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Visual Signaling in Phrynobatrachus Krefftii Boulenger, 1909 (Anura: Ranidae)

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Visual Signaling in Phrynobatrachus Krefftii Boulenger, 1909 (Anura: Ranidae) 18 HERPETOLOGICA [Vol. 62, No. 1 (Amphibia: Plethodontidae). Journal of Morphology WALLS, S. C., A. MATHIS,R.G.JAEGER, AND W. F. GERGITS. 234:131–146. 1989. Male salamanders with high-quality diets have SMUTS, B. B., AND R. W. SMUTS. 1993. Male-aggression faeces attractive to females. Animal Behaviour 38: and sexual coercion of females in nonhuman-primates 546–547. and other mammals—evidence and theoretical WICKLER,W.,AND U. SEIBT. 1983. Monogamy: an implications. Advances in the Study of Behaviour ambiguous concept. Pp. 33–50. In P. Bateson (Ed.), 22:1–63. Mate Choice. Cambridge University Press, New York, DE WAAL, F. B. M. 2000. Primates—a natural heritage of New York, U.S.A. conflict resolution. Science 289:586–590. WISE, S. E., AND R. G. JAEGER. 1998. The influence of tail DE WAAL, F. B. M., AND F. AURELI. 1997. Conflict re- autotomy on agonistic behaviour in a territorial sala- solution and distress alleviation in monkeys and apes. mander. Animal Behaviour 55:1707–1716 Pp. 317–328. In C. S. Carter, I. I. Lenderhendler, and B. Kirkpatrick (Eds.), Integrative Neurobiology of Affiliation. Annals of the New York Academy of Accepted: 6 September 2005 Sciences, Albany, New York, U.S.A. Associate Editor: Troy A. Baird Herpetologica, 62(1), 2006, 18–27 Ó 2006 by The Herpetologists’ League, Inc. VISUAL SIGNALING IN PHRYNOBATRACHUS KREFFTII BOULENGER, 1909 (ANURA: RANIDAE) 1 1,2 WALTER HIRSCHMANN AND WALTER HO¨ DL 1Department of Evolutionary Biology, University of Vienna, Althanstraße 14, A-1090 Wien, Vienna, Austria ABSTRACT: During intraspecific interactions involving males, females and subadults, acoustic as well as visual signals were observed in the diurnal frog Phrynobatrachus krefftii. Strikingly, interactions between adult males are highly dominated by inflations of the bright yellow subgular vocal sac without sound production. We studied the signaling behavior in males of P. krefftii at the Amani Nature Reserve, Usambara mountains, Tanzania, from November 2001 to March 2002. Under nonmanipulated conditions, we registered 641 male- male interactions involving 31 individuals during 323 hours of observation. Most (496 or 77 % ) inflations of the vocal sac were purely visual; the remainder (145 or 23%) of the signals were bimodal, and accompanied by sound production. To test whether exclusive visual signaling can be evoked under experimental conditions, we introduced tethered males into the visual field of 28 residents, registering the elicited responses over 10 minutes after the introduction. Unimodal (i.e., exclusive visual) signals (825 out of 1106 responses) dominated over vocal-sac inflations accompanied with sound production. In seven selected focal males, the rate of exclusive visual signaling (0.57 signals/min), however, was more frequent than under nonmanipulated conditions (0.13 signals/min). Phrynobatrachus krefftii is the first species of anuran amphibians reported to perform nonaudible vocal-sac inflations during intraspecific male-male signaling behavior. Key words: Acoustic communication; Agonistic behavior; Amphibia; Anura; Phrynobatrachus krefftii; Visual signaling; Vocal sac THE MOST prominent mode of communica- 1997; Lindquist and Hetherington, 1996, tion in anuran amphibians is, by far, the use 1998; Richards and James, 1992). All presently of sound production. Compared to acoustic described species performing visual displays cues, visual signals play a subordinate role in emit acoustic signals as well (Ho¨dl and amphibian communication. Some anuran spe- Ame´zquita, 2001), which makes it difficult to cies, however, perform remarkable visual dis- separate the functional role of both communi- plays in variable social contexts (Ame´zquita cation modalities (but see Narins et al., 2003). and Ho¨dl, 2004; Davison, 1984; Haddad and Diurnal species of anurans commonly produce Giaretta, 1999; Harding, 1982; Ho¨dl et al., visual displays (Durant and Doyle, 1975; Wells, 1980) with limb signals being the most distinctive behavior described (e.g., Davison, 2 CORRESPONDENCE: e-mail, [email protected] 1984; Harding, 1982; Heyer et al., 1990; Ho¨dl March 2006] HERPETOLOGICA 19 FIG. 1.—Phrynobatrachus krefftii. Female (above left) and male without (above right), partly (below right) and fully expanded vocal sac (below left). Note the bright yellow subgular region of the male. Photographs taken by W. Ho¨dl at Zigi River (Amani Nature Reserve), Tanzania. et al., 1997; Pombal et al., 1994 ). In addition to 1999). The endemic P. krefftii is diurnal and its acoustic function, the vocal sac also may act inhabits localities of seepage water, rivulets as a conspicuous visual signal. Despite of the and streams in submontane and montane variety of diverse vocal-sac types, their visual rainforest within the East and West Usambara relevance has not yet been explored thoroughly Mountains in Tanzania (Barbour and Lover- (Ho¨dl, 1996). idge, 1928). Within the Amani Nature Reserve Visual signaling so far has been recorded only (ANR) of the East Usambara mountains, P. in tropical frogs, with many species occurring krefftii occurs at elevations above 800 m in ¨ ´ along streams and creeks (Hodl and Amez- a submontane evergreen forest. quita, 2001; Lindquist and Hetherington, A remarkable color dimorphism is found 1998). We searched for other visually commu- between males and females: Males reach up to nicating species along East African mountain 50 mm in snout–vent length (SVL) and possess streams, and found that the diurnally active Phrynobatrachus krefftii appeared to be a good a brightly yellow subgular region. Females candidate. Males of this ranid species are dull reach 40 mm in SVL (Barbour and Loveridge, brown in color, but possess a strikingly bright- 1928), and their throat region is light brown yellow vocal sac. Thus, the aim of this study was colored (Fig. 1). Inflatable vocal sacs are to investigate the possible role in communica- present only in sexually mature males. Two tion of the conspicuously colored gular region more species of the genus Phrynobatrachus, during intraspecific encounters. P. alleni and P. accraensis (5 P. latifrons), also The exclusive African frog genus Phrynoba- possess yellow throat coloration in breeding trachus currently contains 66 species, all of males (Ro¨del, 2000, 2003; Ro¨del and Agyei, which occur in sub-saharan Africa (Poynton, 2003). 20 HERPETOLOGICA [Vol. 62, No. 1 Due to the regular occurrence of egg during both day and night hours. The frogs clutches and tadpoles throughout a nine were located on their nocturnal resting sites in month observation period (July 2001–March the dense vegetation using a head lamp 2002), P. krefftii is considered a ‘‘prolonged between 2000 h–0300 h every night between breeder’’ (sensu Wells, 1977). Males occupy 09 June 2001 and 15 July 2001. All individuals breeding territories containing rocks and fallen larger than 20 mm SVL were toe clipped branches in shallow water. Territories are sites (according to Hero, 1989). The nonmanipu- for mate attraction and oviposition, as well as lated observations started at 0900 h and ended areas for feeding and shelter. Male residents at 1700 h, and corresponded to the typical readily scrutinize conspecific intruders in their period of diurnal activity of P. kreffii. In total, visual field by leaving their calling sites and 14 sites (seven each along a stream and approaching the offenders. At night, adult as a rivulet) with good visibility and access were well as juvenile frogs are found at resting sites selected, each containing at least four adult on leaves within the vegetation along the banks males. The order of observations within the of the stream at approximately one meter pool of individuals was chosen randomly. above the ground. Out of 158 recaptures, 59 Nonmanipulated observations were obtained (30%) toe-clipped individuals were found less from a distance of approximately 2 m from the than 20 cm from the initially marked site on focal individual. Each individual observation leaves each night over a period of seven days. lasted 30 min. For vocal recording, a micro- Calling activities of male frogs were observed phone was placed 0.75 m in front of the focal throughout the day (W. Hirschmann and male. Recordings were made with a Sony Pro- M. Franze´n, unpublished data). fessional Walkman WMDC6 using a directional microphone (AKG C568 EB). To measure the background noise and to test the possible MATERIALS AND METHODS acoustic energy produced during nonaudible The study was conducted near the village of vocal sac inflation, a sound-pressure level me- Amani (5869 S and 388389 E, 900 m), Tanga- ter (Volticraft No. 33-2050; ‘‘A’’ weighted, fast District, Tanzania within a submontane forest response) was positioned 0.75 m in front of of the East Usambara Mountains belonging to the signaling individual. Sonagrams were pro- the Zanzibar-Inhambane transitional rainfor- duced with a MAC computer using Canary est type (White, 1983). Mean annual rainfall (1.2.4) software. (1902–1970) is 1910 mm (Rodgers and Home- To test the importance of visual encounters wood, 1982) with peak precipitation in March, and to confirm that the visual signal is directed April and May. Two study areas were selected towards an introduced male frog during within the boundaries of the Amani Nature territorial interactions, we performed experi- Reserve. A 400-m transect along the margin of ments with tethered frogs. Individual P. krefftii the Zigi river (58099 S and 388639 E) and a 112- males (SVL x 6 SD 5 36.4 6 3.6 mm, n 5 m stretch along the border of a rivulet (58109 S 15) were caught and used at the onset of the and 388639 E) were marked by colored tapes. rainy season between 28 February 2002 and 15 The stream was bordered by gallery forest and March 2002 starting each day at 0900 h and secondary growth, with the soil surface con- ending at 1700 h. Experimental manipulations tinuously being soaked by seepage water. The were restricted to the manual introduction of rivulet was partly covered with dense vegeta- a tethered male P.
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