Herpetologists' League Visual and Acoustic Communication in the Brazilian Torrent Frog, Hylodes asper (Anura: Leptodactylidae) Author(s): Célio F. B. Haddad and Ariovaldo A. Giaretta Source: Herpetologica, Vol. 55, No. 3 (Sep., 1999), pp. 324-333 Published by: Herpetologists' League Stable URL: http://www.jstor.org/stable/3893226 . Accessed: 24/01/2014 11:44 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Herpetologists' League is collaborating with JSTOR to digitize, preserve and extend access to Herpetologica. http://www.jstor.org This content downloaded from 186.217.234.230 on Fri, 24 Jan 2014 11:44:35 AM All use subject to JSTOR Terms and Conditions 324 HERPETOLOGICA [Vol. 55, No. 3 OKADA, Y. 1931. The Tailless Batrachians of the Jap- borea (L.). Pp. 295-4336. In D. H. Taylor and S. I. anese Empire. Imperial Agricultural Experiment Gutmann (Eds.), The Reproductive Biology of Am- Station, Nishigahara, Tokyo, Japan. phibians. Plenum Press, New York, New York, U.S.A. RYAN, M. J., A. S. RAND, AND L. A. WEIGT. 1996. UEDA, H. 1994. Mating calls of the pond frog species Allozyme and advertisement call variation in the distributed in the Far East and their artificial hy- tuingara frog, Physalaemus pustulosus. Evolution brids. Scientific Report of the Laboratory for Am- 50:2435-2453. phibian Biology, Hiroshima University 13:197-232. SAS INSTITUTE. 1988a. SAS User's Guide: Basics. ZHAO, E-M., AND K. ADLER. 1993. Herpetology of Version 6 ed. SAS Institute Inc., Cary, North Car- China. Society for the Study of Amphibians and olina, U.S.A. Reptiles. Contributions to Herpetology 10:1-522. 1988b. SAS User's Guide: Statistics. Version 6 ed. SAS Institute Inc., Cary, North Carolina, U.S.A. SCHNEIDER, H. 1977. Acoustic behavior and physiology Accepted: 24 September 1998 of vocalization in the European tree frog, Hyla ar- Associate Editor: Richard Howard Herpetologica, 55(3), 1999, 324-333 C 1999 by The Herpetologists' League, Inc. VISUAL AND ACOUSTIC COMMUNICATION IN THE BRAZILIAN TORRENT FROG, HYLODES ASPER (ANURA: LEPTODACTYLIDAE) CELIO F. B. HADDAD13 AND ARIOVALDOA. GIARETTAA2 'Museum of Vertebrate Zoology, University of California, Berkeley, CA 94720, USA 2Departamento de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas, C.P. 6109, 13083-970 Campinas, Sio Paulo, Brasil ABSTRACT: We studied the signaling, territorial, and courtship behaviors of the diurnal frog Hylodes asper. Visual and acoustic communication were used during intraspecific interactions in- volving males, females, and subadults. Hylodes asper has a complex visual communication system, of which foot-flagging is the most distinctive display observed in the repertoire of visual signals. The splash zone produced by the waterfalls and torrents creates a high, nearly constant, humidity near the streams, reducing the risk of desiccation which enables the diurnal activity of H. asper Although the ambient sound pressure levels (SPL), measured at the calling sites, are similar to the SPL of the advertisement calls, the high-pitched calls of H. asper are spectrally different from the noise produced by the water current. Thus, the ambient noise produced by the water current may not interfere significantly with the acoustic communication of this species. The noise and the nearly constant and high humidity produced by the torrents and waterfalls, along with the availability of light, probably favored the evolution of contrasting colors and visual communication in H. asper. Males of H. asper excavate underwater chambers that are probably used to shelter the eggs and to prevent the clutch from being drifted downstream. Key words: Hylodes asper; Anura; Hylodinae; Visual communication; Acoustic communication; Reproductive mode; Atlantic Forest; Southeastern Brazil AMONG amphibians, vocalization is munication by acoustic signals is effective highly developed in anurans, most species in several circumstances, including situa- of which have vocal structures capable of tions where vision is limited (Gerhardt, producing a variety of sounds that attract 1983). Acoustic communication must have mates, advertise territories, or express dis- been favored by the pump breathing tress (Duellman and Trueb, 1986). Com- mechanism (Duellman and Trueb, 1986) and by the nocturnal activity observed for the majority of anuran species. The 3 PRESENT ADDRESS: Departmento de Zoologia, In- evo- stituto de Biociencias, Universidade Estadual Paulista, lution of visual communication in some C.P. 199, 13506-900 Rio Claro, Sao Paulo, Brasil. species of frogs has apparently been fa- This content downloaded from 186.217.234.230 on Fri, 24 Jan 2014 11:44:35 AM All use subject to JSTOR Terms and Conditions September 1999] HERPETOLOGICA 325 vored by the availabilityof ambient light h of observations, from March 1988- (Lindquist and Hetherington, 1996, and March 1994, from 0530-2000 h. The study references therein). Diurnal species of an- was conducted at Paranapiacaba, Munici- urans commonly use visual displays, ap- pio de Santo Andre (23047'S, 46018'W, 800 parently for communication (e.g., Durant m above sea level), and Picinguaba, Mun- and Dole, 1975; Wells, 1980), with limb icipio de Ubatuba (23023'S, 44?50'W, 50- signals being the most distinctive behavior 100 m above sea level), State of Sao Paulo, described (e.g., Davison, 1984; Harding, Brazil, in streams with waterfalls and ex- 1982; Heyer et al., 1990; Pombal et al., tensive rapids. 1994). The apparentuse of limb signals for Focal animal and all occurrence sam- visual communication in anuranshas been plings were used for behavioral records observed for the following genera: Brachy- (Lehner, 1979). Experimental manipula- cephalus (Brachycephalidae), Atelopus tions were restricte to the manual inser- (Bufonidae), Dendrobates (Dendrobati- tion of a mirror (15 x 10 cm) next to res- dae), Litoria (Hylidae), Hylodes (Lepto- ident males, to test the importance of vi- dactylidae), Taudactylus (Myobatrachy- sion during the territorial interactions. De- dae), and Staurois (Ranidae) (see revision scriptions of the behaviors are based on in Lindquist and Hetherington, 1996). In individuals observed and videotaped in the addition to these genera, leg signals are field. Individuals were not marked, but performed by males during territorial in- some of them were recognized in succes- natural teractions in two nocturnal species in the sive days by distinctive markings and color patterns. We recorded advertise- genus Phyllomedusa (Hylidae) (P. distinc- ment calls in a Nagra E tape recorder us- ta: Castanho, 1994; P. burmeisteri: obser- ing a Sennheiser ME 80 micro phone; oth- vations by C. F. B. Haddad). er vocalizations were recorde in a Sony The subfamily Hylodinae comprises TCM-74v tape recorder using a Sony car- frogs distributed among the genera Cros- dioid microphone. The sonograms were sodactylus, Hylodes, and Megaelosia. A produced in a Macintosh computer cou- great number of species in this subfamily pled to the MacRecorder Sound System is diurnal, and males of most species in the 2.0.5, using 8 bit resolution, 22 kHz sam- genera Crossodactylus and Hylodes have pling frequency, and FFT with 256 points. loud bird-like voices (Vielliard and Car- For the statistical analysis of the results, doso, 1996; Weygoldt and Carvalho e Sil- we used the Mann-Whitney rank sum test, va, 1992); Megaelosia is apparentlyvoice- with a = 0.05 (Zar, 1996). less (Giaretta et al., 1993). Although some species of Hylodinae are common along RESULTS small forest streams in south and south- Hylodes asper was acoustically active in eastern Brazil, little is known about their all months, showing a peak in activity dur- reproductive behavior (Heyer et al., 1990; ing the rainy season (October-March). Weygoldt and Carvalhoe Silva, 1992). Hy- During the night, males, females, and ju- lodes asper, which has a distribution from veniles of H. asper were observed resting south to southeastern Brazil, occurring in crevices or on branches and leaves up from Santa Catarina to Rio de Janeiro to 1.0 m above the ground, near streams. (Frost, 1985), is known for displayingboth Tadpoles were observed performing feed- acoustic signals and apparentvisual signals ing activities during the day and at night near mountain streams (Heyer et al., 1990; in the streams. Males started to emit ad- Hodl et al., 1997). In the present study, vertisement calls during sunrise and we describe the signaling, territorial,nest stopped during sunset. The communica- tion of H. and courtship behaviors of this asper is based on visual and building, acoustic signals, as described below. diurnal frog. Visual Communication Performed by MATERIALS AND METHODS Isolated Males The behaviors of H. asper were studied During the day, males defended during 18 visits, totaling 45 days and 112 emerged rocks and logs as territories, used This content downloaded from 186.217.234.230 on Fri, 24 Jan 2014 11:44:35 AM All use subject to JSTOR Terms and Conditions 326 HERPETOLOGICA [Vol. 55, No. 3 We quantified the foot-flagging behavior of 11 calling males and observed that both left and right legs are used with similar frequencies (the right leg was used 71 times and the left leg was used 89 times); however, they were not used with regular alternation. The males may call without any observable movement of the limbs (n = 58 calls for 11 observed males). Some- times the foot-flagging display (26 obser- vations for eight males) and the move- ments of the toes (27 observations for two males) were performed by males that were not calling. We observed subtle, but ap- ment call. Note the inflated vocal sacs.