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Emergence times (X ̄ € 1 SE) from a refuge of male Lacerta monticola lizards in presence or absence (control) of a tethered female after being approached directly by an experimenter at one of two different approach speeds (slow vs fast) in the mating season
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Because time spent in refuge may be costly if prey lose opportunities to forage, fight, or mate, prey allow predators to approach closer before beginning to flee when opportunity costs are high. Because the same opportunity costs may apply to refuge use as to escape, prey should make similar trade-offs between risk of emerging and cost of remaining...
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... a predatory attack, males emerged significantly earlier from the refuge when a female was present than in the control situation (two-way ANOVA, female effect: F =85.55, P <0.0001) ( Fig. 2), and emerged significant- ly earlier when the approach was made at slow speed than when it was at fast speed (speed effect: F 1,76 =19.67, P <0.0001). The interaction between female presence and predator speed was not significant ( F 1,76 =2.71, P =0.10). However, post-hoc tests indicated that, in the control situation, males emerged significantly earlier when the speed was slow (Tukey’s HSD test, P =0.0004), but that emergence times did not differ significantly between approach speeds when a female was present ( P =0.21). The approach speed did not significantly affect the propensity of the male to return and court the female after the predatory attack (16 vs 10 males returned to court after being approached rapidly or slowly, respectively; 2- tailed binomial test P =0.33). Emergence times did not differ significantly between males that returned to court (X ̄ €SE=10.0€1.4 s, n =26) and males that did not return (8.7+1.6 s, n =14) (ANOVA: F 1,38 =0.16, P =0.69). When males returned to court the female, latency to resume courtship after emerging from the refuge did not differ significantly between predator approach speeds (fast: X ̄ €SE=4.7+0.8 s; slow: 3.6+1.0 s; ANOVA: F 1,24 =0.21, P =0.65). When a male bit a female during courtship, it might be considered that the interest of this male in the female was greater than when a male licked the female, but did not attempt to mount her. All males that had bitten the female returned to court her after the attack. However, emergence times did not differ significantly between males that bit and those that did not bite the female (two-way ANOVA, bite effect: F 1,36 =0.26, P =0.31) or between approach speeds (speed effect: F 1,36 =1.09, P =0.30), and the interaction was not significant ( F 1,36 =1.03, P =0.30). Nevertheless, in a separate analysis, emergence times did not differ significantly between approach speeds for males that bit the female (fast: X ̄ €SE=7.8+1.7 s; slow: 7.5+1.4 s; ANOVA: F 1,7 =0.001, P =0.98), but for males that did not bite females, emergence times were significantly longer for fast than slow approaches (fast: X ̄ €SE=12.9+2.2 s; slow: 7.5+1.4 s; ANOVA: F 1,29 = 4.22, P <0.05). The different levels of predation risk posed by a predator approaching at different speeds strongly affected latency to emerge from refuge. Rapid approach by a predator is an important indicator of immediate threat of predation to which prey in many species respond by increasing both the flight-initiation distance (e.g., Dill 1974; Martín and López 1996; Cooper 1997a; but see Bonenfant and Kramer 1996) and latency to emerge from refuge (Cooper 1998; Martín and López 1999c). Many animals, such as some fishes (Endler 1987; Candolin 1997), frogs (Ryan et al. 1982), crabs (Koga et al. 1998), or aquatic insects (Sih et al. 1990), may reduce mating activity under predation risk. However, when there is a trade-off between mating opportunities and predator avoidance, the degree of risk taking should be related to the probability of future mating opportunities (Magnhagen 1991; Clark 1994). Thus, for example, young Gobius niger fishes stopped spawning under predation risk, whereas older individuals readily spawned in a similar situation (Magnhagen 1990). Similarly, in L. monticola, where the mating period is very short, sex and season also affected decisions about refuge use, as indicated by the significant interaction term. During the mating season, males had shorter emergence times than in the postreproductive season. Male L . monticola also have greater general activity levels during the mating season and these seasonal differences cannot be explained by thermal constraints or changes in microhabitat use (Aragón et al. 2001). Thus, when mating opportunities are high, males seem to be less sensitive to predation risk, at least in a situation where the predator has disappeared after an unsuccessful attack. Nevertheless, it may be expected that an increase in risk may force prey to increase emergence times. Thus, in the second experiment during the mating season, when we increased risk by remaining close to the refuge, emergence times were longer and were affected by approach speed in the control treatment, but not in the female treatment. Therefore, variations in emergence times seem to reflect different balances between the costs of losing mating opportunities and the benefit of a diminution of predation risk with time. The absence of any effect of season on refuge use of females may be explained in part by the greater reproductive investment of females (Clark 1994) and by the lack of a relationship between reproductive success and number of matings. Whereas males can increase the number of eggs fertilized by mating with multiple females, females may fertilize all their eggs by mating once. Thus, females should risk less than males. Also, during the mating season, female L . monticola remain relatively stationary, but males must search for females (Martín and Salvador 1997; Aragón et al. 2001). Thus, females do not lose reproductive opportunities by increasing refuge use. Nevertheless, it might be expected that gravid females later in the year have greater latency to emerge than males or nongravid females because their risk of predation is greater in many species due to decreased sprint speed, which impairs their ability to escape (Shine 1980; Cooper et al. 1990; Magnhagen 1991). Gravid lizards are known to modify other aspects of antipredatory behavior and may be predicted to spend more time in refuge unless the increased risk is outweighed by decreases in body temperature while in refuge (Martín and López 1999b, 1999c) at a time when basking may be important for development of offspring. Thus, the concurrence of different ecological pressures may lead females to have similar optimal emergence times in the different phases of their reproductive cycle. When a tethered female was presented, males had even shorter emergence times, and these did not differ between approach speeds. When a female is nearby, staying in the refuge may result in losing opportunities to mate. A similar situation was observed in mate-guarding males of the skink Eumeces laticeps (Cooper 1997a, 1999) and the lacertid lizard Psammodromus algirus (Martín and López 1999a), which allowed closer approach than males found alone, before escaping from potential predators. Because costs of fleeing and refuge use increase for a male that must leave his mate to escape, an optimal decision requires that the approach distance and emergence time decrease even if predation risk increases (Ydenberg and Dill 1986; Martín and López 1999a). A male leaving a female not only loses mating opportunities, but also risks sneaked copulations by subordinate males if they are not able to relocate their mates immediately after the predatory attack. Risk of loss of copulations to rivals may be high in dense populations such as the one we studied (Martín and Salvador 1997; Aragón et al. 2001). Risk of losing mating opportunities due to inability to relocate the female may be especially great for transient females and unfamiliar females (Cooper 1985), in which case our results may indicate maximum effects of female presence. Return by males to court tethered females despite the proximity of the predator indicates the willingness of males to take risks to obtain mating opportunities. The predator’s approach speed influenced neither a male’s decision to return to court, nor the latency to resume courtship after emergence. Furthermore, in males that had already attempted mounting (i.e., those that had bitten the female), the value of the female was probably assessed as greater, and emergence times decreased regardless of approach speed. Taken together, the results of both experiments suggest that males adjust emergence times simultaneously to the degree of predation risk and the cost of lost mating opportunities. Therefore, refuge use in the lizard L . monticola reflects a trade-off between survival and ...
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Citations
... We observed a sexual difference in shelter use in P. melisellensis, with males using shelter less than females, that is, engaging in more risky behaviour. This contrasts with the risk-taking behaviours in male P. siculus, and agrees with studies that suggest that antipredator and courtship strategies in males can covary, that is, males usually engage in riskier behaviour to enhance their reproductive success (Clark et al., 2016;Cooper Jr., 1999;Hoefler et al., 2008;Martín et al., 2003;Uzendoski et al., 1993). The relevance of interspecies sexual differences in risk-taking behaviours for the invasive potential of a species is still insufficiently understood. ...
Invasive species are a major threat to biodiversity especially because of their influence on native species. Their behaviours, including those in nonsocial contexts, significantly influence species fitness, drive invasion processes and can result in the competitive exclusion of less dominant species. Understanding these behavioural patterns is vital for the protection of native species and the control of invasive species spread. There is still a controversy regarding the success of invasive species, whether it is attributed to inherent behavioural traits or to adaptations to new environments. This study focused on native populations of two sympatric lizard species in Croatia: a lizard known for its invasiveness, Podarcis siculus, and the eastern Adriatic endemic Podarcis melisellensis. We evaluated nonsocial behaviour associated with competition, including food consumption, exploratory behaviour and risk-taking behaviours, using two primary apparatuses: an open field and a radial maze. Podarcis siculus exhibited higher levels of exploratory behaviour, reduced risk-taking behaviours and greater food consumption than P. melisellensis. These behavioural traits are consistent with behaviour observed in invasive populations of P. siculus, suggesting that they may be intrinsic to the species. Our results indicate that a successful invader is more likely to explore and at the same time be more cautious. Research on behavioural traits in native populations sheds light on preadaptive features of invasive species that could characterize a successful invader, highlighting the importance of researching invasive species in their native range.
... Interspecies mating behaviour in nature has been supported previously by visual observations for males of Darevskia valentini with females of Darevskia armeniaca and Darevskia unisexualis (Galoyan 2013). Mating scars on the venters of lacertid females occur after male mating attempts (Martín et al. 2003) and confirm sexual interest of males towards females. Carretero et al. (2018) attempted to analyse the number of (D. armeniaca and D. unisexualis), and triploid hybrids occur (Carretero et al. 2018). ...
Parthenogenesis has been discovered in a great number of reptiles, indicating that parthenogens can outcompete bisexual species and hybridize with them. This raises intriguing questions: how do parthenogenetic species coexist with gonochoristic relatives, and what outcome does this coexistence produce? To answer these two questions, we studied relationships between male rock lizards, Darevskia valentini, and parthenogenetic Darevskia armeniaca. We compared social and sexual reactions of males to females of their own species and parthenogenetic D. armeniaca and vice versa. We found that females of both species basked together with males and demonstrated mostly submissive behaviour. Females of D. valentini were less aggressive towards males in the mixed-species population than in the single-species population, suggesting female competition for males. The proportion of successful and unsuccessful mating attempts with females of bisexual and unisexual species was equal, and we found no significant differences in male reactions to bisexual and unisexual females in sympatry. The proportion of females bearing mating scars on their venters and the number of these scars per female were greater in the bisexual species. This indicates that females of the bisexual species were more attractive than parthenogenetic females as mating partners to males. We found no triploid hybrids among the offspring of parthenogenetic females that mated with males in the mixed population. Our findings suggest that parthenogenetic species might outcompete bisexual species by mating with males and integrating into the social system.
... Antipredator behavior is often at odds with other behaviors important to an animal's survival and fitness (Ydenberg and Dill 1986). When predators are present, foraging efficiency may decrease as antipredator responses become more pronounced (e.g., Milinski and Heller 1978;Sih 1980;Mateo 2007) and reproductive behavior is often reduced (Herczeg et al. 2008;Brunel-Pons et al. 2011;Lea and Blumstein 2011;Martín et al. 2003). The balance between accepting risk to gain long-term fitness benefits versus prioritizing immediate survival can vary among species, individuals, and the sexes (Ydenberg and Dill 1986;Samia et al. 2016). ...
... Zootoca vivipara females thermoregulate closer to preferred temperatures than males due to their greater risk-taking behavior, which may be due to the greater energetic needs of viviparous females in their high-altitude habitats (Herczeg et al. 2008). By contrast, male Lacerta monticola emerge from refugia sooner than females (Martín et al. 2003), and male Sceloporus virgatus have lower flight initiation distances than females in the presence of a potential predator (Cooper and Wilson 2007). Our own findings suggest that the thermal costs accrued by male A. aquaticus may limit their reproductive potential more severely than that of females. ...
Males and females often differ in use of antipredator behaviors, particularly when antipredator behavior comes at the cost of missed mating opportunities or territory defense. When using thermally suboptimal refugia, ectotherms are especially vulnerable to these costs, as their performance is linked to body temperature. To flee from predators, semi-aquatic Anolis lizards dive underwater for long periods and rebreathe from a bubble of air. We hypothesized that using aquatic refugia would result in body heat loss, that dive duration is influenced by sex, and that oxygen consumption when diving would help explain sex differences. We tested these hypotheses by measuring dive length and body temperatures in A. aquaticus, and by recording oxygen consumption and final oxygen partial pressure during controlled dives in several semi-aquatic Anolis species. Not only was there a significant thermal cost to diving, but A. aquaticus males and females appeared to tolerate different levels of this cost: males re-emerged from water more quickly and at higher body temperatures than did females. Body temperature decreased according to an exponential decay function, dropping up to 6 °C in 5 min. Oxygen consumption rates in semi-aquatic anoles were primarily explained by the expected allometric scaling relationship with mass and, therefore, are unlikely to lead to sex differences in physiological limits to dive times. Instead, shorter male dives may help them maintain physiological performance, mating opportunities or territory defense. Antipredator diving behavior is physiologically costly but undoubtedly beneficial to both sexes, highlighting the need for further study of sex-based antipredator optimization.
Significance statement
To avoid predators, semi-aquatic Anolis lizards can dive underwater and remain there for an extended time by rebreathing a bubble of air over their heads. In this study, we reveal that diving to escape predators also comes with a cost: submersion in water reduces lizard body temperatures. Reduced body temperature can impair a lizard’s ability to move quickly and defend mates or territories, suggesting that there may be divergent diving behaviors in males and females. Our findings confirm that males do indeed spend less time underwater than females. We measured oxygen consumption during dives, and our data suggest that sex differences in diving behavior are unrelated to oxygen use. This study sheds light on the sex-specific balance of antipredator behaviors and the maintenance of optimal body temperatures, and more broadly contributes insight into adaptive responses to environmental challenges.
... In the study of the Iberian rock lizard, havioral changes [11]. In the study of the Iberian rock lizard, antipredator behavior caused the male individuals to spend antipredator behavior caused the male individuals to spend more time in refuge and ultimately lose their mating oppor-more time in refuge and ultimately lose their mating opportunities [12]. tunities [12]. ...
... In the study of the Iberian rock lizard, antipredator behavior caused the male individuals to spend antipredator behavior caused the male individuals to spend more time in refuge and ultimately lose their mating oppor-more time in refuge and ultimately lose their mating opportunities [12]. tunities [12]. ...
It has been recognized in many studies that wildlife tourism practices might generate a negative impact on wildlife, particularly during the reproductive period. Some wildlife may lower their sensitivity towards tourist presence, for instance in Komodo. Understanding to what extenthabituation occurs in Komodo would be necessary for tourism management in Komodo National Park (KNP). Therefore, this study aimed to identify the response of Komodo to tourist presence during mating and nesting activities. The observation was conducted in Loh Buaya, which is one of the tourism sites in KNP. Komodo’s responses were divided into (1) avoidance; (2) neutral; and (3) aggressive under categorized stimulus: tourist number (i.e., < 5 persons; 5-10 persons; and > 10 persons) and distance (i.e., < 5 m; 5-10 m; and > 10 m). Correlation analysis was performed to identify any influences on mating and nesting activities. Our results revealed that Komodo inhabiting tourism facilities havebeen habituated to tourist presence. Different tourist frequencies did not influence Komodo mating activities (r(20)= 0.036, p=0.873), the nest preparing activity (i.e., digging proportion; r(22)= 0.054, p=0.803) and the guarding activity (i.e., nesting proportion; r(22)= 0.314, p=0.135).Nevertheless, our results indicated possible impacts due to tourism activities and its supporting facilities, such as dominated mating pairs, threats to female reproductive success, and human-Komodo conflicts. Therefore, habituation evi- dence must be carefully considered in order todevelop more corresponding strategies and achieve sustainable tourism practices.
... Males in the spider Nephila fenestrata are relatively safe from sexual cannibalism if approaching a feeding female, which is what they prefer to do (Fromhage & Schneider, 2005). Male rock lizards (Lacerta monticola), too, appear to trade-off mating opportunities against predation risk: following a simulated predator attack, they re-emerged more quickly from their Biological Reviews (2023) refuge when mating opportunities seemed good (Martin, Lopez & Cooper, 2003). Because both components of Equation (11) may depend on state, the same is likely to be true of the optimal policy; senescent individuals may avoid activities that require a highly functioning body to mitigate risk [i.e. for which M(u) is forbiddingly high with weakened sense organs and muscles]; or they may make a terminal reproductive investment because they have little to lose [low V (Gadgil & Bossert, 1970;but see McNamara et al., 2009)]. ...
An animal's behaviour can influence many variables, such as its energy reserves, its risk of injury or mortality, and its rate of reproduction. To identify the optimal action in a given situation, these various effects can be compared in the common currency of reproductive value. While this idea has been widely used to study trade-offs between pairs of variables, e.g. between energy gain versus survival, here we present a unified framework that makes explicit how these various trade-offs fit together. This unification covers a wide range of biological phenomena, highlighting similarities in their logical structure and helping to identify knowledge gaps. To fill one such gap, we present a new model of foraging under the risk of predation and damage accumulation. We conclude by discussing the use and limitations of state-dependent optimisation theory in predicting biological observations.
... Tariel et al. (2020) also found that both parental and developmental risk environment affect personality and behavioural plasticity in Physa acuta (Tariel et al., 2020). Apart from foraging, animals also face conflicting demands for mating and antipredator behaviour (Martín et al., 2003;Sih, 1994). Movement and interaction among conspecifics are the first steps towards finding a mate. ...
Predation is a major force that shapes prey phenotypes, species interactions and ecological communities. Prey respond to variation in predation risk and intensity by either increasing or decreasing vigilance and balancing energy requirements under high- or low-risk scenarios. We test the possible mechanism of sensory adaptation and decision making while responding to risk in the pond snail Lymnaea stagnalis. We maintained three risk treatment groups (no risk, low risk and high risk) for 20 days before testing them under multiple cognitive and behavioural challenges involving a predatory stimulus. Under a configural learning procedure where snails learned to forgo feeding when encountering food in the presence of a predatory threat, the high-risk group snails did not show learning/memory formation. However, when high-risk group individuals were transferred to a no-risk environment for 20 days, they were able to form configural learning memory. In a social interaction trial, snails from the no-risk group reduced interactions under threat whereas the high-risk group showed no change between risk environments. All risk groups responded similarly to an operant conditioning training under predation threat that enhanced memory. The decreased antipredator response of the high-risk group in comparison to the low-risk groups shows that snails are able to respond to risk based on past experience, especially when there exists a trade-off between survival and antipredator response. The variation in response to predation threat under different circumstances provides evidence of decision making rather than sensory adaptation as the mechanism involved in prey response. Our study combines multiple phenotypic responses to elucidate animal decision making under different intensities of risk and highlights the importance of experience in shaping antipredator responses.
... In addition, many animals experience limited opportunities to find mates during the period of breeding (Székely et al. 1999, Segoli et al. 2006, Reading & Backwell 2007, Stenzel et al. 2011). Under such circumstances, antipredator responses might be relaxed as a trade-off between reproduction and antipredator behaviour (Cooper 1999, Martín et al. 2003. Conversely, as animals are often exposed to higher predation risk while breeding (Lima & Dill 1990, Magnhagen 1991, their antipredator response could be enhanced to successfully protect their offspring and themselves (Ciuti et al. 2008). ...
... This probably occurs because when females are present, males might lose reproductive opportunities if they immediately flee from a predator. In fact, the few studies that have evaluated whether the presence of a female influences male anti-predatory behaviors have indicated that males adjust their escaping behaviors in response to the female presence (Cooper 1999(Cooper , 2009Martín and López 1999;Martín et al. 2003). However, such studies often do not consider two factors. ...
... The first is that males may delay their escaping behaviors because the presence of another individual may distract the male and increase the time needed to detect a predator (Dukas 2004). The second, and more important point, is that some studies do not consider other factors that may affect male reproductive output, such as male or female condition or reproductive status (Cooper 1999(Cooper , 2009Martín and López 1999; but see Martín et al. 2003;Reaney 2007;Kopena et al. 2015). In particular, the female reproductive status may be an important factor affecting male escaping decisions because, in many species, females reproduce cyclically, being fertile and/or sexually receptive during a short period of the year (Ziegler et al. 2000;Vitt and Caldwell 2014). ...
... For the female exposure treatment, after the 3 min habituation period, we placed a female 50 cm apart from the males, a distance permitting the visual detection of females by males (see Martín et al. 2003;Quintana and Galdino 2017). We collected females in rock outcrops that were apart from those where we were conducting the experiments, ensuring that the females were not neighbors to the experimental males. ...
Studies that test the optimal escape theory often show that males base their escape strategies on the costs and benefits of escaping. Consequently, some studies have shown that males exhibit riskier anti-predatory strategies when they are near females, probably to avoid losing reproductive opportunities. However, since in some species females have a limited reproductive season, this reduction in mating opportunities should be more pronounced during the female reproductive season. Therefore, males should express reduced anti-predatory behaviors when females are fertile. In this study, we used the lizard Eurolophosaurus nanuzae as study subject to evaluate the hypothesis that during the female reproductive season, males will express riskier anti-predatory behaviors than in the non-reproductive season. To accomplish this, we recorded the flight initiation distance (FID) and the time spent in the refuge of males with and without previous exposure to a female, during both the female reproductive and non-reproductive seasons. We found that after exposure to females, males decreased their FID. Although this effect occurred in both seasons, it was stronger during the female reproductive season. Males also spent less time in refuges when females were sexually receptive. Our results indicated that the presence of females induces males to adopt riskier escape decisions, and that the assumed risks are even greater when females represent an immediate chance to mate.
Significance statement
During or immediately after a predatory event, prey must consider the trade-off between the costs and benefits of escaping. In this context, it is known that males of many prey species tend to take a greater predatory risk in the presence of females to avoid losing mating opportunities when fleeing. But only female presence may not be an important determinant of male escaping decisions if females are not always fertile. Here, we did field experiments to show that males assume a higher predatory risk if they had a previous contact with females during the female reproductive season when compared to the non-reproductive station. Thus, our study shows the importance of considering more nuanced reproductive benefits for a better understanding of the escape decisions made by males.
... Searching for mates and mating behavior is also costly, in terms of time and energetic expenditure, and are conspicuous activities that also increases predation risk (Koga et al. 1998;Magnhagen 1991;Engqvist et al. 2015). Many animals modulate their investment in reproductive behavior according to predation risk level and the future opportunities for mating (e.g., Candolin 1997;Koga et al. 1998;Hazlett and Rittschof 2000;Martín et al. 2003). By decreasing investment in mating behavior, D-males might reduce energetic costs and predation risk and increase growth and survival, but, again, at the cost of a lower mating success. ...
Investing in the current reproduction requires diverting energy resources from other metabolic functions, which may compromise future reproduction and lifespan. To solve this trade-off, an individual may consider its labile state to decide how much to invest in current reproduction. We tested experimentally whether the “state quality” of male rock lizards influences their reproductive strategies. To improve the nutritional status of males before the mating season, we captured and supplemented experimental males (N = 20) with dietary vitamin D3 (an essential nutrient for lizards) and had a control group of males (N = 20). Then, we released all these males and females (N = 31) in a large semi-natural outdoor enclosure where lizards could interact and mate freely during the mating period. Activity levels of males did not vary between treatments, but supplemented males started fewer intrasexual agonistic interactions and made fewer mating advances to females. When the mating season ended, we incubated eggs laid by females to obtain the offspring and estimated the paternity of males using DNA microsatellites. Supplemented males sired fewer offspring than control males. These results suggest that vitamin D3 supplemented males used a low risk/less costly mating strategy to protect their assets (i.e., vitamin D reserves), but that still resulted in “some” current reproductive success, while likely increasing longevity and the expected future total reproductive success.
... Such results have been previously observed in fiddler crabs (e.g. Gruber, Kahn, & Backwell, 2019;Reaney, 2007) as well as other taxa including hermit crabs (Hazlett & Rittschof, 2000), lizards (Cooper, 1999;Martín, L opez, & Cooper, 2003) and birds (e.g. Habig, Chiyo, & Lahti, 2017;Kålås, Fiske, & Saether, 1995). ...
Behavioural thermoregulation can ameliorate thermal stress but is costly. For species that court in a thermally stressful microhabitat, sexual selection via endurance rivalry favours individuals that are able and willing to endure harsh conditions in the courtship habitat, as retreats to a thermal refuge will reduce mating opportunities. The relative costs and benefits of refuge use versus continued courtship in the face of thermal risk, which vary across abiotic and biotic contexts, determine the optimal behavioural strategy. We examined the social and abiotic factors driving behavioural decisions related to thermoregulatory retreat in the fiddler crab Austruca mjoebergi. Male fiddler crabs perform a courtship display on the thermally stressful intertidal sediment surface. Time on the surface, and thus time available for display, was limited by high temperatures; as temperature increased, surface time decreased. Yet when presented with a stimulus female, males were more likely to perform the courtship display, displayed at a higher rate and increased time spent on the surface. These results demonstrate that behavioural decisions related to thermal retreat depend both on the abiotic conditions that influence the degree of thermal stress and on the social conditions that influence the reproductive prospects of the individual.
