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Gryllus Texensis) in Response to Acoustic Calls from Conspecific Males

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Gryllus Texensis) in Response to Acoustic Calls from Conspecific Males J Insect Behav DOI 10.1007/s10905-013-9375-7 Phonotactic Behavior of Male Field Crickets (Gryllus texensis) in Response to Acoustic Calls From Conspecific Males Thomas M. McCarthy & John Keyes & William H. Cade Revised: 17 December 2012 /Accepted: 10 January 2013 # Springer Science+Business Media New York 2013 Abstract We studied male phonotactic behaviors elicited by acoustic cues that simulate conspecific male songs in the field cricket, Gryllus texensis. Males exhibited significant positive phonotaxis in response to the simulated song stimuli, but showed no such response to atypical song stimuli. We found no significant relationship between males’ own calling behavior and their phonotactic responses to the stimuli. Analyses indicated that larger males exhibited greater phonotactic responses, which may indicate a greater tendency to engage in aggressive interactions if size is an indicator of fighting ability. Male phonotactic responses were significantly weaker than those exhibited by females, and adult males did not exhibit stronger responses with increasing age as has been documented for females. Observed sex differences in the strengths of phonotactic responses may reflect differences in the fitness-payoffs of responding. That is, females are under strong selection pressure to respond to male songs and subsequently mate. In contrast, males responding to acoustic signals from other males need not precisely locate the signaler but would likely move to areas where females are likely to be found. Alternatively, males might benefit from avoiding areas with calling males and establish- ing their own calling stations away from competing males. Keywords Acoustic signal . phonotaxis . mating system . behavioral strategy . size . orthoptera Introduction A major goal in behavioral ecology is to understand the selective forces that influence the evolution of mating systems. Much effort has gone into increasing our T. M. McCarthy (*) Department of Biology, Utica College, 1600 Burrstone Road, Utica, NY 13502, USA e-mail: [email protected] J. Keyes : W. H. Cade Department of Biological Sciences, University of Lethbridge, Lethbridge, AB T1K 3M4, Canada J Insect Behav understanding of mating processes, and how mating decisions translate into fitness benefits (Bateson 1983; Andersson 1994). Signalling and communication are impor- tant components of many mating systems; signalling facilitates the exchange of information between individuals (both potential mates and rivals) engaged in inter- actions that influence reproductive success. Acoustic signals are among the most extensively studied sexually-selected traits, especially in insects, anurans and birds (Andersson 1994; Gerhardt and Huber 2002). There are likely to be strong selection pressures on both the signallers and the receivers of acoustic cues (Andersson 1994; Johnstone 1997; Gerhardt and Huber 2002): signal- lers must attract mates and dissuade rivals, while those receiving acoustic cues must assess the signal in order to determine either the quality of the signaller as a mate, or the level of risk imposed by the signaller if they are competitive rivals for mates. In crickets, characteristics of males’ acoustic songs have been studied extensively (e.g. Souroukis et al. 1992; Ciceran et al. 1994; Gray and Cade 1999a; Gray and Eckhardt 2001). Additionally, numerous studies have examined females’ phonotactic responses. Male songs generally elicit a positive phonotactic response from females, such that, females move in the direction of the calling male. These studies have shown that females prefer some songs to others (Wagner et al. 1995; Hedrick and Weber 1998; Gray and Cade 1999b; Wagner and Reiser 2000), and that their preferences can be context-dependent (Cade 1979a; Prosser et al. 1997; Lickman et al. 1998). In contrast to the attention focused on female responses, very few studies have considered short-range phonotactic responses of male crickets (e.g. Kiflawi and Gray 2000; Leonard and Hedrick 2009; Jang 2011). Several lines of evidence suggest that males respond to acoustic cues, including: long-range phonotaxis (Cade 1989), spatial distribution patterns of males (French et al. 1986; Cade and Cade 1992; Souroukis and Cade 1993), and alternative male mating strategies (Cade 1981; Cade and Cade 1992; Rowell and Cade 1993; Walker and Cade 2003; Zuk et al. 2006). Consequently, male calls may serve a dual function: to attract females and repel competing males (Alexander 1975; Otte 1977; Cade 1979b; Andersson 1994). However, non-calling males may also eavesdrop (McGregor 1993) and exploit an individual’s calls by employing a satellite- male mating strategy (Cade 1979b, 1981; Cade and Cade 1992; Rowell and Cade 1993; Zuk et al. 2006). These hypotheses lead to specific predictions that can be experimen- tally tested: (1) if competing males are deterred by a calling male, or if calling serves as a mechanism for spatially distributing individuals, then negative phonotaxis should be observed. In various cricket species, calling males are separated by distances greater than 1 m in the field (Cade 1979b). (2) If competing males act agonistically toward a calling male, then positive phonotaxis should be observed. Here, positive male phono- taxis could be an indicator of aggression or competitive ability in a fight (Leonard and Hedrick 2009). Individuals that move toward a calling male might attempt to displace that male and occupy his calling territory. (3) If competing males employ a satellite mating strategy, then either positive phonotaxis or increased non-directed movements should be observed. Positive male phonotaxis may be a mechanism to increase the efficacy of satellite behavior. Silent males roaming in the vicinity of a calling male (Rowell and Cade 1993) should encounter more receptive females than those randomly moving through the environment. Our goals for this study were to determine whether male field crickets exhibit phonotactic responses as has been observed for conspecific females, and if so, J Insect Behav examine the ontogeny of the response and determine whether there is a relationship between male calling behaviors and male phonotaxis. We addressed these questions by exposing individual crickets to synthetic acoustic stimuli and observing their behavioral responses in the laboratory. We predicted that male crickets would exhibit phonotaxis in response to the calls of other males. We also predicted that the responses of males might be state-dependent and change with the age of the individ- ual, as is the case for females (Cade 1979a; Prosser et al. 1997; also see Jang 2011 (G. rubens males)). Finally, we predicted that there would be a significant relationship between calling and phonotactic behaviors of individual males that might reflect alternative male mating strategies. Specifically, we hypothesized that males with low calling rates may adopt satellite mating strategies, and so we predicted that they would have strong positive phonotactic responses. Alternatively, males with high calling rates might respond by moving away from the stimulus source (negative phonotaxis) as a means of spacing between calling males, or by moving toward the stimulus (positive phonotaxis) to displace the calling male. Methods We studied phonotactic behaviors of male field crickets, Gryllus texensis (Cade and Otte 2000). Crickets were raised in laboratory cultures originally founded by wild- caught specimens from Austin, Texas; cultures were supplemented with wild-caught crickets annually (Lickman et al. 1998). The cultures were maintained on a reversed 14:10 light:dark cycle. Individuals were isolated in 500 ml plastic containers and provided food and water prior to use in experiments. Experiment 1: Calling Behavior and Phonotaxis Measuring Calling Behavior We removed newly molted adults from the laboratory cultures daily (within 24 h of adult molt). Adults remained isolated for 7–8 days prior to observations to ensure that they were functionally mature. Males produce spermatophores and begin to call within 3–6 days postecdysis (Cade 1981), and the mean age of females at first mating is 3.6 days (Solymar and Cade 1990). We recorded the calling behavior for each male (N=81) over the course of a single night following the isolation period. This should be a reliable indicator of an individual’s propensity to call as previous work indicated calling duration is a repeatable behavior (Bertram et al. 2007) and is relatively consistent across contexts (Cade 1991). Furthermore, song characteristics do not appear to be condition-dependent (Gray and Eckhardt 2001; but see Hedrick 2005). Observing males for a single night, rather than over multiple nights, allowed us to maximize the number of males that could be observed. Males’ containers were distributed such that they were separated by at least 50 cm during the calling assays (see Cade 1981). Observations were divided into 5-min intervals over the course of 11 h (132 intervals per male): 30 min before the lights turned off, 10 h of darkness, and 30 min after the lights turned on. We recorded whether a male called during a given interval and calculated the proportion of intervals that each male called during J Insect Behav the night. Observations were conducted at temperatures between 25 and 28 °C, and a red-filtered light source was used for illumination. Measuring Phonotaxis We assayed the phonotactic responses of crickets using the Kugel apparatus. The Kugel (described in greater detail by Wagner et al. 1995; Prosser et al. 1997) is a device that measures the relative speed and direction of a cricket’s movement in response to acoustic stimuli played through a series of speakers. Briefly, the device is comprised of a circular anechoic chamber (47.5 cm high, 101.9 cm in diameter) with four speakers (linear, B-4- 5 JO, full range frequency) set at 90° intervals around the wall of the chamber. A plastic sphere (16.2 cm in diameter) floats on a column of air in the center of the chamber, and sensors record the displacement of the sphere as the cricket moves (see Figure 2 in Wagner et al. 1995). Each speaker is 35.4 cm from the sphere. Crickets were tethered in position at the top of the sphere during trials.
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