1 AMPHIBIA: ANURA: CENTROLENIDAE savagei

Catalogue of American and Reptiles 912

Vargas-Salinas, F., A. M. Ospina-L., J. A. Rios-Soto, and M. Rivera-Correa. 2017. .

Centrolene savagei (Ruiz-Carranza and Lynch) Savage’s Cochran

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 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 Lynch 1991; Vargas-Salinas et al. 2007; Var- A humeral spine is absent; the forearm is gas-Salinas et al. 2014; unpublished data).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. (2013) and is absence of nuptial excrescences. as follows, based on 238 calls recorded by 23 males. The call consists of 1–3 ‘peep’ notes PHYLOGENETIC RELATIONSHIPS. Cen- that last 16.65 ± 2.98 ms (range = 10–22 ms); trolene savagei was assigned originally to the consecutive notes are separated by silent in- by Ruiz-Carranza and tervals of 348.77 ± 33.67 ms (range = 302– Lynch (1991), based solely on morpholog- 442 ms). Mean call duration is 0.501 ± 0.26 ical characters such as the absence of a hu- s (range = 0.018–1.057 s). Call rate is 0.112 meral spine. Questions about the homology ± 0.03 calls/s (range = 0.061–0.198 calls/s). of the humeral spine and its implications for The dominant frequency of the call is 6.214 a monophyletic Cochranella, and the possible ± 0.334 kHz (range = 5.822–7.211 kHz). The synonymy with Centrolene were discussed by maximum sound pressure level of the adver- Frost et al. (2006). Subsequently, savagei was tisement call, measured at 50 cm in front of assigned to the genus Centrolene based on an two males, reached 75.4 and 78.6 dB, respec- 5 tively. A separate description of the courtship call in Centrolene savagei was published by Rios-Soto et al. (2017) and Vargas-Salinas et A al. (2014).

ILLUSTRATIONS. Color photgraphs of the venter of a female were published by Bolí- var-G. et al. (2013); color photographs of call- ing and non-calling males were presented by Bolívar-G. et al. (2013), Castro-Herrera et al. (2007), Díaz-Gutiérrez et al. (2013), Guayasa- min et al. (2009), Halliday (2016), Palacio Baena et al. (2006), Stuart et al. (2008); color photographs of amplexus and courtship be- havior were published by Vargas-Salinas et al. B (2014); color photographs of egg attendance by males were published by Bolívar-G. et al. (2013), Rivera-Correa (2013), Rojas-Mo- rales et al. (2011), and Vargas-Salinas et al. (2014). A color picture of a male in anti- predatory position behavior was published by Escobar-Lasso and Rojas-Morales (2012). A series of color drawings and photographs showing courtship behavior, amplexus, ovi- position, and parental care were presented by Vargas-Salinas et al. (2014). Black-and- white photographs of males caring for egg C clutches were presented by Vargas-Salinas et al. (2007). Black-and-white illustrations of tadpole morphology were published by Díaz-Gutiérrez et al. (2013) and black-and- white drawings and photographs of anatomi- cal features of adults were published by Pala- cio Baena et al. (2006) and Ruiz-Carranza and Lynch (1991).

DISTRIBUTION. Centrolene savagei is en- Figure 3. Oscillogram (A), spectrogram (B), demic to Colombia and is found on leaves and power spectrum (C) of the one note adver- between 0.5–5 m height alongside streams in tisement call of Centrolene savagei; dB = decibels; both undisturbed and disturbed forests. This kHz = kilohertz; ms = milliseconds ; SPL = sound species has been recorded between 1440 and pressure level. Snout-vent length of recorded male 2410 m in the central and western Andes of is 22.59 mm, temperature of male is 16.6º C. Re- Colombia in the departments of Antioquia, corded at Filandia, Central Andes of Colombia. Caldas, Quindío, Risaralda, Tolima, and Val- Adapted from Díaz-Gutiérrez et al. (2013). le del Cauca. Detailed data about elevational distribution range in each of those Colombi- an departments were published by Bolívar-G. 6

Figure 4. Development of embryos and egg attendance by the father. (A) Egg brooding after ovoposi- tion. (B) The father begins calling several days after ovoposition. (C) A male can stay beside the egg clutch until embryos are near to hatching. Note change in color of embryos. Photos by Fernando Vargas-Salinas. 7 et al. (2013). However, records in the munic- Dr. Jay M. Savage for his great contributions ipality of Falan, department of Tolima (Gal- to the knowledge about Centrolenidae. lego et al. 2008; Guayasamin et al. 2009) may correspond to another species (Marco Rada; COMMENTS. Experimental studies with personal communication). several species of Centrolenidae show that parental care in glass frogs is mainly to PERTINENT LITERATURE. Published ref- hydrate the eggs and reduce the mortality by erences to this species are listed by topic: ad- desiccation (Delia et al. 2013, 2014; Vocken- vertisement and courtship call (Díaz-Guti- huber et al. 2009). Given that the embryos érrez et al. 2013; Rios-Soto et al. 2017; are visible through their transparent-jelly egg Vargas-Salinas et al. 2014), antipredatory coverings, this hypothesis would be relatively behavior in adults (Escobar-Lasso and Ro- easy to test in Centrolene savagei. In this spe- jas-Morales 2012), breeding behavior, mat- cies, larger males have higher probablilities ing pattern, and parental care (Crump 1995; to mate than smaller males (Vargas-Salinas Ospina-L. et al. in press; Ruiz-Carranza and et al. 2014); the increased mating success of Lynch 1991; Vargas-Salinas et al. 2007, 2014; larger males seems most strongly correlated Wells 2007), conservation status (Bolívar et to a mechanism of endurance related to rival- al. 2004; Halliday 2016; Stuart et al. 2008), ry competition (intrasexual selection) rather geographic distribution (Acosta-Galvis than to female choice (intersexual selection) 2000, Bernal and Lynch 2008; Cadavid et al. based on egg fertilization efficiency or -pa 2005; Duellman 1993; Gallego et al. 2008; rental care quality (Ospina-L et al. in press). Palacio Baena et al. 2006; Rojas-Morales et al. Additionally, males are commonly observed 2011; Rueda-Almonacid 2000; Ruiz-Carran- attending egg clutches from ovoposition za and Lynch 1997; Ruiz-Carranza et al. 1996; until hatching (Vargas-Salinas et al. 2014). Stuart et al. 2008), morphology of adults and At present, it is unknown whether males of tadpoles (Cisneros-Heredia and McDiarmid Centrolene savagei adjust their parental effort 2007; Díaz-Gutiérrez et al. 2013; Ruiz-Car- according to environmental conditions as re- ranza and Lynch 1991), popular press books ported for other glass frogs (Delia et al. 2013, (Halliday 2016), , systematics, 2014), an important point in the context of and phylogenetic relationships (Castrovie- global climatic change. jo-Fisher et al. 2014; Frank and Ramus 1995; Frost 2017; Glaw et al. 1998, 2000a, 2000b; ADDITIONAL VERNACULAR NAMES. Guayasamin et al. 2008, 2009; Hutchins et Rana de cristal (Ospina-L. et al. in press; Var- al. 2003; Hutter et al. 2013; Pyron and Wiens gas-Salinas et al. 2014). 2011; Wrobel 2004). ACKNOWLEDGMENTS. Some of the dis- REMARKS. Unpublished genetic diversi- tributional records for the municipality of ty for mitochondrial markers in some pop- Salento, department of Quindío were provid- ulations of Centrolene savagei suggest that ed by Carlos M. Gómez. Geographical distri- there could be cryptic taxa within this spe- bution map was made with the help of Carlos cies (Marco Rada, personal communication). A. Londoño-G. Thanks to the journal Zoot- Future analyses that include different lines of axa for allowing us to replicate some images. evidence will help to elucidate the taxonomic limits of the species. LITERATURE CITED Acosta-Galvis, A. R. 2000. Ranas, Salaman- ETYMOLOGY. The specific epithetsavagei dras y Caecilias (Tetrapoda: Amphibia) de is a tribute to the recognized herpetologist Colombia. Biota Colombiana 1:289–319. 8

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FERNANDO VARGAS-SALINAS, Gru- po de Investigación en Evolución, Ecología y Conservación (EECO), Programa de Bi- ología, Universidad del Quindío, Armenia, Colombia ([email protected]), ANA MARÍA OSPINA-L., Grupo de Inves- tigación en Evolución, Ecología y Conser- vación (EECO), Programa de Biología, Uni- versidad del Quindío, Armenia, Colombia ([email protected]), JULIÁN A. RI- OS-SOTO, Grupo de Investigación en Evolu- ción, Ecología y Conservación (EECO), Pro- grama de Biología, Universidad del Quindío, Armenia, Colombia (rsoto.julian@gmail. com), and MAURICIO RIVERA-CORREA, Grupo Herpetológico de Antioquia, Instituto de Biología, Universidad de Antioquia, Me- dellín, Colombia (mauriciorivera79@gmail. com).

Primary editors for this account, Christopher J. Bell and Travis J. LaDuc.

Published 21 June 2017 and Copyright © 2017 by the Society for the Study of Amphibians and Reptiles.