767.1

AMPHIBIA: ANURA: LEPTODACTYLIDAE LEPTODACTYLUSMYSTACINUS

Catalogue of American Amphibians and Reptiles.

Heyer, M.M., W.R. Heyer, S. Spear. and R.O. de Si.2003. Lep- toc/actylus n~gstacinus. Leptodactylus mystacinus (Burmeister) Mustached Frog

Cystigncrthus mysracinus Burmeister 1861532. Type locality, "Bei Rozario" [Argentina, Santa Fe, Rosario, 32'57's 60°40'W, see Nomenclatural History]. Holotype, Martin- Luther-Universitat. Halle (Saale),no number, adult male,col- lector and date unknown (examined by WRH). Cysrignarhus labialis Cope 1877:90. Leptoduchlus my.s~acinus:Boulenger 1882:244. First use of currently recognized name.

CONTENT. The species is monotypic (see Nomenclatural History).

DEFINITION. Adult Lcptodacrylus mystacir~usare of mod- erate size, the head is as wide as long, and the hind limbs are moderately short (see Table: Heyer and Thompson 2000 pro- vided definitions of adult size and leg length categories for Lcptodnctylus). Male vocal sacs are not visible externally or at best are weakly expanded laterally and slightly darker than fe- male throats. Male snouts are more spatulate than those of fe- males. Male forearms are not hypertrophied. Males lack asperi- ties on the thumbs and chest. One or two pairs of dorsolateral I I I folds (indicated by darkllight outlining in indifferently preserved MAP. Di5tribution of L~~prodocr)~111.~~YSIOC~~US. The type loiality is specimens) are present: one distinct more dorsal pair extends indicated by a circle. DO^ mark-other localities. A dot may represent from behind theeye (often with a gap with the fold beginning at more than one site. Published locality data used to generate the map a level above the tympanum) to the upper groin: a second pair, should be considered as secondary sources of information. as we did either complete or interrupted, extends from above the forearm not confirm all specimen identities. insertion at the same level as the dorsal portion of the supratympanic fold and extends to the groin along the flanks. The toe tips are narrow. The toes lack fringes or fleshy ridges. The upper shank has many or scattered distinct white tubercles. The outer tarsus almost always (94%) has many or scattered distinct white tubercles. The sole of the foot usually (75%) has distinct scattered to many white tubercles,sometimes (25%) the white tubercles are absent. The upper lip usually (86%) has a distinct light stripe, some- times (14%) the stripe is indistinct. A dark interorbital bar is absent.Thedorsum is uniform,striped,or with small dark spots. The upper pair of dorsolateral folds have a distinct dark brown stripe or band ventrally and usually have a light pinstripe high- lighting the fold dorsally; the light stripe is often broader and more distinct posteriorly. The fold along the flanks may have dark or cream highlights or both. The species lacks light mid- of A. Olmos). dorsal stripes. The belly ranges from being immaculate to 3(1). The spiracle is sinistral and the vent tube is median. The mottled, with the mottling more intense on the lateral portions dorsal fin ends at the body and does not extend onto it. At Gosrler of the belly behind the arm insertions. The posterior thigh sur- stages 35-36, larval total lengths range from 4-8 mm and faces usually (94%) lack light horizontal stripes on their lower aspects; a few specimens (6%) have indistinct light stripes. The upper shanks have narrow dark crossbands, often interrupted TABLE. Summary measurement data for Leprodacrylus mysrtrcinus medially. (means are in parentheses). Larvae have a typical pond morphology and are members of Measurement Males Females the lentic. benthic guild (Altig and Johnston 1986, guild 12). The oral disk is anteroventrally positioned, entire (not emargin- SVL (rnm) 44-65 (54.1) 53.547.1 (57.3) ate), with an anterior gap lacking marginal papillae. A single Head IengthISVL (70) 33-39 (37) 32-38 (36) row of marginal papillae is on either side of the anterior gap; Head widthlSVL (%) 32-38 (35) 33-38 (35) two rows of marginal papillae occur laterally; one or two rows Thigh length/SVL (%) 3345 (39) 33-42 (38) occur ventrally, the second row usually not complete. No sub- Shank IengthJSVL (%) 3745 (42) 3W(42) marginal papillae are present. The tooth row formula is 2(1)/ Foot length/SVL (%) 36-47 (42) 3746 (42) body lengths from 17-18 mm; eye diameters are9-11% of body character descriptions of adults, including color patterns, are in lengths: and the widths of the oral disks are 21-23% of body Gallardo (1964~)and Hensel(1867,as L. mystacercs).Cei (1980), n lengths. The dorsum and sides of the body are brown: brown Cochran (1955).Heyer(1978),A. Lutz (1926),Mthely (1904), melanophores encroach on the venter anterior to the gut, other- F. Miiller (1880, as sp. questionably rnystacinus) and Sazima wise the venter lacks pigment. The tail musculature is mottled (1975) furnished morphological and color descriptions. Miranda- brown and cream. The tail fins range from having distinct dark Ribeiro's (1926) description of L. pygmaeus is not of L. to little mottling. mystacinus. Descriptions of juvenile L. mvsrcrcin~iswere pro- The advertisement call consists of a single note per call, given vided by Cope (1877, as Cystignathus labialis), Gallardo at a rate of 250JK)Wmin. Call duration ranges from 0.04-0.06 s (1964~).and Sazima (1975). Larval characteristics and color (Barrio's 1965b value of 0. I0 s seems to be due to over-record- are described in Sazima (1975) and Wogel et al. (2000). ing and/or microphone ringing). The call lacks or has negligible Wassersug and Heyer (1988) detailed larval oral features. Ad- amplitude modulation.The call lacks or has negligible frquency vertisement call descriptions are in Abrunhosa et al. (2001). modulation; when modulation is present, the average rise is 50 Barrio (1964b, 1965b). Kwet (2001). ancl Sazima (1975). Kwet Hz from beginning to end of call. The call attains near maxi- (2001) and Straneck et al. (1993) produced a compact disk and mum intensity almost immediately; the final third of the call cassette tape, respectively, that include L. my.stacinus advertise- falls offevenly.The dominant frequency is the fundamental fre- ment calls. A description of the karyotype is in Bogart (1974). quency. The dominant frequency ranges from 2050-2500 Hz. Harmonics are either absent (calls analyzed from Paraguay for ILLUSTRATIONS. Color photographs were provided by this account) or present (Barrio 1965b from Argentina). Couturier (l985), De la Riva et al. (2000), Kwet (2001), and Lavilla et al. (1995,as L. cf. my.stacinus). Photos of adult speci- DIAGNOSIS. The species having a combination of no toe mens are in Bogart (1974),Cei ( 1956), Freiberg (1972),Gallardo fringing, no light stripe on the posterior surface of the thigh, (1 964c, 1974), Klappenbach ( 1969), Martinez Achenbach and distinct white tubercles on the posterior surface of the tar- ( 1963). and Sazima ( 1975). Gallardo ( 1964c, 1987b) furnished sus are Leptodacrylus bufonius, L. labrosus, L. mystacinus, L. drawings of adult specimens. Illustrations by Cei (1980),A. Lutz troglodytes, and L. venrrinracnlztis. Leptodacrylus mj~stacinus (1926),and MChely (1904) are in color. Depictions of adult ana- has distinct dorsolateral folds (at least indicated by color pat- tomical features are in Cei (1980), Gallardo (1987b), Langone tern); dorsolateral folds are indistinct or lacking- in L. bufonius (1995). Limeses (1969). Limeses et al. (1972), and Sazima and L. rroglodvres. Lrprodacty1u.s mystacin~rsis difficult to di- (1975). Sazima (1975) presented an illustration of the head pro- agnose consistently from L. lubrosus and L. ventrimacularus file of a juvenile specimen. A larval photo is in Wogel et al. based on external morphological features. Most L. mystacinus (2000) and a SEM micrograph of the larval oral cavity is in have a very distinct light upper lip stripe; no L. labrosus or L. Wassersug and Heyer (1988). Drawings of larvae and larval ventrimaculatus have distinct upper lip stripes. The advertise- anatomical features were included in Cei (1980), Larson and de rn ment call of L. mystacinus differs markedly from those of L. Si (1998). Sazima (1975), and Wogel et al. (2000). Abrunhosa labrosus and L. ventrimacularus; the latter two species have calls et al. (2001), Barrio (1964b, 1965b), Kwet (2001), Salas et al. with pronounced rising frequency modulation (WRH, unpubl. (1 998), Sazima (1975), Straneck et al. (1993), and Wogel et al. data). (2002) provided audiospectrograms of the advertisement call. Abrunhosa et al. (2001) included an oscillogwm of the adver- DESCRIPTIONS. Burmeister (1861) (Spanish translation, tisement call and audiospectrograms and oscillograms of calls 1944) described the type specimen in detail and L. Miiller ( 194 I) of an unknown function. Photographs and illustrations of nests provided a morphological description of the holotype. Detailed and nest construction are in Sazima (1975). Bogart (1974) pro- vided a photograph of the karyotype.

DISTRIBUTION. Leptodactylus mystacinus is found in sub- tropical arid and rnesic habitats of the eastern slopes of the Andes in Bolivia and the Yungas of northwestern Argentina,extending eastward through Mato Grosso and Goiris to Bahia, Brazil. It ranges southward to central Argentina and Uruguay. Leptodac- rylus mystacinus occurs throughol~tthe Chaco and much of the FIGURE 2.Tadpole of Leptodac~lusniystocit~lrs (USNM 556038) from Cerrado Morphoclimatic Domain, but is not found in the Quinta de Santos, Montevideo, Uruguay; Gosner stage 37. Bar = 5 mm. Caatinga Morphoclimatic Domain (Ab7Sriber,1977). Where L. mystacinus occurs in mesic regions, as in the central and south- ern Atlantic Forests of Brazil, it inhabits clearings such as pas- tures rather than closed canopied forests. Harding ( 1983) listed its distribution by countries in the New World. Duellman (1999) tabled its distribution as Atlantic For- est, Caatinga-Cerrado-Chaco, and Pampean-Monte Domains. Citations for its general distribution are found in Cei (1980), Gallardo (1964~.1987b), Klappenbach and Langone (1992). Langone (1995). and Lavilla et al. (2000b). Boettger (1 892) re- corded a Leptorl~ictylusmystacinus collected from "Carricas, Venezuela" and Berg (1 896). Cei (1956). Freiberg (1942), A. Lutz (1927, 1928), Marelli (1924), and Nieden (1923) included n the mistaken locality in their distribution statements. Gallardo (1964~)indicated that the Venezuelan locality error was based FIGURE 3. Oral disk of Lcprodoctylus m~smrinuslarva (USNM on species misidentification. Beebe (1925. 1946), Crawford 556038) from Quinta de Santos, Montevideo. Uruguay; Gosner stage (193 l),and Crawford and Jones (1933) placed L.mysracinus in 37. Bar = l mm. British Guiana and Venezuela based on mistaken identifications. Vizotto and Castro 1964, as L. myscucin~is;Wogel et al. 2000); Paraguay (Aquino et al. 1996; Bertoni 19 14, 1939; Boettger 1885; Gallardo 1964~;MChely 1904; Scott and Lovett 1975); Uruguay (Achival and Olmos 1997; Borteiroet al. 1999; Bou- lenger 1882: Braun and Braun 1974: Canavero et al. 2001: Cei 1968 (M): Dei-Cas et al. 1976; Gallardo 1964~;Gonzilez 1997; Gonzilez et al. 1998: Gudynas 1980, 1989; Heyer 1978; Lagomarsino et al. 1988; Langgi~th1976: Langone 1995,1999; Maneyro et al. 1995: Meneghel 1992a: Prigioni and Langone 1984; Sierra et al. 1977).

FOSSIL RECORD. None.

PERTINENT LITERATURE. Gallardo (1964~)and Hensel (1867, as C. mj'staceus) published the most comprehensive works on the species. The following literature is listed by topic and the symbol (M) indicates a species mention. whereas (S) stands for a secondary source: bibliographic information and lists (Lavilla and Cei 2001, Liner 1992, Vizotto 1964b): bioge- ography (Baumann 1912; Braun and Braun 1980: Bridarolli and di Tada 1994b; Bucher 1980: Cei 1953. 1954, 1982: Cei and Pierotti 1957: CCspedez et al. 1995; Contreras and Contreras Time (seconds) 1982; Feio and Caramaschi 1995: Gallardo 1964a.b. 1965,1966, FIGURE 4. Wave form and ;~udiospectrogramdisplays of the adver- 1968a. 1972a.b. 1979,1985: Gudynas 1981: Heyer 1978.1988: tisement call of LPprodacty1~i.sn~j~smcinus (USNM recording 182. cut Heyer and Maxson 1982; Lavilla and Manzano 1995; Lynch 2) from El Tirol, near Encarnacibn. Ipapua. Paraguay. Upper wave form 197 1, 1979; Roig and Cei 196 I; Vellard 1948); call and call ofthe fifth call in the audiospectrogram: lower wave form of the begin- parameters (Abrunhosa et a1.200 1 ; Barrio 1964 a,b; Martinez ning of the upper wave form. Achenbach 1962,1963: Paillette's 1986 citation of L. mystcrcinits is L. niysfaceus: Salas et al. 1901. 1998; Straughan and Heyer 1976);catalogue accounts (Boulenger 1882; Giinther 1858, as In addition, the localities cited by Miranda-Ribeiro (1927) for C. schornhut:qkii; F. Miiller 1880. as C. sp.); conservation (Achh- n L. niysracinrrs, as L. p.ygmaeus, are contingent on a mix of spe- val et al. 1984; Anonymous 1997: Brand50 and Peres Jlinior cies (P. E. Vanzolini. pers. comm.). De la Riva et al. (2000), 200 1, as L. rnistucinus; di Tada et al. 1996; Gudynas 1989; Lago- Gonzales (1998). and Machado et al. (1999) collected the spe- marsino et al. 1988; Lavilla 2001 : Lavilla et al. 2000a; de Ledo cies in open formations. Sazima ( 1975), Bemarde and Kokubum et al. 1977; Maneyro and Langone 1999a.b. 2001; Sierra et al. (1999). and Striissrnann et al. (2000a) found it in gallery and 1977: Vaz-Ferreira and Achival 1983); ecology, natural his- primary forests. Finally, altitudinal limits were noted as 1200 m tory, reproduction (Abe and Garcia 1990: Anonymous 1877 by Barrio ( 1977) and 1800 m by di Tada et al. ( 1976). (S); Barbour 1926 (S); Barrio 1973; Bernarde and Kokubum The following references to distribution and localities are 1999; Blair 1976; Boettger and Pechuel-Loesche 1892 (S); listed by country; the letter (M) indicates a species mention: Boulenger 1886a; Brand50 and Araujo 200 1; Brandes and Argentina (Alvarez et al. 1996,2002; Barrio 1965b; Berg 1896; Schoenichen 1901 (S); Braun and Braun 1987; Bridarolli and Bosso and CCspedez 1995; Bucher 1980; Cei 1961 : Cei and di Tada 1994a; Cascon and Peixoto 1985 (M); Cei 1953 (S), Roig 1961 : CCspedez et al. 1995; Contreras 1982; Contreras 1954 (S); Cei and Roig 1961 ; Contreras and Contreras 1982; and Contreras 1982; Cruz et al 1992; di Tada et al. 1976,1996; Cope 1889 (S): Eisentraut 1932 (S); Fernindez and Fernindez Freiberg 1942: Gallardo 1961, 1964ac, 1965, 1966, 1968a.b, 1921 (S); Freitas et al. 2001 (S); Gadow 1901 (S): Gallardo 1972a, 1974, 1980, 1982, 1986, 1987a; Heyer 1978; Lajrnano- 1958 (S), 1964c, I965,1970,1972a, 1980,1987b. 1993; Garcia vich 1993: Laurent 1973, as L. mystacin~~s;Laurent and Terin 1972; Gonzhlez et al. 1998: Gudynas 1989; Gudynas and de 1982; Lavilla and Scrocchi 1992,1999;Lavillaet al. 1992,1995, Lecin 1992; Gudynas and Gehrau 198 1; Gundlach 188 1 (S): as L. cf. mystacinus, 2000b; Limeses 1969; Limeses et al. 1972; Hensel 1867, as C. niystaceus; Hoffrnann 1873- 1878 (S), as C. Lobo 1995; Martinez Achenbach 1963,1965; Martori and Aun mystaceus; Klingelhoffer 1956 (S); Knauer 1878 (S), 1883 (S); 1995; Martori et al. 1999; Montanelli and Acosta 199 I ; Peltzer Kwet 1999; Langone 1999; Lavilla and Manzano 1995; Lavilla and Lajmanovich 1999; Peracca 1897; Perotti 1997. as L. cf. et al. 1995, as L. cf. myslacinus; Livezey and Wright 1947 (S); mysracinus; Richard and Belrnonte de Richard 1985; Ringulet A. Lutz 1927; B. Lutz 1947 (S); Machadoet al. 1999; Maneyro and Ararnburu 1957: Vellard 1948; Weyenbergh 1876; Widmer et al. 1995; Martinez Achenbach 1962,1963,1965; Martins 1988 1994; Yanosky et al. 1993); Bolivia (De la Riva 1990, De la (M)(S); Martori et al. 1983; Meneghel 1992b,as L. misracinns; Riva et al. 2000, Gallardo 1964c, Gans 1960, Gonzales 1998, Mertens 1929 (S); Milstead 1972, as L. pygrnaeus; Noble 1917 Gonzales and Reichle 2002, Heyer 1978, Kohler 2000, Lavilla (S), as L. rnystuceniis; Perotti 1994, 1997, as L. cf. mysrocinu.~; et al. 1996, Peracca 1897); Brazil (Abrunhosa et al. 2001; W. Peters 1877 (S); Rossa-Feres and Jim 1994,1996; Sampson Bernarde and Kokubum 1999; Boettger 1892; Boulenger 1882, 1900 (S), as C. niystaceus; Santana and Menin 1994 (S); Sazima 1885a, 1886b; Brand50 and Arat~jo200 1; Brand50 and PCres 1975; Spirandeli-Cruz 1985; Striissmann et al. 2000a; Vaz- Jlinior 2001, as L. mistaciniis; Braun and Braun 1976a,b, 1980; Ferreira 1984 (S); Vaz-Ferreira and Gehrau 1974, 1975 (M); p Cei 1968 (M): Cochran 1955: Cope 1885; Feio and Caramaschi Vellard 1947 (M), 1948; Werner 1912 (S); Wunder 1932 (S)); 1995,2002; Gallardo 1964~;Heyer 1978: Izecksohn and Car- evolution (Heyer 1969a, 1978; B. Lutz 1948, as L. mystaceus; valho e Silva 200 1 ; Kwet 1999,2001 ; Lema and Braun 1993; Lynch 1971); habitat (Brand20 and PCres Jlinior 2001, as L. A. Lutz 1924; Machado et al. 1995; Mainieri and Gayer 1996; mistncinus; Feio and Caramaschi 1995; Gallardo 1964c, 1982, Maxson and Heyer 1988; F. Miiller 1885; Muramatsu and Cruz 1987b; Gans 1960; Gonzales 1998; Gonzales and Reichle 2002; 1996 (M); Sazima 1975; Striissmann et al. 2000b: Vizotto 1964a; Gudynas 1980 (M), 1989; Hensel 1867, as C.mystaceus; Kohler 2000: Kwet 2001; Maneyro et al. 1995: Martinez Achenbach al. 1988, Langguth 1976,Maneyro et al. 1995,Meneghel 1992a, 1962,1963; Martori et al. 1999):karyotype(Bogat 1974, Heyer Sierra et al. 1977), "Ranita de bigotes" (Martinez Achenbach 1978 (S),Heyer and Diment 1974. Rada cie Martinez 1976 (M)): 1962). and "Rana Bigote" (Cei 1956) are Spanish variations of keys (Boulenger 1882, 1886b: Lavilla 1994b; Lavilla and the common name,"Mustached Frog" (Frank and Ramus 1955). Scrocchi 1992; Lavilla et al. 1992,1993:A. Lutz 1926: Nieden Other common names include "Pfeifer" (Boettger and Pechuel- 1923: Prigioni and Achival 1992): listsof specimens in indi- Loesche 1892), "RI" (Anonymous 1997, Brand50 and PCres vidual collections (Aquino et al. 1996; Avila and Martori 1991: Jlinior 2001, as L. mi.sracinus), "Ri dourada" (Braun and Braun Boettger 1892: Elter 198 I ;Gavetti and Andreone 1993; F. Miiller 1976b), "Rana" (Schulz 1944), "Rana Criolla" (Alvarez et al. 1880, as sp. questionably n~,y.smcinus,1885: Yanosky et al. 1993); 2002), "Ranita de Burmeister" (Freiberg 1972), "Schnurr- morphology (Altig and Johnston 1986: Barrio 1965a; Cei and bartpfeifer" (Boettger and Pechuel-Loesche 1892,Gliesch 1927, Roig I961 ; Echeverria 1995 (S): Gallardo 1987b: Heyer 1978; Werner 1912). "Sqnderbare Pfeifer" (Knauer 1878, 1883). and Langone and de S6 200 1 ;Larson and de S6 1998; Limeses 1968. ''YLI? (Guarani) (Alvarez et al. 2002). 1969; Limeses et al. 1972: Liu 1935a.b: Martinez Achenbach Beebe's (1946) citations of L. mysracinus and L. mysricenu.~ 1962,1963; Mertens 1929; Noble 1922: Ponssa 1999); nomen- [sic] from Guyana andlor Venezuela as snake prey items, re- clature and taxonomy (Berg 1896; Boulenger 1882; Cei 1980. peated by Michaud and Dixon (1989), are most certainly not L. 1987 (M); Cochran 1955; De la Riva 2000: Dubois and Heyer mystacinus and are probably L. mysmceus. 1992: Heyer 1974,1978,2002: Klappenbach and Langone 1992; Lavilla 1992; A. Lutz 1930; Peters 1877 (M)); non-technical ETYMOLOGY.The specific name is derived from the Greek accounts (Cuenot 1932; Freiberg 1954, as L. pygtnaeus, 1972; word mystax, meaning upper lip or mustache. Gadow's (1901) Gadow 1901; Gallardo 1974; Gliesch 1927; Klappenbach 1969; statement, "Its specific name refers to the dark brown stripe Maneyro et al. 1995: Santos 1942); parasite host (Baker and which runs from the tip of the mouth through the eye to the Vaucher 1984,1986; Cordero 1933, as L.pygmueus; Dei-Cas et tympanum," refers to the canthal stripe. We believe the specific al. 1976: Fabio 1985): physiology (Barrio 1965a, Cei and name refers to a distinct light stripe on the upper lip that occurs Erspamer 1965, Cei et al. 1967, Daly and Witkop 197 1 (S), on most individuals. Erspamer et al. 1986. Roseghini et al. 1986): relationships and systematics (Bogart 1974. Cei and Erspamer 1965, Cei et al. ACKNOWLEDGMENTS. Reuber BrandIo, JosC Langone, 1967, Gallardo 1964b. Heyer 1978. Heyer and Diment 1974, Esteban Lavilla, Oswaldo Peixoto, and P.E. Vanzolini provided Larson and de S6 1998, Lirneses 1969, Limeses et al. 1972, information on literature and localities. Rainer Guenther clari- Lobo 1995, Lynch 197 I, Maxson and Heyer 1988, Melin 1941 fied the status of museum specimens and JosC Rosado loaned (M), Montero 1986. Savage 2002 (M), Wassersug and Heyer material. Esteban Lavilla critically reviewed the manuscript.The 1988); species accounts (Ach6val and Olmos 1997; Cei 1980; Smithsonian Institution Library staff of the Natural History Cochran 1955: di Tada et al. 1976: Gallardo 1964c: Gallardo branch helped obtain difficult-to-find publications. Especially and Varela de Olmedo 1992; Heyer 1978: Klappenbach and helpful were Martha Rosen, Claire Catron, Ron Lindsey, Leslie Langone 1992: Langone 1995; A. Lutz 1926: Nieden 1923: Overstreet, Courtney Shaw, David Steere, and Wanda West. Widmer 1994, as L. mysmceus): species comparisons (Anders- Work on this account was supported in part by the Neotropical son 1906 (M): Baumann 19 12 (M), 19 17 (M): Gallardo 1964~; Lowlands Research Program, Smithsonian Institution (Richard Heyer 1969b (M): A. Lutz 1930 (M); Melin 1941 (M)); species P. Vari, Principal Investigator) and the National Science Foun- or taxonomic lists (Ach6val 1989; Boulenger 1885a,b: De la dation (award # 9815787) to ROdS and WRH. Riva et al. 2000: de Si 1983: Frost 1985; Glaw et al. 2000; Gorham 1963,1966,1974; Harding 1983: Knauer 1878,1883; LITERATURE CITED Lavilla 1994a: Lavilla and Cei 2001: A. Lutz 1924; Ringulet and Aramburu 1957). Abc,A.S. and L.S. Garcia. 1990. Altera~desde fluidos corpdreos na r5 Leprodnc1v1u.sfuscus durante a estivas50 (Anura, Leptodactylidae). NOMENCLATURAL HISTORY. Leptoclnctylus mystacinus Revista Brasileira de Biologia 50:243-247. was identified as C~~stignathusmystaceus, Leptodactylirs pyg- Abrunhosa, P.A.. H. Wogel, and J.P. Pombal, Jr. 2001. Vocali.~a$iiode nlaeus, or Cysti,pnathus schotnbnrgkii prior to its recognition as quarto espicies de anuros do Estado do Rio de Janeiro, sudeste do Brasil (Amphibia, Hylidae, Leptodactylidae). Boletim do Muscu a distinct species in 1861. The influential paper by Peters (1 873) Nacional, Nova SCrie, Zoologia, Rio de Janeiro (472):1-12. erroneously concluded that Hensel's L. mystaceus was not the Ab'S6ber.A.N. 1977. Osdominios morfoclim,lticos na AmCrica do Sul. same species as L. mystacinus. Primeira aproximaqiio. Geomorfologia 52: 1-23 + I map. Leptodacty1u.r labialis (Cope 1877) was considered to be the Achsval, F. 1989. Lista de especies de vertebrados del Umguay. Parte valid name for the taxon now recognized as L.f,.agilis (Brocchi 2. Anfibios, reptiles, aves y mamiferos. Facultad de Humi~nidadesy 1877) prior to 1978 (Heyer 1978,2002). Ciencias. Servicio de Publicaciones Doccntes Internas, Montevideo: Heyer ( 1978) was in error in stating that no locality data other 146. than Argentina existed for the holotype of C~~stignathusmysta- - and A. Olmos. 1997. Anfibios y Reptiles del Uruguay. Serie Fauna No. I, Montevideo, Uruguay. cinus. Other authors (Barrio 1965b, Cei 1980, Lavilla 1992) -, M.D. Meneghel,and A.R. Melgarejo. 1984 ("IYXO").Conservacidn correctly identified the type locality as Rosario, Argentina. de la fauna de vertebrados del Uruguay. 11 Reuni6n Iberoamericana Schulz (1944,Agregado 2, Pane B, Planilla I) gave the coordi- de Consewaci6n de Zo6logos de Vertebrados:573-583. nates for the Rosario published by Burmeister as 32'56'37" lati- Altig, R. and G.F. Johnston. 1986. Major characteristics of free-living tude (South) and 60°39'00" longitude West. tadpoles. Smithsonian Herpetological Information Sewice (67):1- Lavilla et al. (1995) referred to specimens from the Los 75. Colorados Biological Station in Salta, Argentina as L. cf. ~lvarez,B.B., J.A. CCspedez, M.L. Li0ns.A. Hemando,and R.Aguirre. mystacinus.The males call from above ground, low on tree trunks 1996. Herpetofauna de las Provincias de Corrientes,Chaco y Fomlosa (Argentina).Facena 12: 119-134. and are being evaluated as to whether the Chaco frogs represent -, R.H. Aguirre, J.A. CCspedez, A.B. Hemando, and M.E. Tedesco. a speciesdistinct from L. mptacin~~s(E.O. Lavilla, pers.comm.). 2002. Atlas de Anfibios y Reptiles de Corrientes, Chaco y Formosa (Argentina).I.Anuros,Cecflidos, Saurios,AnfisMnidos y Serpientes. REMARKS. "Rana de bigotes" (de Ledo et al. 1977, de SA Editorial Universitaria. Universitaria Nacional Nordeste (Corrientes). 1983, di Tada et al. 1976, Klappenbach 1969, Lagomarsino et Andersson, L.G. 1906. On batnchians from Bolivia, Argentina, and Peru,collected by Erland Nordenskiold 190 1-1902 and Nils Holmgren - and Dr. Pechuel-Loesche. 1892. BrehmsTierleben.AllgemeineKunde P 1904-1905. Arkiv f6r Zoologi 3:l-19 + 1 plate. desTierreichs.Die Kriechtiere und Lurche. BibliographischesInstirut, Anonymous. I877.The development of batrachians without metamorph- Leipzig und Wien. osis. Nature 15:49 1492. Bogart. J.P. 1974. A karyosystematic study of frogs in the genus Lepro- - . 1997. Aspectos ecol6gicos dos vertebrados terrestres e semi-aquiti- dacrjdus (Anura: Leptodactylidae). 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