and II are of equal length. Web- is 31.0–37.3% of the SVL (mean: 34.2%, SD: bing is absent between Fingers I and II, vesti- 1.747). Head with a short face, rounded in gial between Fingers II and III, and moderate both dorsal and lateral views; the distance between Fingers III and IV. The hind limbs between eye and nostril is 66.7–95.0% of the are thin; metatarsal tubercle is ovoid and dif- anteroposterior diameter of the eye; loreal ferentiated, being twice as long as it is wide. region is straight and widening toward the There is a row of tubercles along a dermal fold superior lip. Nostrils prominent with elipti- in the external edge of the tarsus that extends 2 COLOMBIA Map. Known geographic distribution of Centrolene savagei. Red pentagon indicates the type locality, white dots represent additional localities. 3 to the base of Toe IV. Discs at the top of toes mm. Tail length is 23.3 mm (70.4% of total are smaller than those on hands. The outlines length). Myotomes are visible across the tail. of the discs are truncated to rounded. The medial line is straight and visible, sepa- Dorsal skin smooth with small white, rating the dorsal and ventral myotomes. Tail rounded warts in the trunk and the flanks of muscle width is 2.9 mm; tail muscle height the head. Skin on venter and ventral surfac- is 3.0 mm. The dorsal fin begins at the tail- es of the thigh is granular. The cloacal fold is body joint, increases in size at the middle of short, in horizontal position, and located at a the tail, and gradually tapers to a rounded tip. higher level than the thigh. Granular sub-clo- The ventral fin originates almost at the base of acal area with white granules. Individuals the tail muscle in the posterior body region; it are pale green with white, yellow, and green is slightly arched, and reaches its maximum spots; small blue blotches are usually present height throughout the distal two-thirds of the as well. Cloacal warts and tarsal tubercles are tail. Dorsal fin depth is 1.3 mm; ventral fin white. The upper lip has a white line. The iris depth is 1.1 mm. is gray-yellowish with dark brown reticles. The oral disc of the tadpole is directed an- The ventral area exhibits a white color in the teroventrally and is not emarginated; oral disc parietal peritoneum that extends almost until the liver. The large intestine and pericardi- um are covered by a white pigmentation. The heart is not visible. Bones are pale green. The tadpole was described by Díaz-Guti- érrez et al. (2013) based on an individual in developmental stage 39 (Gosner 1960). In that individual, the body is elongate and depressed (wider than high) in lateral view and oval in dorsal view. The total length is 33.1 mm; the body length is 9.8 mm (33.7% of total length), body width is 5.9 mm, and body height is 4.8 mm. Chondrocranial ele- ments are not visible. The snout is truncated in dorsal view and rounded in lateral view. A lateral-line system with neuromasts is evi- dent, mainly in the anterior part of the body, infraorbital line, angular line, and mid-lateral line. The eyes are positioned dorsolaterally; eye diameter is 1.1 mm; interorbital distance is 1.3 mm; eye-snout distance is 3.0 mm. Nar- ial apertures are small, elliptical, and dorsally directed. The distance between nostrils is 2.1 mm. The spiracle, at the posterolateral region Figure 2. Lateral (A), dorsal (B) and ventral (C) views of the tadpole of Centrolene savagei at of the body, is short and sinistral; diameter Gosner stage 39 (Gosner 1960); Museo de Her- of spiracle aperture is 0.7 mm. The spiracle petologia Universidad de Antioquia, Medellín is located slightly below the body midline at [MHUA]-L 0197–1, total length = 33.1 mm); oral 61.6% of the body length. The vent tube is disc at stage 39 (D) and at stage 26 (E). Illustration short and medial, free posteriorly, and opens by Mauricio Rivera-Correa. Reprinted with per- posteriorly; the cloacal tube length is 0.8 mission of the journal Zootaxa. 4 width is 3.0 mm (50.8% of the body width). extensive revision of the systematics of Cen- There are marginal and uni-serial papillae trolenidae using nuclear genes (c-myc exon distributed around the oral disc, with the lon- 2, RAG1, POMC) and mitochondrial genes gest papillae mainly on the lower labium; five (12S, 16S, ND1) (Guayasamin et al. 2009). submarginal papillae are present on each side Centrolene savagei is morphologically simi- of the jaw sheath (one papilla with a diminu- lar to Centrolene daidaleum (Ruiz-Carranza tive tooth row); marginal papillae are inter- and Lynch 1991); the two species were re- rupted dorsomedially in the anterior labium, covered as sister species in different phylo- forming a wide gap. The upper jaw sheath is genetic hypotheses constructed with molec- completely keratinized with a serrated edge, ular evidence (Castroviejo-Fisher et al. 2014; and with a pronounced arch in the middle. Catenazzi et al. 2012; Dugo-Cota et al. 2015; the lower jaw sheath is keratinized, V-shaped, Guayasamin et al. 2008, 2009; Hutter et al. and with a serrated edge. Broad-based serra- 2013; Pyron and Wiens 2011). tions are short and oriented straight medially. Labial tooth row formula is 1(1)/2(2); tooth PUBLISHED DESCRIPTIONS. Descrip- row A-1 is interrupted medially and is locat- tions of Centrolene savagei and taxonomic re- ed laterally on both sides of the upper jaw views of the family Centrolenidae were pub- sheath, forming a wide gap. lished by Cisneros-Heredia and McDiarmid (2007), Guayasamin et al. (2008, 2009), and DIAGNOSIS. Diagnostic characters of Cen- Ruiz-Carranza and Lynch (1991). Observa- trolene savagei are: presence of vomerine tions of breeding behavior, mating pattern teeth; individuals with pale green bones in and parental care were presented by Var- life; extension of the parietal peritoneum al- gas-Salinas et al. (2007, 2014). Antipredatory most to the liver; pale green coloration with behaviors were described by Escobar-Lasso white and green spots; webbing formula in and Rojas-Morales (2012). The tadpole, ad- fingers II (2+–2½)–(3⅓–3⅔) III (2⅓–2½)– vertisement call, and courtship call of Centro- (2–2⅓) IV; webbing formula of toes I (1⅓– lene savagei were described by Díaz-Gutiér- 1½)–2+ II 1–(2–2⅓) III (1–1½)–(2–2⅓) IV rez et al. (2013), Rios-Soto et al. (2017), and (2–2⅓)–1 V; rounded snout when viewed Vargas-Salinas et al. (2014). Spanish language dorsally; smooth dorsal skin with low, round- species accounts were published by Bolí- ed, and white warts; a thin row of tubercles var-G. et al. (2013) and Palacio Baena et al. along the outer margins of the hand, forearm, (2006). tarsus, and foot; absence of a humeral spine; The advertisement call was described in presence of a large and rounded tympanum; detail by Díaz-Gutiérrez et al.