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Home , Adenomera, Aglossa, Chacophrys, Leptodactylinae, Leptodactylus, Leptodactylus bufonius

905.1 AMPHIBIA: ANURA: bufonius

Catalogue of American and .

Schalk, C. M. and D. J. Leavitt. 2017. .

Leptodactylus bufonius Boulenger Oven

Leptodactylus bufonius Boulenger 1894a: 348. Type locality, “Asunción, .” Lectotype, designated by Heyer (1978), Museum of Natural History (BMNH) Figure 1. Calling male Leptodactylus bufonius 1947.2.17.72, an adult female collected in Cordillera, Santa Cruz, . Photograph by by G.A. Boulenger (not examined by au- Christopher M. Schalk. thors). See Remarks. Leptodactylus bufonis Vogel, 1963: 100. Lap- sus. sally positioned. Te oral disc is ventrally po- CONTENT. No subspecies are recognized. sitioned. Te tooth row formula is 2(2)/3(1). Te oral disc is slightly emarginated, sur- DESCRIPTION. Leptodactylus bufonius rounded with marginal papillae, and possess- is a moderately-sized of the es a dorsal gap. A row of submarginal papil- (following criteria established by Heyer and lae is present. Te spiracle is sinistral and the Tompson [2000]) with adult snout-vent vent tube is median. Te tail fns originate at length (SVL) ranging between 44–62 mm the tail-body junction. Te tail fns are trans- (Table 1). Head width is generally greater parent, almost unspotted (Cei 1980). Indi- than head length and hind limbs are moder- viduals collected from the Bolivian Chaco ately short (Table 1). Leptodactylus bufonius possessed tail fns that were darkly pigment- lacks distinct dorsolateral folds. Te tarsus ed with melanophores, especially towards contains white tubercles, but the sole of the the terminal end of the tail (Christopher M. foot is usually smooth. Mature males have a Schalk, personal observation). strongly sloped, spatulate snout. Sexually ma- Two advertisement calls have been doc- ture males lack both thumb and chest spines. umented in Leptodactylus bufonius. Adver- Te dorsum is grey and may contain black tisement call type 1 (Figure 2A) consists of a spots or irregular shaped splotches; black single note. To the human ear, the call sounds bars are present on the tibia and the venter is like a metallic “huinc, hunic” (Cei 1980). Call white. Tis species lacks a white stripe on its duration is approximately 0.5 seconds; calls/ posterior thigh and tarsus. A middorsal stripe minute, 38; maximum call energy, 2 kHz; and light lip stripe are absent. Males possess a average dominant frequency, 1.488 kHz, fre- pair of black bilateral gular spots, but a single quency range, 1.25–2.0 Hz. Advertisement vocal sac. Juveniles and adults have a similar call type 2 (Figure 2B) consists of three notes. color pattern. Call duration is 0.39 seconds with each note Larvae are exotrophic, lentic, benthic lasting 0.13 seconds; calls/minute, 16; max- guild members (Altig and McDiarmid 1999). imum call energy, 1.75 kHz; average domi- Te body is ovoid-shaped and color can vary nant frequency, 1.567 kHz, frequency range, from brown to a grayish color. Eyes are dor- 1.0–2.0 kHz. 905.2

Map 1. Distribution of Leptodactylus bufonius. Te lectotype is designated by the white circle. Tese locality data should be considered secondary sources because we did not confrm the identifcations of specimens from all localities.

DIAGNOSIS. Adult Leptodactylus bufonius Leptodactylus bufonius is smaller than lack toe fringes and dorsolateral folds, two (female SVL: 68.1– traits found in , Lep- 97.6 mm; male SVL: 65.4–94.3 mm; de Sá et todactylus labyrinthicus, Leptodactylus lati- al. 2014), Leptodactylus labyrinthicus (female ceps, , Leptodactylus SVL: 141.8±12.3 mm; male SVL: 148.9±16.3 lithonaetes, , Leptodac- mm; Heyer 1979), tylus rugosus, , Lepto- (female SVL: 88.0–117.0 mm; male SVL: dactylus turimiquensis, and Leptodactylus 94.2–109.7 mm; de Sá et al. 2014), Leptodac- troglodytes. However, Leptodactylus bufonius tylus myersi (female SVL: 78.9–112.9 mm; has a Chacoan distribution and is found in male SVL: 74.2–123.4 mm; Heyer 2005), and sympatry with Leptodactylus chaquensis, Lep- Leptodactylus turimiquensis (female SVL: todactylus elenae, , Lep- 122.4–128.0 mm; male SVL: 127.2–160.0 todactylus lacticeps, Leptodactylus latinasus, mm; Heyer 2005), but generally larger than Leptodactylus leptodactyloides, and Leptodac- (female SVL: 42.8 ± 3.1 tylus mystacinus. mm; male SVL: 42.7 ± 2.5 mm; Heyer 1978), 905.3

Leptodactylus fuscus (female SVL: 43.6 ± 4.4; male SVL: 42.8 ± 4.0; Heyer 1978), Leptodac- tylus latinasus (female SVL: 33.0 ± 1.9 mm; male SVL: 31.2 ± 1.7 mm; Heyer 1978), and Leptodactylus leptodactyloides (female SVL: 34.8–56.2 mm; male SVL: 28.3–47.9 mm; Heyer 1994). Distinct dorsolateral folds are lacking in Leptodactylus bufonius, but one pair of folds is present in Leptodactylus laby- rinthicus; one or two pair of folds are present in Leptodactylus elenae, Leptodactylus lepto- dactyloides, and ; three pair of folds are found in Leptodactylus fuscus, and four pair of folds are present in Leptodactylus chaquensis. Te dorsum of Leptodactylus bufonius contains dark spots or irregular shaped splotches, ofen black against a grey-colored dorsum. Te colors are duller compared to Leptodactylus laticeps, which contains a tile- like pattern of aposematic colors of black and red rectangles against white (in life). Lepto- dactylus bufonius lacks a white stripe on the posterior surface of the thigh, but this stripe is present in Leptodactylus elenae, Leptodac- tylus fragilis, Leptodactylus fuscus, Leptodac- tylus latinasus, and generally in Leptodactylus leptodactyloides. Longitudinal dorsal stripes are present in Leptodactylus chaquensis and Leptodactylus latinasus, but absent in Lepto- dactylus bufonius. A middorsal white stripe is sometimes present in Leptodactylus fuscus, but always absent in Leptodactylus bufonius. Te venter is solid white in Leptodactylus bu- fonius, but melanophores are almost always present on the venter of Leptodactylus lepto- dactyloides. White tubercles are present on the foot of Leptodactylus elenae, Leptodactylus fragilis, Figure 2. Audiospectrogram (top) wave form and Leptodactylus latinasus, but these are (center), and expanded waveform (bottom) of the generally absent in Leptodactylus bufonius, advertisement calls A) type 1 (recorded at 23:00h which tend to have a smooth foot. Te shank at 30.3°C ) and B) type 2 (recorded at 21:00h at of Leptodactylus lithonaetes has a few to many 27.9°C ) of Leptodactylus bufonius recorded by a black and/or white tubercles, but the shank SM2 Song Meter in Cordillera, Santa Cruz, Boliv- of Leptodactylus bufonius is smooth. Tumb ia. No voucher specimens. spines are absent in sexually mature males of Leptodactylus bufonius, but one pair of spines 905.4

Table 1. Summary measurements for adult cies group was provided by Ponssa (2008) in a specimens of Leptodactylus bufonius. Ranges of cladistic analysis using external morphology, trait/SVL proportions are presented with the osteology, larval morphology, and behavior; average values in parentheses. Individuals mea- Leptodactylus troglodytes was the sister spe- sured for this table were from museum specimens cies of Leptodactylus bufonius (Ponssa 2008). and captured-and-released individuals from A more recent phylogenetic analysis of molec- localities in the Gran Chaco of Bolivia. Abbre- ular and non-molecular characters by de Sá et viations: SVL = snout-vent length, HW = head width, HL = head length, TL = thigh length, SL al. (2014) also supports the monophyly of the = shank length, FTL = foot and tarsal length. fuscus species group. Leptodactylus bufonius is part of the basal clade in the fuscus species Measurement Males Females group that includes Leptodactylus troglodytes (n=38) (n=48) and Leptodactylus mystacinus as sister taxa SVL (mm) 44–57 (52) 47–62 (54) (de Sá et al. 2014). Tese three species com- HW/SVL (%) 33–38 (36) 33–40 (35) prise the sister clade to the remaining species in the fuscus group (de Sá et al. 2014). HL/SVL (%) 29–37 (36) 27–40 (32) TL/SVL (%) 30–45 (35) 27–44 (35) PUBLISHED DESCRIPTIONS. Descrip- SL/SVL (%) 37–48 (41) 38–46 (41) tions of Leptodactylus bufonius were provid- FTL/SVL (%) 47–67 (54) 46–60 (53) ed by Achenbach (1962), Andersson (1906), Berg (1896), Boulenger (1894a), Budgett (1899), Cei (1980), de Sá et al. (2014), Gal- are present in Leptodactylus labyrinthicus, lardo (1987), Gallardo and Varela de Olmedo Leptodactylus leptodactyloides, Leptodactylus (1992), Heyer (1978), Lutz (1930), Mertens lithonaetes, and Leptodactylus myersi; two (1929), Müller and Hellmich (1936), Nei- pair of thumb spines are present in Leptodac- den (1923), Norman (1994), Straneck et al. tylus chaquensis, Leptodactylus laticeps, and (1993), and Weiler et al. (2013). A description Leptodactylus syphax. of the tadpole was published by Cei (1980). Adult Leptodactylus bufonius are most Audiospectrograms and descriptions of the similar to Leptodactylus troglodytes that are call were provided by Heyer (1978), Heyer similar in size, and lack both toe fringes and Scott (2006), Straneck et al. (1993), and and distinct dorsolateral folds (Heyer 1978). Straughan and Heyer (1976). Márquez et al. However, these species are not sympatric; (2002) and Straneck (1992) provide record- Leptodactylus troglodytes occur in northeast- ings of the call. ern , whereas Leptodactylus bufonius is restricted to the Gran Chaco ecoregion. ILLUSTRATIONS. Color photographs of the adult were provided by De la Riva et al. PHYLOGENETIC RELATIONSHIPS. Lep- (2000), de Sá et al. (2014), Gonzales et al. todactylus bufonius is a member of the fuscus (2006), Heyer and Scott (2006), Lavilla et species group, which is the largest species al. (1995a), Márquez et al. (2002), Norman group of the fve recognized species groups (1994), Pansonato et al. (2011), Ponssa and of genus Leptodactylus (Barrio 1973; de Sá Medina (2016), and Weiler et al. (2013). A et al. 2014; Heyer 1969; Ponssa 2008). While color photograph of a metamorph being con- the fuscus species group has undergone revi- sumed during a predation event was pub- sions (Heyer 1978, 1998), Leptodactylus bufo- lished by Schalk (2010a). Color photographs nius has remained in the fuscus species group of stained skeletal features were published by since its initial designation by Heyer (1969). Vera and Ponssa (2014). Color photographs Support for the monophyly of the fuscus spe- of the nest were published by Crump (2015), 905.5

Faggioni et al. (2017), and Weiler et al. (2013). cies of anurans in the Gran Chaco ecoregion Black-and-white photographs of adults of South America. Oven are known can be found in Achenbach (1962), Bogart from southeastern Bolivia to northern Ar- (1974), Cei (1949a), Cei (1956), Pisanó et al. gentina, Paraguay, the Brazilian Chaco, and (1993), Reading and Jofré (2003); black-and- the northwestern Pantanal of Brazil. white photographs of the tadpole were pub- lished by Pisano´ (1965b), Pisano´ (1966), FOSSIL RECORD. None. and Pisano and Barbieri (1965). Black-and- white photographs of the nest were provid- PERTINENT LITERATURE. Relevant cita- ed by Cei (1949a), Cei (1956), Crump (1995, tions are listed by topic: biogeography (Álva- 2010), Eisentraut (1932), Pisano´ (1965b), rez et al. 1996; Alvarez et al. 2002; Blair et al. Pisanó et al. (1993), and Reading and Jofré 1976; Bridarolli and di Tada 1994; Cei 1955a, (2003). A black-and-white photograph of the 1957, 1980, 1982; Contreras and Contreras karyotype was published by Bogart (1974); a 1982; Duellman 1999; Gallardo 1966, 1979, black-and-white photograph of the chromo- 1985; Gonzales et al. 2006; Heyer 1988; Heyer some and sperm was published by Barbieri and Maxson 1982a, 1982b; Roig and Contre- (1950). Black-and-white photographs of the ras 1975), call and call parameters (Barrio prepollex of an adult and primary carpal car- 1965b; Cei 1980; de Sá et al. 2014; Eggermont tilages of a tadpole were published in Fabrezi 1988; Gingras and Fitch 2013; Heyer 1978; (2001). Pisano´ (1966) provided a black-and- Heyer and Scott 2006; Kerr 1950; Márquez white photograph cross-sections of the me- et al. 2002; Straneck 1992; Straneck et al. sonephros of the tadpole. A black-and-white 1993; Straughan and Heyer 1976), checklists drawing of the jaw musculature was present- and catalog lists (Álvarez et al. 1996, 2009; ed by Manzano et al. (2003). Color illustra- Alvarez et al. 2002; Aquino et al. 1996; Aqui- tions of adult Leptodactylus bufonius were no-Schuster et. al. 1991; Avila et al. 1998; Avi- published by Cei (1980), Te Encyclopedia la and Carrizo 2003; Berg 1896; Bertoni 1914, of (2004), and Te Encyclopedia of 1939; Boulenger 1894a, 1894b, 1898; Briguera Reptiles Amphibians & Invertebrates (2006a, et al. 2006; Brusquetti and Lavilla 2006; Bu- 2006b). Black-and-white illustrations of cher 1980; Budgett 1899; Cei 1955a, 1955b, adult Leptodactylus bufonius were published 1956, 1957, 1960; Cei and Roig 1964; Ces- by di Tada et al. (1976), Gallardo (1987), Hey- pedez and Klein 2002; Cespedez et. al. 1995, er (1978), Lutz (1930), Müller and Hellmich 2004; Contreras 1982; Cruz et. al. 1992; De la (1936), and Straneck et al. (1993). Black-and- Riva 1990; De la Riva et al. 2000; di Tada et al. white illustrations of the tadpole were provid- 1976, 1996; Doria et al. 2001; Duellman 1999; ed by Cei (1949a) and Cei (1980). Black-and- Elter 1981; Freiberg 1942; Gavetti and An- white illustrations of the musculature were dreone 1993; Glaw et al. 1998, 2000a, 2000b; published by Limeses (1964). Black-and- Godoy 1963; Gonzales 1998; Gonzales et al. white illustrations of the nest were provided 2006; Gorham 1963, 1966, 1974; Harding by Eisentraut (1932) and Norman (1994). 1983; Hutchins et al. 2003; Jofré et al. 2013; Black-and-white illustrations of the nesting Kacoliris et. al. 2006; Köhler 2000; Lanza et behavior were published by Faggioni et al. al. 2006; Laurent and Teran 1981; Lavilla and (2017) and a black-and-white illustration of Cei 2001; Lavilla and Scrocchi 1991; Lavilla the calling site was published by Straneck et et al. 1995a, 1995b, 2000a, 2000b; Manzano et al. (1993). al. 2004; Martori and Aun 1995; Müller and Hellmich 1936; Padial et. al. 2003; Pansonato DISTRIBUTION. Leptodactylus bufonius is et al. 2011; Peracca 1895 [and 2007a reprint], one of the most common and abundant spe- 1897 [and 2007b reprint]; Pérez Iglesias and 905.6

Natale 2013; Reichle 2004; Sanabria and Qui- 1998; Limeses 1964, 1968; Limeses et al. 1972; roga 2009; Schalk et al. 2013; Scott and Lovett Manzano et. al. 2003; Medina et al. 2013; Per- 1975; Soria Modesto and Noss 2000; Souza acca 1895 [and 2007a reprint]; Pisanó 1965a; et al. 2010; Vellard 1948; Vogel 1963, 1964a, Pisano´ 1966; Pisano and Barbieri 1965; 1964b; Weiler et al. 2013; Wrobel 2004; Ya- Pisanò and del Rio 1968; Pisanò and Rengel nosky et al. 1993; Ziegler 2002), conserva- 1967; Ponssa 2008; Ponssa and Medina 2016; tion (Briguera et al. 2006; Brusquetti and Ponssa and Vera Candioti 2012; Schalk 2016; Lavilla. 2006; Crump 2005; De la Riva and Sebben et al. 2007; Vera and Ponssa 2014; Vi- Reichle 2014; Duellman 1999; Lavilla 2001; ertel and Richter 1999), physiology and bio- Lavilla and Brusquetti 2010; Lavilla and Cei chemistry (Barrio 1965a; Bertini and Rathe 2001; Lavilla and Heatwole 2010; Lavilla et al. 1962; Cei and Erspamer 1965, 1966; Cei et al. 2000a, 2000b, 2004; Stuart et al. 2008; Wei- 1961, 1967; Duellman and Trueb 1986, 1994; ler et al. 2013), ecology (Blair 1970, 1976a, Erspamer et al. 1964; Fernández and Fernán- 1976b; Bucher 1980, 1982; Cei 1949a, 1953, dez 1975; McClanahan 1975; Montero 1986; 1955b, 1956, 1980; Cei and Roig 1964; Con- Rastogi et al. 2005; Shoemaker and McClana- treras and Contreras 1982; di Tada et al. 1976; han 1973; Warburg 1997; Wells 2007), popu- Duré and Kehr 2004; Gallardo and Varela de lar press books (Mattison 1987 [and reprints, Olmedo 1992; Heyer 1969; Jofré et al. 2010; e.g., 1989 and 1994]; Te Encyclopedia of Lutz 1969; Mares et al. 1977; Mattison 1987 Animals 2004), predators and parasites [and reprints, e.g., 1989 and 1994]; Perotti (Baker 1987; Baker and Vaucher 1984, 1986; 1997; Pisanó et. al. 1993; Ponssa and Medina Durette-Desset et al. 1985; González and Ha- 2016; Reading and Jofré 2003; Sanabria and mann 2006, 2014, 2015; Hamann et al. 2012a, Quiroga 2009; Schalk 2016; Schalk and Saenz 2012b; Heyer and Scott 2006; Hulse 1978; 2016; Schalk and Sezano 2014; Schalk et al. Lent et al. 1946; Schalk 2010a, 2010b, 2016; 2013; Schalk et al. 2017; Straneck et al. 1993; Schalk and Montaña 2012; Schalk et al. 2014; Vellard 1948), evolution (Blair 1970, 1976a, Scott and Aquino 2005; Senzano and Schalk 1976b; Emerson 1976; Heyer 1969), genetics 2013; Smales 2007; Vucetich and Giacobbe and karyotypes (Barbieri 1950; Bogart 1967, 1949), reproductive biology (Barbieri 1950; 1974; Brum-Zorrilla and Saez 1968; Faggioni Budgett 1899; Cei 1949a, 1949b; Crump 1995, et al. 2014; Green and Sessions 2007; Heyer 2005, 2010, 2015; Duellman and Trueb 1986, and Diment 1974; Maxson and Heyer 1988; 1994; Eisentraut 1932; Faggioni et al. 2011, Morescalchi 1979; Rabello 1970), keys (Berg 2017; Klingelhöfer 1956; Lavilla et al. 2000b; 1896; Cei 1980; Heyer 1978; Lavilla et al. Pérez Iglesias and Natale 2013; Perotti 1994, 1993; Nieden 1923; Weiler et al. 2013), larvae 1997; Philibosian et. al. 1974; Pisanó 1962, and larval characteristics (Cei 1980; Crump 1965a; Pisano´ 1965b; Pisano and Barbieri 2015; De La Riva 1995; de Sá et al. 2014; Du- 1965; Pisanó et al. 1993; Rastogi et al. 2005; ellman and Trueb 1986, 1994; Faivovich and Reading and Jofré 2003; Ridley 1978; Schalk Carrizo 1992; Langone and de Sá 2005; Larson 2016; Schalk and Saenz 2016; Schalk et al. and de Sá 1998; McClanahan 1975; Motte et 2013; Shoemaker et. al. 1973; Weiler et al. al. 2004; Pisanó 1965a; Pisano´ 1965b; Pisanò 2013; Wells 2007), and , systemat- and del Rio 1968; Pisanò and Rengel 1967; ics, and phylogenetics (Barrio 1973; Cei Schalk 2016; Schalk et al. 2017; Shoemak- 1987; Cei and Erspamer 1965, 1966; Cei et al. er and McClanahan 1973; Vera and Ponssa 1967; De la Riva 1996; de Sá et al. 2005, 2014; 2014), morphology and development (An- Frank and Ramus 1995; Frost 1985, 2017; dersson 1906; Fabrezi 1992, 1996, 2001, 2012; Heyer 1978, 1995, 1998; Larson and de Sá Fabrezi and Barg 2001; Goin 1959; Hayek and 1998; Lavilla 1992; Limeses et al. 1972; Max- Heyer 2005; Heyer 1978; Larson and de Sá son and Heyer 1988; Montero 1986; Ponssa 905.7

2008; Sokolov, 1988; Zhao et al. 1998). ACKNOWLEDGEMENTS. We thank C. Vanek for assistance with the literature search ADDITIONAL VERNACULAR NAMES. and M. Jansen for creating the call graphs and Rana de los hormigueros (di Tada et al. 1976), providing constructive comments on the call Vizcacheras’ White-lipped Frog (Faggioni et description. We also thank W. R. Heyer for al. 2011; Te Encyclopedia of Animals 2004; providing constructive comments on this ac- Te Encyclopedia of Reptiles Amphibians & count. Support for CMS was provided by the Invertebrates 2006a, 2006b). NSF-GRF program and the Applied Biodiver- sity Sciences NSF-IGERT program at ETYMOLOGY. Tere were no comments A&M University. Tis is publication number on the etymology of the species in the origi- 1498 at the Research and Teach- nal description (Boulenger 1894a). Te word ing Collections at Texas A&M University. bufo is Latin for toad, and Budgett (1899) stat- ed the species name “bufonius” refers to the LITERATURE CITED presence of the spots. Presumably Leptodac- Achenbach, G. M. 1962. Nota sobre leptodac- tylus bufonius is considered more toad-like as tilidos de la zona del Rio Salado en el De- compared to other species of Leptodactylus, partmento La Capital de La Provincia de referring to the small wart-like bumps pres- Santa Fe. Anales del Museo Provincial de ent on the dorsum. Ciencias Naturales “Florentino Ameghi- no” 1:41–47. REMARKS. In the original description, there Altig, R. and R. W. McDiarmid. 1999. Diversi- was no holotype designated by Boulenger ty. Familial and generic characterizations. (1894a). Boulenger (1894a) reported the SVL Pp. 295–337 in Tadpoles. Te Biology of measurement of the specimen in his descrip- Anuran Larvae (R.W. McDiarmid and R. tion as 48 mm. Parker (1931) reexamined the Altig, eds.). Te University of Chicago type series (n = 4) and reassigned the smallest Press, Chicago, Illinois. two of the series to Leptodactylus diptyx. Te Álvarez, B. B., J. A. Cespedez, M. L. Lions, A. four syntypes collected by Boulenger (1894a) Hernando, and R. Aguirre. 1996. Herpe- were examined by Heyer (1978). Te frst tofauna de las Provincias de Corrientes, specimen of the series, BMNH 1947.2.17.72, Chaco y Formosa (). FACENA: had an SVL of 46.4 mm, the closest measure- Revista Científca Periódica de la Fac- ment to Boulenger’s (1894a) description, and ultad de Ciencas Exactas y Naturales y is likely the specimen upon which Bouleng- Agrimensura de la Universidad Nacional er’s description was based. Tis specimen was del Nordeste 12:119–134. designated as the lectotype by Heyer (1978). Alvarez, B. B., R. H. Aguirre, J. A. Cespedez, A. B. Hernando, and M. E. Tedesco. 2002. SPECIMENS EXAMINED. All Leptodactylus Atlas de Anfbios y Reptiles de las Pro- bufonius specimens examined were collected vincias de Corrientes, Chaco y Formosa, in Bolivia and are housed in the Herpetology Argentina. I. (Anuros, Cecilidos, Sauri- Collection in the Museo de Historia Natural os, Anfsbenidos, y Serpientes). Editorial Noel Kempf Mercado (MNKA): MNKA 399, Universitaria de la Universidad Nacional 452, 505, 1363, 1365, 1365, 1366, 1367, 1370, del Nordeste, Corrientes, República Ar- 1987, 4646, 4655, 4656, 5239, 5558, 5559, gentina. 156 pp. 5560, 5612, 5613, 5614, 5615, 5644, 5645, Álvarez, B. B., J. A. R. García, J. A. Céspedez, 5694, 5907, 7537, 7571, 8483, 8484, 8507, A. B. Hernando, V. H. Zaracho, C. C. 8526. Calamante, and R. H. Aguirre. 2009. 905.8

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Christopher M. Schalk, Department of Bio- logical Sciences, Sam Houston State Univer- sity, Huntsville, Texas 77341, USA (cschalk@ tamu.edu) and Daniel J. Leavitt, Arizona Fish and Game Department, Phoenix, Arizo- na 85086, USA ([email protected]).

Editors for this account, Christopher J. Bell and Travis J. LaDuc.

Published 2 May 2017 and Copyright © 2017 by the Society for the Study of Amphibians and Reptiles.