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2003 Leptodactylus mystacinus M. M. Heyer
W. R. Heyer
Rafael O. de Sá University of Richmond, [email protected]
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Recommended Citation Heyer, M. M., W. R. Heyer, and Rafael O. de Sá. "Leptodactylus Mystacinus." Catalogue of American Amphibians and Reptiles, 2003, 767.1-67.11.
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AMPHIBIA: ANURA: LEPTODACTYLIDAE UPTODACTYWS MYSTA.CINUS
Catalogue or American Amphibians and Reptiles.
Heyer,M.M ., W.R. Heyer.S. Spear,and R.0.deSa.2003. up todacry/us m.vstacinus. Leptodactylus mystacinus (Burmeister) Mustached Frog
Cystigna1lms mys1aci11us Bunneisler 1861 :532. Type localiry, I "Bei Rozario" [Argentina, Santa Fe, Rosario, 32°57'S (J() 0 40'W. see Nomenclatural History). Holotype. Mnnio Lulher-Univel'llitll1. lialle (Saale), no number. adult ma.le, col· leclor and date unknown (examined by WRH). Cystlg11ath11s labia/is Cope 1877:90. leptodactylus mystacinus: Bouleoger 1882:244. Finn use of currently recognized name.
• CONTENT. The species is mono1ypic (see Nomcnclatural History).
• DEFINITION. Aduh leptodactylus 111ystad1111.T are of mod erale size, the head is as wide as long, and the Mod limbs are moderalely short (!U Table; Heyer and Thompson 2000 pro vided defini1 ions of adult size and leg length categories for Lcptodacry/11.s) . Male vocal sacs are not visible externally or a1 best are weakly expanded laterally and slightly darker lhrut fe· male 1hroa1S. Male snouts :ue more spatulate than those of fe m3Jes. Male forearms are not hypertrophied. Males lack asperi- 0 6CIO le• 1ies on the thumbs and chest. One or two pairs of dorsoJaternl I. t I I I folds (indiCDted by darlc/light outlining in indifferently pre.served MAP. Distribution of l.Lptodacry/u:s 111ys1oci111a. 1be rype localiry is specimens) are present: one distincc more dorsal pair extends Indicated by a circle. Dool marlc other localities. A dot may represent from behind the eye (often with a gap with the fold beginning at more thnn one ite . Published locality dalll U.'ICd to geocrete the map a level above the tympanum) to the upper groin; a second pair. $houJd be considered as secondary wun:cs of information. as we
body lengths from 17- 18 mm; eye diameters are9-l 1%ofbody character descriptions of adu lts, including color pauerns, are in lengths; and the widths of the ocal disks are 21- 23% of body GaUardo (l 964c) and Hensel (1867 ,as L. mystaceus). Cei (1980), lengths. The dorsum and sides of the body are brown; brown Cochran ( 1955), Heyer ( 1978). A. Lutz ( 1926), Mchel y ( 1904), melanopbores encroach on the venter anterior to the gut, other F. MUiier (1880, as sp. questionably mystacinus) and Sazima wise the venter laclcs pigment. The tail musculature is mottled ( 197 5) furnished morphological and color descriptions. Miranda browi4 and cream. The tail fins range from having distinct dark Ribeiro 's ( 1926) description of L. pygmaeus is not of L. to little motWng. mystocinus. Descriptions of juvenile L. mystacinus were pro The advertisement call consists of a single note per call, given vided by Cope ( 1877, as Cystignathus labialis), Gallardo al a rate of250-400/min. Call duration ranges from 0.04--0.06 s (1964c), and Sazima (1975). Larval characteristics and color (Barrio's 1965b value ofO.lOs seems 10 be due lo over-record arc described in Saiima (1975) and Wogel et al. (2000). ing and/or microphone ringing). The call lacks or has negligible W11sse.rsug and Heyer (1988) detailed L1rval oral features. Ad amplitude modulation. The call laclcs or has negligible frquency vertisement call descriptions are in Abrunhosa ct al. (2001), modulation; when modulation is present, the average rise is 50 Barrio (1964b, 1965b), Kwet (2001), and Sazima (1 975). Kwct Hz from beginning to end of call. The call attains near maxi (2001) and Straneck ct al. (1993) produced a compact disk and mum intensity almost immediately; the final third of the call cassette tape, respectively, that include L . mystadnus advertise falls off evenly. The dominant frequency is the fundamental fre ment calls.A description of the karyorypc i<; in Bogart (1974). quency. The dominant frequency ranges from 2050-2500 Hz. Harmonics arc either absent (calls analyzed from Paraguay for • ILLUSTRATIONS. Color photogruphs were provided by this account) or pre..;ent (Barrio l965b from Argentina). Couturier (1985), De la Riva ct al. (2000), Kwet (2001), and Lavilla ct al. (1995 , as L.cf.mystacinu.s). Photos of adult speci • DIAGNOSIS. The species having a combination of no toe mens arc in Bogart (1974),Cei (1956),freiber:g (1972),Gallardo fringing, no light stripe on the posterior surface of the thigh, (I964c, 1974), Klappenbach (1969), Martinez Achenbach and distinct white ntbcrcles on the posterior surface of lhe tar (1963), and Sazima (1975). Gallardo ( 1964c, 1987b) furnished sus are leptodactylus b11/onius, L. labrosus, L. mystacinus, L. drawings of adult specimens. lUustration.~ by Cci (1980),A. Lutz troglodytes, and L. ventrimaculatis. Leptodactylus mystacinus (1926),and MChely (1904) are in color. Depictions of adult ana has distinct dorsolatcraJ 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 ec al. (1972), and Sazima and L. troglodytes. Leptodactylus mystacinus is difficult to di (1975). Sazima (1975) presented an illustration of the bcadpr<> agnose consistently from L. lahrosus and L. ventrimaculatus file of a juvenile specimen. A larva l photo is in Woge! et al. ba~ on e.xtemal morphological features. Most L. mystacim1s (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. Was.~rsug and Heyer (1 988). Drawings of larvae and larval venirimaculatus have distinct upper lip stripes. The advertise analomical features were includec.I in Cei (1980), Larson and de ment call of L . my.~tacinus diITers mark:edJy from those of L. Sa ( 1998), Sazima (1975), and Woge! et al. (2000). Abrunhosa /abrosus and L. ventrimaculatus; the latier two species have calls et al. (2001), Harrio (J964b, 1965b), Kwet (2001), Salas et aJ. with pronounced rising frequency modulation (WRH , unpubl. (1998),Sazima (1975), Straneck ct al. ( 1993), and Woge! et al. daCa). (2002) provided audiospectrogr.uns of the advertisement call. Abrunhosa et al. (2001) included an o.~cillograrn of the adver •DESCRIPTIONS. Burmeister (1861) (Spani.;h translation, tisement call and audiospectrogr.ims and osci ll ogra.m~ of call~ 1944) described the type specimen in detail and L. MillJer ( 1941) of an unknown function. Photographs and illustrations of nests provided a morphological description of the holotype. Detailed and nestcon.~truction are in Sa7.ima ( 1975). Bogart (1974) pro vided a photograph of the karyotype.
• DISTRIBUTION. Leptodactylus mystacinus i~ found in sub tropical arid and rnesic habitats of the ca.~tcm slopes of Che Andes in Bolivia and the Yuogas of northwestern Argentina.extending eastward through Mato Grosso and Goias to Bahia, Brazil. It r.mges southward to central Argentina and Uruguay. Leptcdac tylus mystacinus occurs throughout the Chaco and much of the ~ · FIGURE 2. Tadpole of Leptodactylus mystacinus (USNM 556038) from Cerrado Morphoclimatic Domain, but is not fount.I in th e Quinta de Santos, Montevideo, Uruguay; Gosoer stage 37. Bar= 5 mm. Caatinga Morphoclimatic Domain (Ab'Slibcr, 1977). Where L. mystacinus occurs in mesic regions, as in the central and south ern Atlantic r'Orcsts of Brazil, it inhabit~ clearings such as pas tures rarhcr tl1an 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-Oiaco, and Parnpean-Monte Domains. Citations for its general distribution are found in Cei (1980), Gallardo (1964c, 1987b), Klappenbach and Langone (1992), Langone ( 1995), and Lavilla et al. (2000b). Boettger (1892) re corded a Leptodactylus mystacinus collected from "Caracas, Venezuela" and Berg ( 1896), Cei (1956), Freiberg (1942), A. Lutz ( 1927, 1928), Marclli (1924),and Nicden (1923) included the mistaken locality in their distribution statements. Gallardo w 11 1 !ii!llii (l 964c) indicated that the Venezuelan locality error was based · . --·on spec'1es m1"sidentification. Beebe ( 1925 1946) Craw~ord1 FIGURE 3. Oral disk of Lc}Uodac1yl11s mystacm11s larva (USNM ' • . ' • 556038) from-Quinta de Santos, Montevideo, Uruguay; Gosner suige - -(193 I), and Crawford and Jooes-(1933) P_laced L: mystacmus m - 3'7-:-Bar-= 1 mm. British Guiana and Venezuela-based on rrustaken identifications. 767.3
Virotto and Castro 1964, as L. myscacinus; Woge! et al. 2000); Paraguay (Aquino ct al. 1996; Bertoni 1914, 1939; Boettger 1885; Gallardo 1964c; M6hcly 1904; Scott and Lovett 1975); Uruguay (Acb Sms •FOSSIL RECORD. None. 6.0 ,....._ •PERTINENT LITERATURE. Gallardo (1964c) and Hensel :rN (1867, as C. mystace11s) published the most comprehensive ~ 4.0 works on the species. The following li terature is listed by topic g and the symbol (M) indicates a species mention, whereas (S) Cl stands for a secondary source: bibliographic information and ~ 2.0 ... ~- lists (La villa and Cei 200 I, Liner 1992, Vizotto 1%4b ); bioge ~ • ·• ..' • • • ' ography (Baumann 1912; Braun and Brauo 1980; Bridarolli and di Tada 1994b; Bucher 1980; Cei 1953, 1954, 1982; Cei 1.S 2.0 and Pierotti 1957; CCspedez.et al. 1995; Contreras and Contreras Time (seconds) 1982; Feio and Carnm.1schi 1995; Gallanlo 1964a,b, 1965, 1966, FlGURE 4. Wave fonu and audiospcctrogr.un displays of the adver 1968a, 1972a,b, 1979, 1985; Gudynas 1981; Heyer 1978, 1988; tisement call of vptodactylu.t mystocinus (USNM recording 182, cut Heyer and Maxson 1982; Lavilla and Manuno 1995; Lyoch 2) from El Tirol, near Encamaci6n, Ipapoa, Paraguay. Upper wave form 1971 , 1979; Roig and Cei 1961; Vellard 1948); caU and call of the fif1h call in the aucliospeclrOgram; lower wave form of the begin parameters (Abruobosa ct al. 200 l; Barrio I 964 a,b; Martinez ning of lhe upper wave fonn. Achenbach 1962, 1963; Pai llette's 1986 citation or L. mystacinus is L. mystaceus; Salas et al. 1991, 1998; Straughan and Heyer 1976); catalogue accounts (Boulcnger 1882; Gunther 1858, as In addition, lhe loca l iti~ cited hy Micanda-Ribeiro (1927) for C.schomb11rgkii; !:'.Muller 1880,as C.sp.); conservation (Acha L. mystacim's. as L. pygmaeu3, are contingent on a mix of spe val et al. 1984; Anonymous 1997; Brandiio and Peres Junior cies (P. E. Vanzolini, pcrs. comm.). De la Riva et al. (2000), 200 I , as L. mistacinus; di Tada ct al. 1996; Gudynas 1989; Lago Gontilles (1998), Md Machado ct al. (1999) collected the s~ marsino et al. 1988; Lavilla 2001; LaviJla ct aJ. 2000a; de Ledo cies in open fonnations. Sa.iirna ( 1975), Bemanle and Kokubum ct al. 1977; Maneyro and Langone 1999a,b, 2001; Sierra et al. (1999}, aod Strilssmann et al. (2000a) found it in gallery and 1977; Vaz-Ferreira and Ach6val 1983); ecoto,:y, natural his primary fo1es1s. 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; Bcmardc and Kokubum The following references to disuihution and localities are 1999; Blair 1976; Boettger and Pechuel-Locsche 1892 (S); listed by country; lhe letler (M) ind icates a species mention: Boulcnger 1886a; Brandlio and Araujo 2001; Brandes and Argentina (Alvarez et al. 1996,2002; Barrio J 965b; Berg 1896; Schoenichen 1901 (S); Braun and Braun 1987; Bridarolli and Bosso and ~pedez 1995; Bucher 1980; Cei 1961; Cei and di Tada 1994a; Cascon and Peixoto 1985 (M); Cei 1953 (S), Roig 1961; Cespcdez et al. 1995; Contreras 1982; Contrc.ras 1954 (S); Cei and Roig 1961; Contreras and Contreras 1982; and Contreras 1982; Cruz et al 1992; di Thda et al. 1976, 1996; Cope 1889 (S); Eisentraut 1932 (S); Femande'l. and Fernandez Freiberg 1942; Gallardo 1961, 1964a,c, 1965, 1966, 1968a,b, 1921 (S); Freitas et al. 2001 (S); Gadow 190 1 (S); Gallardo 1972a, 1974, 1980, 1982, 1986, 1987a; Heyer 1978; Lajmano 1958(S), 'l964c, 1965, 1970, 1972a, 1980, l987b, l993; Ga.rda vich 1993; Laurent 1973, as L. mystacinys; Laurent and Teriln 1972; Gomalez et al. 1998; Gudynas 1989; Gudynas and de 1982; LaviUa and Scrocchi 1992, 1999; Lavillact al. 1992, 1995, Le6n 1992; Gudynas and Gehrau 198 1; Gundlach '188 1 (S); asL.cf. mystacinu.s, 2000b; Limeses 1969; Limescs ct al. 1972; He.nsel I 867, as C. mystaceus; Hoffmann 187'.l- I 878 (S). as C. Lobo 1995; Martinez Achenbach 1963, 1965; Martori and Aun mystaceus; KJingelhoffer 1956 (S); Knauer 1878 (S). 1883 (S); I 995; Martori et al. 1999; Montanelli and Acosta 1991; Peltzer Kwet l 999; Langone 1999; La villa and Man1.ano 1995; Lavilla and Lajmanovich 1999; Peracca 1897; Perotti 1997, as L. cf. et al. 1995, as L. cf. mystacinus; Livezey and Wright 1947 (S); mystacinus; Richard and Belmonte de Richard 1985; Ringulet A. UltZ 1927; B. Lotz 1947 (S); Machado ct al. 1999; Maneyro andAramburu 1957; Vcllard 1948; Wcycnhergh 1876; Widmer et al.1995; Martfne-t.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 l 992b, as L. mistacinus; Riva ct al. 2000, Gallardo 1964c, Gans 1960, Gonzales 1998, MCTtens 1929 (S); Milstead 1972, asL.pygmaeus; Noble 1917 Gonzales and Reichle 2002, Heyer 1978, Kohler 2000, Lavilla (S), as f,. mystacen11S; Perotti 1994, 1997, as L. cf. mystacinus; ct al. 1996; Peracca 1897); Brazil (Abrunhosa ct al. 2001; W. Peters 1877 (S); Rossa~Fercs and Jim 1994, 1996; Sampson Bcmarde and Kokubum 1999; Boettger 1892; Boulcngcr 1882, 1900 (S),as C.mystaceu.r, Santana and Meoin 1994 (S); Sazima 1885a, 1886b; Brandao and Araujo 2001; Brandao and P~rcs 1975; Spirandeli-Cruz 1985; Strilssmann et al. 2000a; Vaz Jdoior 200 1, as L. mistacinus; Braun and Braun 1976a,b, 1980; Fcrreira 1984 (S); Vaz-Ferreira and Gehrau 1974, 1975 (M); Cci 1968 (M); Cochran 195.5; Cope 1885; Fcio and Caramaschi Vella.rd 1947 (M), 1948; Werner 1912 (S); Wunder 1932 (S)); 1995, 2002; Gallardo 1964c; Heyer 1978; lzecksohn and Car evolution (Heyer 1969a, 1978; B. Lutz 1948, as L. mystacellS; valho e Silva 2001; Kwet 1999, 2001; Lema and Braun 1993; Lynch 197 1); habitat (Brandiio and P6res J6nior 2001, as L. A. Lotz 1924; Machado ct al. 1995; Mainieri and Gayer 1996; mistacinus; Fcio and Caramaschi 1995; Gallardo I 964c, 1982, Maxson and Heyer 1988; F. MULier 1885; Muramatsu and Cruz I 987b; Gans 1960; Gonzales 1998; Gonzales and Reichle 2002; l 996 (M); Sazima 1975; Str!issmann ct al. 2000b; Vizotto I964a; Gudynas 1980(M), 1989; Hensel 1867,as C.mystaceus; K<>hlcr 767.4 2000; Kw el 2001; Maneyro et al. 1995; Martinez Achenbach al. 1988, Langguth 1976,Maneyroet al. 1995,Meneghel 1992a, 1962, J963; Marton et al. 1999); karyotype (Bogart 1974, Heyer Sierra et al. 1977), "Ranita de bigotcs" (Martinez Achenbach 1978 (S), Heyer and Dimenl 1974,RadadeMartinez 1976 (M)); 1962), and "Rana Bigote" (Cei 1956) arc Spanish variations of keys (Doulenger 1882, 1886b; Lavilla 1994b; Lavilla and ll1e common name, "Mustached Frog" (Frank a11d Ramus 1955). Scrocchi 1992; Lavilla et al. 1992, 1993; A. Lutz 1926; Nieden Other common names include "Pfeifer" (Boettger and Pechuel 1923; Prigioni and Achaval 1992); lists of specimens in indi Locsche 1892), "Ra" (Anonymous 1997, Brandiio and Peres vidual collections (Aquino et al. 1996; Avila and Martori 1991; Junior 2001, as L. mistacinus), "Rii dourada" (Braun and Braun Boettger 1892; Elter 1981; Gaveni and Andreone 1993: F. MUiier 1976b), "Rana" (Schulz 1944), "Rana Criolla" (Alvarez el al. 1880,as sp.questionably mysracinus, 1885; Yanosky et al. 1993); 2002), "Ranita de Burmeister" (Freiberg 1972), "Schnurr morphology (Altig and Johnston 1986; Barrio 1965a; Cci and bartpfcifer" (Boettger and Pechuel-Loesche 1892. Gliesch 1927, Roig 1961; Echeverria 1995 (S); Gallardo I 987b; Heyer 1978; Werner 1912), "Sonderbare Pfeifer" (Knauer 1878, 1883), and Langone and de Sa 200 I ; Larson and de Sa 1998; Li meses 1968, "Yur• (Guarani) (Alvarez ct al. 2002). 1969; Limeses et al. 1972; Liu 1935a,b; Martinez Achenbach Beebe's (1946) citations of L. mystacin11s and L. mysticenus 1962, 1963; Mertens 1929; Noble 1922; Ponssa 1999); nomen [sic) from Guyana and/or 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 mysracitius and are probably L. mystaceus. 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 (Cu~nol 1932; Freiberg 1954, as L.pygmaeus, 1972; word mysuu, meaning upper lip or mustache. Gadow's (1901) Gadow 1901; Gallardo 1974; Gliesch 1927; Kl appenbach 1969; statement, "lts 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. pygmaeus; Dei-Cas ct tympanum," refers 10 the canthal si:ripe. We believe the specific al. 1976; Fabio 1985); physiology (Barrio 1965a, Cei and name refers 10 a dist.im:.1 light stripe on the upper lip that occurs Erspamer 1965, C<:i et al. 1967. Daly and Witkop 1971 (S), on most individuals. Erspamer el al. 1986, Roseghini el al. 1986); relationships and system atics (Bogart 1974, Cei and Erspamer 1965, Cei et al. • ACKNOWLEDGMENTS. Reuber Brandiio, Jose Langone, 1967, Gallardo 1964b, Heyer 1978, Heyer and Diment 1974, Esteban Lavilla, Oswaldo Peixoto, and P.E. Vaozolini provided Larson and de Sli 1998, Limeses 1969, Limeses et al. 1972, infom1ation on literature and localities.'Rainer Guenther clari Lobo 1995, Lynch 1971, Maxson and Heyer 1988, McUn 194 I fied the status of museum specimens and Jose Rosado loaned (M), Montero 1986, Savage 2002 (M), Wassersug and Heyer material. Esteban Lavilla critically reviewed the manuscript. The 1988); species accounts (Achaval and Olmos 1997; Cei 1980; Smithsonian Institution Library staff of the Natural History Cochran 1955; di Tada et al. 1976; Gallardo 1964<:; Gallardo branch helped obtaiu difficult-to-find publications. Especially and Varela de Olmedo 1992; Heyer 1978; Klappeobach and helpful were Martha tdien, Claire Catron, Ron Lindsey, Leslie Langone 1992; Langone 1995; A. Lutz 1926; Nieden 1923; Overstreet, Courtney Shaw, David Steere, and Wanda West. Widmer I 994, as L. mys1ace 11s); species comparisons (Anders Work on this account was supponed in pan by the Ncotropical son 1906 (M); Baumann 1912(M),1917 (M); GaJlardo 1964c; Lowlands Research Program, Smithsonian Institution (Richard Heyer 1969b (M): A. Lutz 1930 (M); Melin 194 1 (M)); species P. Vari, Principal Investigator) and the National Science fQun or taxonomic lists (Achaval 1989; Boulenger 1885a,b; De la dation (award I 9815787) to ROdS and WRH. Riva et al. 2000; de Sli 1983; Frost 1985; Glaw et al. 2000; Gorham 1963, 1966, 1974; Harding 1983; Knauer 1878, 1883; LITERATURE CITED Lavilla 1994a; Lavilla and Cei 2001 ; !>,.. Lut.z 1924; Ringulct and Aramburu 1957). Abe,A.S. and L.S. Gan:ia. 1 990.Altera~ISes de fiuidos corp6reos narll Leptodactylus fuscus durante a esti\la~ (Anura, Leptodactylidac). • NOMENCLATURAL ID STORY. Leptodactylus mystacinus Rcvista Brasilcira de Biologja 50:243-247. was identified as Cystignathus mystaceus, Leptodactylus pyg Abrunhosa. PA., H. Wogel, and J.P. Pombal, Jr. 2001. ~liza~ de maeus, or Cystignathus schomburgkii prior lo its recognition as quano esp6cies de anuros do Estndo do Rio de Janeiro, sudcste do Brasi.1 (Amphibia, Hylidac, Leptodaetylidae). Boletim do Museu a distinct species in 1861. The influential paper by Peters ( 1873) Nacional, No\la ~rie, Z.OO!ogia, Rio de Jaoeiro (472):1-12. erroneously concluded that Hensel's L. mystaceus was not the Ab'S:ibcr,A.N. 1977. Os domfnios monoclio1~ticos na Am~ricado Sul. same species as L. mystacinus. Primeira aprox ima~o. Gcomorfologia 52: 1- 23 + I map. Leptodactylus labia/is (Cope 1877) was considered to be the Ach:ival, F. 1989. Lista de cspccies de vertcbrados del Uruguay. Partc valid name for the taxon now recognized as L.fragilis (Brocch.i 2. Anfibios, reptiles,a"es y mruruferos. Facultad de Humanidades y 1877) priorto 1978 (Heyer 1978, 2002). Ciencias. Servicio de Publicaciones Doccntes lntcmas. Montevideo: Heyer ( 1978) was in error in staling that no locality data other 1-46. than Argentina existed for ll1e holotype of Cystignathus mysta - and A. Olmos. 1997. Anfibios y Reptiles del Uruguay. Serie Fauna No. l,Monte\lideo, Uruguay. cinus. Other authors (Barrio J965b, Cei 1980, Lavilla 1992) - , M .D. Mcncghcl, and A.R. Melgarejo. 1984 (" 1980"). Conservaci6n correctly identified the type locality as Rosario, Argentina. de la fauna de vertcbrados dcl Uruguay. LI Reuni6n lbcronmcricnna Schulz (1944,Agregado 2, Parte B. Planilla l) gave the coordi de Conservaci6n de Zo6logos de Vertebrados:573-583. nates for the Rosario published by Burmeister as 32°56'37" lati Altig, R. and OF. Johnston. 1986. Major characteristics of free-living tude (South) and 60°39'00" longitude West. tadpoles. Smithsonian Herpctologjcal lnfonnation Service (67):1- Lavilla er al. (1995) referred to specimens from lhc Los 75. Colorados Biological Station in Salta, Argentina as L. cf. Alvarez, B.B.,J.A. ~spede:z, M.L. Lions, A. Hernando, and R. Aguirre. mystacinus. The males call from above ground.low on tree trunks 1996. Herpetofaunadc las ProvinciasdcCorricntcS,Chacoy Formosa and are being eval.uated as to whether the Chaco frogs represent (Argentina). Faccna 12:1 19-134. -, R.H. Aguirre, JA. C~pcdcz, A.B. Hemando, and M.E. Tedesco. a species distinct from L. mystacinus (E.O. La villa, pers.comm.). 2002. Atlas de Anfibios y Reptiles de Corrientcs, Chaco y Formosa __(Argentina).l.Anuros,Cecnidos ;Saurios,Anfis~nidosySerpientes. • RE MARKS. "Rana de big~lcs " (He Ledo et al. 1977. de Sli Editorial Universitaria, Univcrsitaria Nacioll'iiTI'lordestc(Corriootes)::· 1983, di Tada et al. 1976, KJappenbach 1969, I:.agomarsino et · ~ AlldcrSSOn, L.G. 1906. On batmchians ·frotn Bolivia, Argentina, and· 767.5 PCni ,collected by Er land NordcnsldOld 1901-1902 Md Nils 1lol mgren - and Dr. ~hue I-Loesche. I 892. Brduns T"1edeben . Allgcmeine Kunde 1904-1905.Aikiv fllrZoologi 3: 1- 19 +I plate. desTierrcichs. Die Kricchtiere und Lurche.Bibliographischcs lns1i1u1, Anooymous . I 877. The development ofbatrochians without metrunorpb Leipzig und Wien. osis. Nature I 5:491-492. Bogart, J.P. 1974. A karyosystcmatic study of fr~ in che genus up10- - . 1997. Aspec1os ccol6gicos dos vcnebrados tem:strcs e semi-aqu6ti dactylus (Anura: Leptocbctylidae). Copcia 1974:728-737. cos no Pantanal, p. 197-433. /n Pl ano de Conserva~iio da Bacio do Boneiro, C., F. Kolene, G. Niquicbcnco, J. Rendo, and M.Tcdros. 1999. Aho Paraguai, Diagn6stico dos Meios Ffsico e Bi6tico. Volume U, Anfibios y reptiles del Cerro de Montevideo y atrededores [Abslractj. Torno 111. hmpresa Brasilcira de Pesquisa. Agropecuaria, ProjclO Bolel!n de la Sociedad Zool6gica dcl Uruguay, Segunda apoca (Ac1as Pani.n:1l, Proarama Nacional de Mcio Ambicn1e, Brasilia, Brasil. de las V JOOl3das de 1.ooloifa del Uruguay) 11 :9. Aquino, AL., NJ . Scoct, ond M. 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