The Behavioral Ecology of Anuran Communication
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The Behavioral Ecology of Anuran Communication
Wells, K. D. and J. J. Schwartz. 2007. The behavioral ecology of anuran communication. Pp. 44-86. In: Narins, P.M., Feng. A.S.. Fay, R.R.. and Popper, The Behavioral Ecology of Anuran Communication, A.N. (eds) Hearing and Sound Communication in Amphibians, springer Handbook of Auditory Research (Vol. 28). New York: Springer-Verlag.The original publication is available at www.springer.comhttp://www.springer.com/life+sciences/animal+sciences/book/978-0-387-32521-73The Behavioral Ecology of Anur The Behavioral Ecology of Anuran Communicationan CommunicationKentwood D. Wells and Joshua J. Schwartz1IntroductionAs the last rays of sunlight disappear from the evening sky. a shallow marsh in PThe Behavioral Ecology of Anuran Communication
anama begins to come alive with the calls of frogs and toads. Among these is a small yellow and brown hourglass treefrog, Hyla ebraccata (Fig. 3.1C). Wells, K. D. and J. J. Schwartz. 2007. The behavioral ecology of anuran communication. Pp. 44-86. In: Narins, P.M., Feng. A.S.. Fay, R.R.. and Popper, The Behavioral Ecology of Anuran Communicationbegins to develop. The first male responds to the calls of his neighbors by placing his own calls immediately after their calls, and he soon increases his calling rate and begins to add clicklike secondary notes to his calls in an attempt to outsignal his rivals. Suddenly another male calls only a f The Behavioral Ecology of Anuran Communicationew centimeters away, and the first male responds by modifying the introductory notes of his calls, producing aggressive notes with a pulse repetitionThe Behavioral Ecology of Anuran Communication
rate about three times that of his advertisement calls. As the two males approach each other, they gradually increase the duration of their aggressiveWells, K. D. and J. J. Schwartz. 2007. The behavioral ecology of anuran communication. Pp. 44-86. In: Narins, P.M., Feng. A.S.. Fay, R.R.. and Popper, The Behavioral Ecology of Anuran Communicationirst male returns to advertisement calling.Having sorted out spacing within the chorus, most of the males soon settle into a regular rhythm of advertisment calling, punctuated by occasional aggressive calls. Periodically they stop calling as their calls are overpowered by bursts of calling from grou The Behavioral Ecology of Anuran Communicationps of males of another frog, the small-headed treefrog (Hyla microcephala) (Fig. 3.ID). The males of H. ebraccata have difficulty making their calls aThe Behavioral Ecology of Anuran Communication
udible when surrounded by the other species, and they attempt to place their calls in the silent periods between bursts of H. microcephala calling actWells, K. D. and J. J. Schwartz. 2007. The behavioral ecology of anuran communication. Pp. 44-86. In: Narins, P.M., Feng. A.S.. Fay, R.R.. and Popper, The Behavioral Ecology of Anuran Communicationiately switches to a rapid series of repeated introductory advertisement call notes. The female turns toward the male, and with a few zigzag hops, approaches his calling site and allows him to clasp her in amplexus. The pair then moves off to find asuitable leaf on which to lay their eggs, positione The Behavioral Ecology of Anuran Communicationd a half meter or so above the shallow water where the tadpoles will complete their development. After mating,EFFigure 3 1. Calling males of some anurThe Behavioral Ecology of Anuran Communication
ans in which vocal communication has been studied in detail. (A) Pseudacris crucifer (Hylidae) (B) Hyla versicolor (Hylidae). (C) Hyla ebraccata (HyliWells, K. D. and J. J. Schwartz. 2007. The behavioral ecology of anuran communication. Pp. 44-86. In: Narins, P.M., Feng. A.S.. Fay, R.R.. and Popper, The Behavioral Ecology of Anuran Communicationtwood D. Wells.the male takes no further interest in his offspring, but moves to a nearby calling perch and resumes calling.These scenes, which can be repeated dozens of times each night in a single breeding assemblage in Panama (Schwartz and Wells 1984a; Wells and Schwartz 1984a,b), illustrate the The Behavioral Ecology of Anuran Communicationcomplexity of vocal interactions that can occur in a chorus of frogs. Many anurans have repertoires consisting of several distinct types of calls, andThe Behavioral Ecology of Anuran Communication
they often exhibit considerable plasticity in their use of different call elements, responding to changes in local chorus density, the presence of neWells, K. D. and J. J. Schwartz. 2007. The behavioral ecology of anuran communication. Pp. 44-86. In: Narins, P.M., Feng. A.S.. Fay, R.R.. and Popper, The Behavioral Ecology of Anuran Communicationle frogs have the same ultimate goal: to outsignal their competitors and attractfemales, eventually fertilizing their eggs to contribute their genes to the next generation of frogs. The ways in which they accomplish this goal vary among species, however. Some anurans have relatively simple calls, wh The Behavioral Ecology of Anuran Communicationereas others have exceedingly complex calls. Males of some species are very aggressive toward other males in a chorus, whereas males of other speciesThe Behavioral Ecology of Anuran Communication
seldom react to their neighbors.This chapter reviews the behavioral ecology of anuran vocal communication. First, the influence of sexual selection onWells, K. D. and J. J. Schwartz. 2007. The behavioral ecology of anuran communication. Pp. 44-86. In: Narins, P.M., Feng. A.S.. Fay, R.R.. and Popper, The Behavioral Ecology of Anuran Communicationocal signals of females are discussed. The interactions among males in choruses and the ways in which these interactions affect the ability of males to attract mates are covered in some detail, followed by a brief discussion of ways in which features of the anuran auditory system contribute to commu The Behavioral Ecology of Anuran Communicationnication within a chorus setting.2Sexual Selection, Energetic Constraints, and Signaling System EvolutionWhen Charles Darwin originally outlined his tThe Behavioral Ecology of Anuran Communication
heory of sexual selection in his book, he had relatively little to say about sexual selection in amphibians. He did suggest that the calls of frogs arWells, K. D. and J. J. Schwartz. 2007. The behavioral ecology of anuran communication. Pp. 44-86. In: Narins, P.M., Feng. A.S.. Fay, R.R.. and Popper, The Behavioral Ecology of Anuran Communicationn driving force in the evolution of anuran acoustic communication (Gerhardt and Huber 2002). Many features of anuran calls can be shaped by sexual selection, including call intensity, calling rate, call duration, call pitch, and the temporal pattern of interaction among competing males. In addition, The Behavioral Ecology of Anuran Communication all of the morphological, physiological, and biochemical machinery involved in call production is molded by sexual selection. Because the energetic cThe Behavioral Ecology of Anuran Communication
ost of calling in many species is quite high, selection should favor mechanisms to increase the efficiency of sound production and transmission, thereWells, K. D. and J. J. Schwartz. 2007. The behavioral ecology of anuran communication. Pp. 44-86. In: Narins, P.M., Feng. A.S.. Fay, R.R.. and Popper, The Behavioral Ecology of Anuran Communication Physiology of Call-Producing MusclesThe basic mechanisms of sound production during calling are discussed in the next chapter (see Walkowiak, Chapter 4). The power for sound production is provided mainly by the sexually dimorphic trunk muscles, and sexual selection has produced a number of morpholo The Behavioral Ecology of Anuran Communicationgical and biochemical adaptations for call production by males, including highly aerobic muscle fibers, high concentrations of mitochondria, high actiThe Behavioral Ecology of Anuran Communication
vities of aerobic enzymes, heavy vascularization, and ample supplies of lipid and carbohydrates to fuel call production. There is a strong interspecifWells, K. D. and J. J. Schwartz. 2007. The behavioral ecology of anuran communication. Pp. 44-86. In: Narins, P.M., Feng. A.S.. Fay, R.R.. and Popper, The Behavioral Ecology of Anuran Communication high calling rates (Wells 2001).The repeated contraction of the trunk muscles to produce calls can be energetically expensive in species with high calling rates (Wells 2001). The North American spring peeper (Pseudacris crucifer. Fig. 3.1A) produces single-note calls (Fig. 3.2A), each representing The Behavioral Ecology of Anuran Communicationone contraction of the trunk muscles, and can produce up to 100 call notes per minute. Similarly, the tiny Neotropical smallheaded treefrog (Hyla micrThe Behavioral Ecology of Anuran Communication
ocephala) (Fig. 3.ID) produces long trains of notesgrouped into multinote calls (Fig. 3.2E) and also can produce up to 100 notes per minute (Wells andWells, K. D. and J. J. Schwartz. 2007. The behavioral ecology of anuran communication. Pp. 44-86. In: Narins, P.M., Feng. A.S.. Fay, R.R.. and Popper, The Behavioral Ecology of Anuran Communicationuse anaerobic metabolism plays little role in call production. For these small treefrogs, rates of oxygen consumption during calling can be up to 25 times resting rates. This results in a significant drain on stored energy reserves. Shortterm performance probably is limited mainly by carbohydrate re The Behavioral Ecology of Anuran Communicationserves in the form of glycogen stored in the muscle tissue (Bevier 1997b), whereas lipid reserves are depleted over longer time intervals (Ressel 2001The Behavioral Ecology of Anuran Communication
). The high energetic cost of calling probably explains the relatively short average chorus tenure of many male frogs (Murphy 1994), which exerts stroWells, K. D. and J. J. Schwartz. 2007. The behavioral ecology of anuran communication. Pp. 44-86. In: Narins, P.M., Feng. A.S.. Fay, R.R.. and Popper, The Behavioral Ecology of Anuran Communicationt in calling each night, but can remain in a chorus for several months (Bevier 1997a: Wells 2001). For these species, the ability to remain active for long periods of time probably is a more important determinant of mating success than nightly calling performance.2.2Vocal Sacs as Sound Radiators and The Behavioral Ecology of Anuran Communication Visual SignalsBecause the metabolic cost of calling is high for many anurans, any adaptation to increase the efficiency of sound transmission will beThe Behavioral Ecology of Anuran Communication
favored by selection. Most anurans that call in air have inflatable vocal sacs that radiate sound to the external environment, providing greater enerGọi ngay
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