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Visual Signaling in Anuran Amphibians

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Visual Signaling in Anuran Amphibians .. Hödl, W. and Amezquita, A. (2001). Visual signaling in anuran amphibians. In: Anuran communication, (M.J. Ryan, ed.). .. Smithsonian lust. Press, Washington. Pp. 121-141. 10 WALTER HÖDL AND ADOLFO AMEZQUITA Visual Signaling in Anuran Amphibians lntroduction cation. social behavior, or natural history. visual signaling was either not considered or was treated as a minor subject Acoustic communication plays a fundamental role in an- (Wells 1977a, 1977b; Arak 1983; Duellman and Trueb 1986; uran reproduction and thus is involved in evolutionary Rand 1988; Halliday and Tejedo 1995; Stebbins and Cohen processes such as mate recognition. reproductive isolation. 1995; Sullivan et al. 1995). The most detailed review ofthe speciation. and character displacement (Wells 1977a. 1977b. subject is now more than 20 years old (Wells 1977b). Never- 1988;Rand 1988;Gerhardt and Schwartz 1995;Halliday and theless some authors have discussed the possible evolution- Tejedo 1995;Sullivan et al. 1995).Visual cues. however. have ary link between visual signaling and the reproductive ecol- been thought to function only during dose-range inter- ogy of species, such as reproduction associated with streams actions (Wells 1977c; Duellman and Trueb 1986). Visual sig- (Heyer et aI. 1990; Lindquist and Hetherington 1996. 1998; naling is predicted to be predominantly employed by diur- Hödl et al. 1997;Haddad and Giaretta 1999) or reproduction nal species at sites with an unobstructed view (Endler 1992). within feeding territories (Wells 1977c). Diurnality. however. is not common for the majority offrog Our aim in this review is (1) to propose a dassmcation of species. Thus vocalizations. which are highly efficient for reported behavioral patterns of visual signaling in frags; (2) communicating at night or in dense vegetation, are by far to describe the diversity of visual signals among living an- the best studied anuran signals (Duellman and Trueb 1986; uran taxa; and (3) to apply a comparative approach to explor- Fritzsch et aI. 1988;HödlI996). Nevertheless there are anec- ing any associations between the diversity of visual signals dotaI reports of apparent visual signaling in intra- or inter- and the ecological conditions in which they may have sexual interactions, or both (for summary see Lindquist and evolved. Hetherington 1996). Although the function of visual cues has only occasionally been tested experimentally (Ernlen Visual Signals and Communication 1968;Lindquist and Hetherington 1996) the reports suggest that visual signaling is a significant mode of communication A communication process involves a transfer ofinformation in at least a few anuran species. from asender to a receiver by means of specmcally designed In both dassic and re cent reviews on anuran communi- signals. Signals are traits that have evolved specifically to 121 ;0, 122 W. HÖOL AND A. AMEZQUITA manipulate the receiver's behavior (Harper 1991; Krebs and nals, for example, the inflation of the gular sac during vocal- Davies 1993; Bradbury and Vehrencamp 1998). Signaling ization and the vibration of flanks du ring ventilation (Rand may occur at both the intra- and interspecific level (Brad- 1988;Zimmermann and Zimmermann 1988). However we bury and Vehrencamp 1998). do not ignore the fact that the accompanying movements Because of the complexity of visual stimuli and signal pro- may provoke changes in the behavior of potential receivers cessing and the difficulty of quanti.fying what animals actu- hearing a call or looking at another individual. In the den- ally perceive, the identification and analysis of visual signals drobatid frog Colostcthuspalmatus, the presence offrog dum- is problematic (Bradbury and Vehrencamp 1998). In addi- mies and simultaneous playback of advertisement calls elic- tion to their information content, features of rituatized sig- its in females low postures associated with courtship that are nals generally involve redundancy, conspicuousness, stereo- intensified when the dummy's gular sac is vibrated; how- typy, and alerting components (Hailman 1977; Harper ever, gular sac vibrations without simultaneous playback of 1991). In general. communication systems that increase the advertisement calls elicits predatory attacks (Lüddecke signal-to-noise ratio will be favored by natural selection 1999). A serious reason to ignore epiphenomena is in many (Endler 1992; Alcock 1998). Conspicuousness in visual sig- cases the trouble of separating, either experimentally or by nals is achieved by enhancing the contrast between the sig- observation, the concomitant effects of the sound or the nals and the brightness, color, spatial pattern, or movement presence of the individual. We also consider as epiphenom- oftheir background (Hailman 1977; Endler 1992; Bradbury ena actions thatindividuals perform to improve their visual - and Vehrencamp 1998).Most reflected light signalsinvolve or acoustic field or to prepare themselves for another be- muscular movement to change body shapes, to perform havioral pattern. For example, in some contexts "upright stereotyped displays or gestures, or to orient or position the posture" or "body elevation" are followed by jumping in individual in space. Often signal-gene rating movements are C. palmatus and may constitute orientation or preparatory accentuated by strikingly colored or structured parts ofthe movements (Lüddecke 1974, 1999). The stereotyped move- body (Bradbury and Vehrencamp 1998). ments at underwater calling sites in pipid frogs (Rabb and We exclusively defioe a visual signal if it is reported or Rabb 1960, 1963) have been classified as "postural or other personally observed that the behavioral event (1) provides a visual displays" (Wells 1977b). However we do not include visual cue during an intra- or interspecific interaction, (2) is these behavioral traits in our considerations since it is not redundant, conspicuous, and stereotypical, and (3) most known whether these movements are performed to pro- likely provokes an immediate response by the receiver that duce mechanical cues that propagate through water or benefits the sender. The presence of alerting components in whether they represent visual signals themselves. visual signals is difficult to prove (Lindquist and Hethering- Additionally we do not classify as signals visual cues that ton 1998). are permanently "on" (Bradbury and Vehrencamp 1998 and Any visually perceivable state (sensu Lehner 1979;Martin references therein), such as coloration or size, even though and Bateson 1993), such as persistent or temporarily limited we cannot rule out the fact that som; states provide sub- sexual dimorphism, is excluded. Thus seasonal variation in stantial information in communication processes. Examples color (as in breeding males of the European brown frog, include aposematic body coloration in dendrobatid frogs, Rana arvalis woltcrsto1jfi,Mildner and Hafner 1990)or shape thought to indicate toxicity to potential predators (Myers (e.g., hypertrophied forelimb musculature and nuptial ex- and Daly 1976; Myers et al. 1978); sexual dimorphism in crescences in sexually mature males; Duellman and Trueb color patterns in Atclopus spp. (Lötters 1996) and Bufo spp. 1986) is not included in our analyses. We ooly considered (Duellman and Trueb 1986), considered to playa role in changes of body coloration as visual signals if they occur intersexual communication; and intermale differences in within short time intervals and immediately affect the be- the amount of ventral pigmentation, correlated with differ- havior of a conspecific receiver. Many postures and, in gen- ences in reproductive success (Burrowes 1997). eral, behavioral states can be used by conspecifics or het- erospecifics as cues that provide information about the Deimatic Behavior and Visual Signaling of status of asender. However they barely satisfy our definition Anurans during Interspecmc Communication of visual signals-that the behavioral change provoked in the receiver by the posture most probably benefits the Signals Addressed toward a Potential Predator sender. The same reasoning applies to many locomotory movements. Some anuran species perform a deimatic behavior consist- Consequently we do not consider visual cues that obvi- ing of intimidating postures or actions when caught by a ously originate as epiphenomena to be rituatized visual sig- pursuing predator (Edmunds 1981). The unken reflex (Un- Visual Signaling in Anuran Amphibians 123 ~ km is the German term for the genus Bombina), a common in same individuals of C. calcaratain the presence of a small behavioral response to perceived threat in the discoglossid leptodactylid frag of the genus Pleurodema. In this case not species Bombina bombina and B. variegata, is characterized oo1y the fourth and fifth toes of a hind foot were curled and primarily by lifting all four legs and arching the back, draw- undulated, but the foot itself was lifted and rotated. As in ing attention to the bright ventral surface. Accompanying C. omata, individuals do not signal to all types of prey. For this posture and subsequent immobility is a slowing down example, the presence of crickets failed to elicit pedalluring of respiratory movements and an increase in skin secretion (Murphy 1976). (Hinsche 1926; Noble 1955; Bajger 1980). The term unken reflexhas been used broadly to describe a wide range of sirn- Visual Signaling in Anurans during Intraspecific ilar defensive postures restricted to species with ventral Interactions warning coloration (for anurans, see Haberl and Wilkinson
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