Johnson and Hupton: Behavioral Syndromes in the Sagebrush Cricket: A Pilot Study to Q 105 BEHAVIORAL SYNDROMES IN THE SAGEBRUSH CRICKET: A PILOT STUDY TO QUANTIFY INDIVIDUAL VARIATION IN MALE CALLING BEHAVIOR J. CHADWICK JOHNSON GINA MARIE HUPTON ARIZONA STATE UNIVERSITY TEMPE ABSTRACT In contrast to this the notion of unbridled plasticity, the field of behavioral ecology has recently The field of behavioral ecology has recently been reinvigorated by the notion of behavioral been reinvigorated by the addition of the notion of syndromes (i.e. animal personalities; reviewed in Sih behavioral syndromes (a.k.a. animal personality). et al. 2004, 2010; Figure 2). A behavioral syndrome Behavioral syndromes imply the existence of is individual variation that is consistent across individual variation in behavioral expression that is distinct functional contexts. For example, Riechert consistent across distinct functional contexts (e.g. and colleagues studied behavioral syndromes in the foraging, mating, anti-predator). The syndromes funnel web spider, Agelenopsis aperta. Funnel web paradigm suggests that the behavioral phenotype is spiders demonstrate positive correlations among best viewed as an integrated phenomenon wherein aggression toward prey, including superfluous killing, any given behavior can only be fully understood by boldness toward predators, aggression toward studying selection pressures in all contexts. Here we conspecifics, and the size of territory defended. report on a pilot study on behavioral syndromes in Controlled crosses revealed a genetic basis (probably the Sagebrush cricket (Cyphoderrris strepitans), an pleiotropy) to these behavioral correlations (see acoustic Orthopteran insect that inhabits high altitude references in Maupin and Riechert 2001 Riechert and sagebrush meadows of Grand Teton National Park. Maupin, 2008). In addition to being correlated at the The results of our preliminary analysis suggest very individual level, these behaviors are also correlated at little consistent repeatability in the mating behavior the population level. Riparian populations of A. of C. strepitans. In addition, we make note of the aperta enjoy six times higher prey availability but synergistic collaboration in our group between faculty researchers and graduate, undergraduate and high school research collaborators. INTRODUCTION The traditional adaptive paradigm suggests that, over time, natural selection acts to drive phenotypes towards an optimal expression (Figure 1). While the evolutionary power of natural selection is not in question, most evolutionary ecologists work from the underlying assumption that many forces constrain adaptation. For example, available sources of genetic variation limit the types of traits that can Figure 1. The traditional adaptive paradigm suggesting be favored by natural selection. extensive adaptive plasticity. Published by Wyoming Scholars Repository, 2011 1 University of Wyoming National Park Service Research Center Annual Report, Vol. 34 [2011], Art. 15 106 and Henriksson 1997). The adaptive foraging hypothesis (Johnson and Sih 2005) explains precopulatory sexual cannibalism by positing that females that had poor foraging success on other prey (e.g., crickets) might be willing to risk the cost of lost reproduction in favor of the benefits of food. Contrary to this hypothesis, Johnson and Sih found that females that most readily attack males are the voracious females that had the highest levels of recent foraging success on other prey. That is, female spiders exhibited positive behavioral correlations between voracity toward heterospecific prey, boldness toward predation, and tendency to attack males. These data are consistent with the hypothesis that precopulatory sexual cannibalism reflects a carryover of voraciousness toward prey, in general, on to voraciousness toward potential mates. Figure 2. The behavioral syndromes paradigm suggesting extensive individual variation and limits to plasticity. Subsequent studies showed that this carryover is not absolute: if a female grew up suffer significantly higher predation risk relative to experiencing low availability of males, then she is their counterparts from grassland populations. As one less likely to engage in sexual cannibalism (Johnson might predict then from a traditional adaptive and Sih 2005). paradigm, spiders from riparian populations are significantly shyer toward predation risk, less Thus, the extent to which animal behavior is voracious toward prey, less agonistic toward driven by adaptive plasticity versus constraint is a conspecifics, and defend smaller territories than pressing question in the current field of behavioral grassland spiders. However, what is most interesting ecology. To address this issue, more studies on more is that within the adaptive modulation going on taxa need to follow individuals through their lifetime: across populations, we continue to see correlations at 1) documenting the repeatability of an individual‘s the individual level. In other words, although riparian behavioral expression within a context, and 2) spiders are, on average, shy toward predation risk, looking for the across –context correlations that are some individuals from riparian populations continue the hallmark of a behavioral syndrome. Here we to show high levels of boldness toward predators, and approach the mating behavior of the sagebrush this individual variation can be explained by the fact cricket, Cyphoderris strepitans, with the behavioral that these bolder individuals are also more voracious syndromes approach. toward prey. Conversely, grassland populations are, on average, more voracious toward prey, but some STUDY SYSTEM individuals continue to show low voracity toward prey, and this can be explained, at least in part, by the C. strepitans is an acoustic Orthopteran fact that these individuals are also exceedingly shy insect from the family Haglidae. Sagebrush crickets toward predators (Riechert and Hedrick 1993). This emerge soon after snowmelt in high-altitude work provides a beautiful illustration of the tug-of- sagebrush meadows. Several populations residing war occurring between adaptation to one‘s within and around Grand Teton National Park have environment at the population level and potential received considerable attention (e.g. Sakaluk 1991, constraints arising from behavioral correlations Sakaluk and Sneddon 1990, Sakaluk et al. 1987, occurring at the level of individuals. 1995, 2004). More recently, Johnson and Sih (2005, Of particular interest is the mating system of 2007), working with the semiaquatic fishing spider C. strepitans. Male sagebrush crickets stridulate Dolomedes triton, investigated a connection between (call) to attract females for mating. Calling song is behavioral syndromes and precopulatory cannibalism produced by a file and scraper mechanism found on by females on courting males. Precopulatory sexual opposing edges of the male‘s calling wings, the cannibalism can be puzzling from an evolutionary tegmina (Figure 3). However, upon mounting a male, point of view particularly when it results in females females begin to chew on a pair of fleshy hind wings not mating enough to fertilize their eggs (Arnqvist located underneath the tegmina (Figure 3). These 2 Johnson and Hupton: Behavioral Syndromes in the Sagebrush Cricket: A Pilot Study to Q 107 Figure 3. A copulating pair. Wingless females mount the male‘s dorsum and feed on the male‘s fleshy hindwings while male‘s struggle to transfer a spermatophore. (Photo: S. K. Sakaluk) Figure 4. A mated male. Males displaying a dried scab over their hindwings can reliably be scored as mated. fleshy hind wings then produce hemolymph which (Photo courtesy of Scott K. Sakaluk) the female consumes while the male is busy transferring an external spermatophore to the female. South and West. We broke this area up into 4 Thus, the mating system of sagebrush crickets is quadrants, each approximately 50 X 100 m. characterized by a non-lethal form of sexual cannibalism—females consume irreplaceable male body parts, but males are not prevented from mating METHODS again in the future (Figure 4). Our 2011 field season ran from June 1 Sexual cannibalism has been argued to be an through June 11. Much of our time in Wyoming was ideal target behavior in the study of behavioral limited by rain, which combined with cold syndromes as it so naturally combines the distinct temperatures can limit calling behavior for the entire functional contexts of foraging and mating behavior night. Nevertheless, we measured the calling into an integrated context (Johnson, 2001). For behavior of 18 different males in the field. After this example, the fishing spider case study discussed initial field measurement we collected all males and above nicely illustrates a case where sexual transported them to the laboratory where they were cannibalism integrates foraging, mating and anti- weighed to the nearest milligram and given a unique predator behavior. Here we present preliminary data I.D. tag secured to the pronotum. Marked males designed to measure the repeatability of male calling were then returned the next day to the base of the behavior in the field. As per the behavioral bush they were found calling from. On subsequent syndromes hypothesis we predicted we would find nights we recorded repeated measures of the calling repeatable individual variation (behavioral types) in behavior of these males (N=15 males scored twice; calling behavior. N=9 males scored three times; N=6 males