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Vocalizations of Eight Species of Atelopus (Anura: Bufonidae) with Comments on Communication in the Genus

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Vocalizations of Eight Species of Atelopus (Anura: Bufonidae) with Comments on Communication in the Genus Copeia, 1990(3), pp. 631-643 Vocalizations of Eight Species of Atelopus (Anura: Bufonidae) With Comments on Communication in the Genus REGINALD B. COCROFT, ROY W. MCDIARMID, ALAN P. JASLOW AND PEDRO M. RUIZ-CARRANZA Vocalizations of frogs of the genus Atelopus include three discrete types of signals: pulsed calls, pure tone calls, and short calls. Repertoire composition is conservative across species. Repertoires of most species whose calls have been recorded contain two or three of these identifiable call types. Within a call type, details of call structure are very similar across species. This apparent lack of divergence in calls may be related to the rarity of sympatry among species of Atelopus and to the relative importance of visual communication in their social interactions. COMMUNICATION in frogs of the genus lopus by comparing the structure and behavioral Atelopus has been httle studied despite sev- context, where available, of calls of these species eral unique features of their biology that make with calls of other species reported in the lit- them an important comparative system in light erature. of the extensive work on other species of an- METHODS urans (Littlejohn, 1977; Wells, 1977; Gerhardt, 1988). First, the auditory system oí Atelopus is Calls were analyzed using a Kay Digital Sona- highly modified, and most species lack external Graph 7800, a Data 6000 Waveform Analyzer and middle ears (McDiarmid, 1971). Second, in with a model 610 plug-in digitizer, and a Mul- contrast to most anurans, species oiAtelopus are tigon Uniscan II real-time analyzer. Call fre- diurnal and often brightly colored, and visual quencies were measured from waveform and communication plays an important role in the Fourier transform analyses; pulse rates and call social behavior of at least some species (Jaslow, lengths were measured from waveform analyses 1979; Crump, 1988). Finally, recent behavioral or from wide-band audiospectrograms. Call rates studies by Crump (1988) indicate that in at least were measured from the real-time analyzer one species (Fig. 1), vocalizations primarily me- screen or with a stopwatch directly from the diate male-male interactions and not mate at- tapes. Original tapes of some species and copies traction as is the case in most anurans. of all tapes are on file in the tape archive, Di- Detailed information on vocalizations oï Ate- vision of Amphibians and Reptiles, USNM; lopus has been reported for only four species original tapes of other species are on file as in- (Jaslow, 1979; Lescure, 1981; Asquith and Al- dicated below. tig, 1987), and laryngeal morphology has been Few field recordings of Atelopus are available, examined in three species (Martin, 1972). From and therefore some of the recordings utilized our fieldwork, recordings made by colleagues, in this and other studies (Asquith and Altig, and a review of the tape archives of the National 1987; Jaslow, 1979; Lescure, 1981) were made Museum of Natural History (USNM), the from animals in captivity. We tested the poten- American Museum of Natural History (AMNH), tial for distortion of calls from animals calling and the Museum of Natural History, University from plastic bags by analyzing recordings of a of Kansas (KU), we have analyzed recordings of single male of Hyla koechlini Duellman and Trueb the calls of eight species of Atelopus, including calling in and out of a bag (this species has a seven species for which calls have not been re- pulsed call somewhat similar in structure to calls ported previously. These species include A. cru- of Atelopus); our analyses showed no discernible ciger, A. minutulus, A. nicefori, A. senex, A. spu- difference between the two recordings. We also marius (two populations), A. varius (two carefully inspected laboratory recordings for populations), A. zeteki, and A. sp. Our purposes evidence of reverberation or smearing of the are: 1) to summarize the available information call waveform. Calls with waveforms showing on calls of Atelopus; and 2) to ask what this in- evidence of degradation were eliminated from formation suggests about call evolution in Ate- the temporal analyses. © 1990 by the American Society of Ichthyologists and Herpetologists 632 COPEIA, 1990, NO. 3 Fig. 1. Male Atelopus varius from a population along the Rio Lagarto, 3 km SSW Santa Elena, Puntarenas Province, Costa Rica. Males in this population apparently use the pulsed call to communicate with other males and rely on visual cues to locate mates (Crump, 1988). Photo courtesy of Martha L. Crump. We use the following abbreviations in re- northern Venezuela. Although Atelopus is poor- porting call data: call length (cl); pulse rate (pr); ly known as a group, A. cruciger is an exception number of pulses (pn); dominant frequency (df); primarily because of its relative abundance and and frequency modulation (fm). All snout-vent the published observations on its life history length (SVL) measurements are in millimeters. (Sexton, 1958) and larval development (Mebs, Institutional abbreviations follow Levitón et al. 1980). Even though Sexton (1958) reported that (1985). males of this species call, no literature on vocal- izations of A. cruciger exists. MATERIALS Several calling males of A. cruciger (SVL about 30) were located by RWM along a rocky stream Frogs of the genus Atelopus often are con- about I km south of Cuyagua, Estado Aragua, spicuous members of Neotropical anuran fau- Venezuela, on 19 Dec. 1976. About 12 males nas in evergreen wet forest habitats, especially were seen in a 20 m section of the shoreline. at elevations between 800 and 2000 m from Seven single males and an amplexing pair were Costa Rica south to Bolivia and in the Guianas collected and transported to Caracas where they and northern Brazil. These diurnal, slow-mov- were put into a planted terrarium and observed. ing frogs are usually brightly colored and fre- Some of the males resumed calling. At least quently concentrate along small to moderate- three distinct types of calls given by captive male sized streams, in which they breed. In spite of A. cruciger were recorded in the laboratory: a their conspicuousness and diversity (Frost [1985] pulsed call, a pure tone call, and a short call listed 43 species), relatively little is known about (temperature information not available). their reproductive ecology and behavior. Male size, habitat, and general range of the nine Atelopus minutulus Ruiz-Carranza, Hernandez-Ca- species studied in this work are reported in Ta- macho and Ardila.•This small species was re- ble I. cently described from specimens taken at a sin- gle site in wet forest at 1560 m near Vereda de Atelopus cruciger (Lichtenstein and Martens).•This Portachuelo (vie. Manzanares), Departamento species is relatively common along several of the de Meta, Colombia. W. Lámar located males streams that drain the wet forests of Parque (SVL X = 19.8, n = 25) calling from rocks along Nacional Henri Pittier, which includes the bi- the edge of a small stream at 1640 m between ological research station of Rancho Grande in 1530 and 1700 h on 8 July 1980. He recorded COCROFT ET AL.•VOCALIZATIONS IN ATELOPUS 633 TABLE 1. SPECIES OF Alelopus FOR WHICH CALL DATA ARE REPORTED IN THIS STUDY. Male SVL, habitats and distributions, and sources of information are indicated. Species/SVL (mm) General distribution Source Atelopus cruciger cloud forests, coastal range of northern Venezuela Sexton, 1958 28-33 Atelopus minutulus cloud forests, eastern Andes, Meta, Colombia Ruiz-Carranza et al., 1988; 17-20 Lamar, pers. comm. Atelopus nicefori montane wet forests, near Caicedo, Antioquia, Co- Rivero, 1963; Ruiz-Carranza 17.5-22.5 lombia etal., 1988 Atelopus spumarius Amazonian lowlands, Peru, Ecuador, northern Bra- Peters, 1973; Lescure, 1973, 23-29 zil and Guianas 1981 Atelopus senex montane wet forests, central and southern Costa Savage, 1972 28-32 Rica Atelopus varius wet forests, Costa Rica, Panama and northern Co- Savage, 1972 27-41 lombia Atelopus zeteki wet forests. Valle de Anton area of western Pana- Dunn, 1933; This study 34-43 ma Atelopus sp. Caribbean lowlands, eastern Panama Myers, pers. comm. 24-29 at least three call types from individuals in a captive specimens (USNM 269068-269069,19- plastic bag (temperature information not avail- 21 SVL) of A. spumarius spumarius collected at able). 640 m on the Rio Tambopata, Departamento Madre de Dios, Peru (temperature information Atelopus nicefori Rivero.•This small species (male not available). Thirteen calls of a single male holotype measured 19 SVL) has been reported (KU 129958, 29.4 SVL) referred to A. s. hoog- from two localities, at 1800 and 2670 m, near moedi (Lescure, 1981) from the SUDAM Floral Caicedo, Antioquia, Colombia (Rivero, 1963; Reserve, 74 km SE Satarem, Estado Para, Brasil, Rivero and Serna, 1985). Frogs at the higher were recorded by Martha Crump on 5 July 1970; elevation were found in wet forest near the temperature 27 C (KU tape 989, Reel 88). mouth of a small stream. Five calls were re- corded by PRC from a male A. nicefori (ICN Atelopus varius group.•Atelopus chiriquiensis, A. 1304, SVL 21.3) found in amplexus with a fe- senex, A. varius (an extremely variable species), male during the day on 30 April 1972, at "La and A. zeteki (considered a subspecies of varius Nevera," Vereda El Chuscal, Departamento by many authors) are closely related members Antioquia, Colombia. This locality is in a resid- of a Middle American species group reviewed ual patch of cloud forest at 2585 m elevation by Savage (1972). These forms are diurnal and on the western slope of the Western Andes. The usually found near fast-moving, small streams pair was taken with several other individual in wet evergreen forests from near sea level to males and females along a small, clear creek; above 2150 m.
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