风险对两种南非蜥蜴逃跑启始距离和逃避策略的影响

逃避理论预测,不逃跑若增大适合度代价则导致逃跑启始距离加长,逃跑若增大代价则导致逃跑启始距离缩短。逃跑路径和去向等受生境结构影响。作者通过模拟捕食者逼近研究喀拉哈里树石龙子(Trachylepis sparsa)和黑环蜥(Cordylus niger)逃避策略和风险因子对逃跑启始距离的影响。与迂回逼近相比较,直接逼近不仅提高蜥蜴逃跑几率还能缩短其逃跑启始距离。喀拉哈里树石龙子在两种逼近方式下的逃跑启始距离有显著差异,这种差异对黑环蜥而言是边缘性的。喀拉哈里树石龙子以树为避所,树上个体可逼近的距离短于地面个体;快速逼近地面个体的逃跑启始距离比慢速逼近更长。习惯于有人环境的黑环蜥逃跑启始距离比人迹罕至环境中的个体更短。地面喀拉哈里树石龙子多遁至树上而很少逃入倒木或倒伏编巢中。树上喀拉哈里树石龙子通常奔逃至远侧和高处,有时遁入树洞或编巢中;黑环蜥则逃入石缝中。所有发现都证实逃避理论中有关逃跑启始距离的预测。逃跑策略的种间差异表明每一种蜥蜴都利用其生境中逃跑路径和避所的有利条件。在风险不同的生境中,生境结构可影响逃跑启始距离,似乎对逃跑策略亦有重要影响。     

Optimal flight initiation distance

Decisions regarding flight initiation distance have received scant theoretical attention. A graphical model by Ydenberg and Dill (1986. The economics of fleeing from predators. Adv. Stud. Behav. 16, 229-249) that has guided research for the past 20 years specifies when escape begins. In the model, a prey detects a predator, monitors its approach until costs of escape and of remaining are equal, and then flees. The distance between predator and prey when escape is initiated (approach distance = flight initiation distance) occurs where decreasing cost of remaining and increasing cost of fleeing intersect. We argue that prey fleeing as predicted cannot maximize fitness because the best prey can do is break even during an encounter. We develop two optimality models, one applying when all expected future contribution to fitness (residual reproductive value) is lost if the prey dies, the other when any fitness gained (increase in expected RRV) during the encounter is retained after death. Both models predict optimal flight initiation distance from initial expected fitness, benefits obtainable during encounters, costs of escaping, and probability of being killed. Predictions match extensively verified predictions of Ydenberg and Dill's (1986) model. Our main conclusion is that optimality models are preferable to break-even models because they permit fitness maximization, offer many new testable predictions, and allow assessment of prey decisions in many naturally occurring situations through modification of benefit, escape cost, and risk functions.     

Determinants of sexual differences in escape behavior in lizards of the genus Anolis: a comparative approach

Males and females are known to differ in a whole suite of characteristics,such as morphology, physiology, ecology, and behavior. Intersexualdifferences are generally believed to arise because of differencesin selective pressures on either sex. In this study, we investigatedwhether intersexual differences in escape behavior exist inlizards of the genus Anolis, and whether these could be explainedby intersexual differences in body size and/or microhabitatuse. To do so, we compared the behavioral response to an approachinghuman predator in the field in males and females of 12 Anolisspecies. We found that ecomorphs and sexes differ greatly withrespect to escape behavior. Twig anoles have the shortest approachdistance (i.e., distance between the observer and the lizardwhen it starts fleeing) and final distance (i.e., distance betweenthe observer and the lizard when it stops moving), comparedwith the other ecomorphs. The distance fled, on the contrary,is greatest in twig anoles. Also, females flee less soon andrun over shorter distances than do males. Since twig anolesare considered the most cryptic anoles, and females may be lessconspicuous than males, these results corroborate the idea thatwell camouflaged animals allow predators to come closer. Theinterspecific variation in sexual dimorphism in escape behavior,however, cannot be explained by the interspecific variationin sexual size dimorphism or sexual dimorphism in microhabitatuse. Thus, escape behavior appears determined by different factorsin males and females.     

Flight initiation distance and escape behavior in the black redstart (Phoenicurus ochruros)

There are many anti‐predatory escape strategies in animals. A well‐established method to assess escape behavior is the flight initiation distance (FID), which is the distance between prey and predator at which an animal flees. Previous studies in various species throughout the animal kingdom have shown that group size, urbanization, and distance to refuge and body mass affect FID. In most species, FID increases if body mass, group size or distance to refuge decreases. However, how age and sexual dimorphism affect FID is rather unknown. Here, we assess the escape behavior and FID of the black redstart (Phoenicurus ochruros), a small turdid passerine. When approached by a human, males initiated flights later, that is allowing a closer approach than females. Males of this species are more conspicuous, and therefore, may exhibit aposematism to deter potential predators or are less fearful than females. Additionally, juveniles fled at shorter distances and fled to lower heights than adults. Lastly, concerning escape strategy, black redstarts, unless other passerine birds, fled less often into cover, but rather onto open or elevated spots. Black redstarts are especially prone to predation by ambushing predators that might hide in cover. Hence, this species most likely has a higher chance of escaping by fleeing to an open spot rather than to a potentially risky cover.     

Nuptial Coloration and Mate Guarding Affect Escape Decisions of Male Lizards Psammodromus algirus

Males of many species show conspicuous breeding colours that are important for status signalling, but that may decrease crypsis and increase predation risk. However, prey may adjust their escape response, such that the optimal distance at which an animal starts to flee (approach distance) would be the point where the costs of staying exceed the cost of fleeing. We examined in the field the escape response of Psammodromus algirus lizards, to test the hypothesis that more conspicuous old males, showing orange nuptial coloration on most of the head, which presumably have a higher probability of being detected (i.e. increased costs of staying), have longer approach distances, independent of other environmental variables. As predicted, old brightly coloured males had significantly longer approach and flight distances than young dull males, and young males had longer ones than females. In contrast, neither the distance to the nearest refuge nor the air temperature when the lizards escaped were significantly different. In addition, although male and female lizards differed in their use of microhabitats, old and young males did not differ. We also found that old males guarding a female (i.e. increased costs of fleeing) had shorter approach distances than old males that were found alone.     

Pre- and post-copulatory sexual selection increase offspring quality but impose survival costs to female field crickets

Whether sexual selection increases or decreases fitness is under ongoing debate. Sexual selection operates before and after mating. Yet, the effects of each episode of selection on individual reproductive success remain largely unexplored. We ask how disentangled pre- and post-copulatory sexual selection contribute to fitness of field crickets Gryllus bimaculatus. Treatments allowed exclusively for (i) pre-copulatory selection, with males fighting and courting one female, and the resulting pair breeding monogamously, (ii) post-copulatory selection, with females mating consecutively to multiple males and (iii) relaxed selection, with enforced pair monogamy. While standardizing the number of matings, we estimated a number of fitness traits across treatments and show that females experiencing sexual selection were more likely to reproduce, their offspring hatched sooner, developed faster and had higher body mass at adulthood, but females suffered survival costs. Interestingly, we found no differences in fitness of females or their offspring from pre- and post-copulatory sexual selection treatments. Our findings highlight the potential for sexual selection in enhancing indirect female fitness while concurrently imposing direct survival costs. By potentially outweighing these costs, increased offspring quality could lead to beneficial population-level consequences of sexual selection.     

Female egg dumping and the effect of sex ratio on male egg carrying in a coreid bug

Phyllomorpha laciniata Vill (Heteroptera, Coreidae) is uniqueamong terrestrial insects in that females glue eggs on the backsof other conspecifics. Egg carrying byP. laciniatamales haspreviously been considered as paternal care. We explored femaleoviposition with respect to previous mating experience of femalesand tested whether sex ratio affects male egg-carrying. Thehypothesis that male egg-carrying is a form of paternal carepredicts that a male should always accept eggs after matingwith a female. However, if male egg-carrying is a form of postcopulatorymate guarding rather than paternal care, egg carrying shouldincrease in the presence of other males. When two couples wereplaced together, females laid eggs on the backs of all individualsenclosed, including the backs of other females. However, whena female was accompanied by 2 males, 22 out of 26 females ovipositedon their mating partner. Thus, sexual competition rather thanpaternity alone, affects a male's eagerness to carry eggs. However,even if males sometimes carry their own eggs, females lay eggson the backs of all conspecifics they can easily acquire. Thus,egg carrying in P. laciniata is partially voluntary and partiallythe result of female egg dumping     

Maintenance of variation in sexually selected traits in females: a case study using intrasexual aggression in tree swallows Tachycineta bicolor

A fundamental question in evolutionary biology is how phenotypic variation is maintained in the face of selection that ought to deplete that variation. Much research has investigated this question in traits favored via sexual selection in males, with a common solution implicating the condition dependence of sexually selected phenotypes. Despite growing interest in sexual selection on females, it is not clear if the same mechanisms maintain variation in female ornaments, weaponry or other female behaviors targeted by sexual selection. An important step in testing condition dependence in females is thus to identify whether sexually selected female phenotypes are associated with condition and also with potential costs. Here, I examine these two components of condition dependence for a sexually selected behavior, intrasexual aggression, in female tree swallows Tachycineta bicolor. I asked whether high levels of intrasexual aggression map onto natural variation in female condition and whether aggression is associated with one potential behavioral cost: performance in a vertically challenging test of flight. More aggressive females were heavier for their body size, heavier for their wing size and showed decreased flight ability, relative to less aggressive females. These findings are consistent with condition dependence, where only females in better condition are able to be highly aggressive. The association between high aggression and reduced flight ability may result from the additional lift required to power these relatively heavier birds. These associations between natural variation in aggressive behavior, morphology and flight ability are consistent with condition dependence because they confirm two basic assumptions of condition dependence: a link between aggression and condition, and a link between aggression and a behavioral cost, the speed of escape flight. As the first study to examine these assumptions for a conspicuous behavior favored by intrasexual selection in females, this study suggests broad relevance of condition dependence.     

Cross-generational effects of sexual harassment on female fitness in the guppy

Sexual harassment is a common outcome of sexual conflict over mating rate. A large number of studies have identified several direct costs to females of sexual harassment including energy expenditure and reduced foraging ability. However, the fitness consequences of sexual harassment for descendants have rarely been investigated. Here, we manipulated the level of sexual harassment and mating rate in two groups of female guppies, Poecilia reticulata, a live-bearing fish in which sexual conflict over mating rate is particularly pronounced. Each female was allowed to interact with three males for one day (low sexual harassment, LSH) or for eight days (high sexual harassment, HSH) during each breeding cycle throughout their life. Female lifetime fecundity did not differ between the groups, but we found a strong effect on offspring fitness. HSH females produced (1) daughters with smaller bodies and (2) sons with shorter gonopodia, which were less attractive to females and less successful in coercive matings than their LSH counterparts. Although these results may be influenced by the indirect effects of sex ratio differences between treatments, they suggest that sexual harassment and elevated mating rate can have negative cross-generational fitness effects and more profound evolutionary consequences than currently thought.     

The costs of copulating in the dung fly Sepsis cynipsea

Finding, assessing, rejecting, and copulating with a mate isassumed to carry fitness costs, particularly for females, thathave to be traded off against fitness benefits of mating suchas increased fecundity, fertility, longevity, or better qualityoffspring. Female dung flies, Sepsis cynipsea (Diptera: Sepsidae),typically attempt to dislodge mounted males harassing themby vigorous shaking. Shaking duration has been shown to reflect both direct and indirect female choice in this species. Thelatter is an expression of the females' general reluctanceto mate due to presumed costs of mating. We investigated thecosts of copulation in the laboratory. Females were randomlyassigned to one of three treatment groups and allowed to copulateeither not at all, once, or twice. The males' armored genitalia injured females internally during copula. Injuries were visibleas sclerotized scars in the female ovipositor, and their occurrenceincreased with mating frequency. Presumably due to these injuries,mated females showed higher mortality. This effect was statisticallyindependent from additional costs of reproduction related tooviposition, as copulation also increased lifetime egg productionand tended to augment oviposition rate (eggs per day), while fertility (proportion of offspring emerged) was unaffected.We thus found high mortality costs of copulating, indicatingsubstantial sexual conflict, which helps explain female reluctanceto mate in this species.     

Male and female mate choice affects offspring quality in a sex-role-reversed pipefish.

Where both sexes invest substantially in offspring, both females and males should discriminate between potential partners when choosing mates. The degree of choosiness should relate to the costs of choice and to the potential benefits to be gained. We measured offspring quality from experimentally staged matings with preferred and non-preferred partners in a sex-role-reversed pipefish, Syngnathus typhle L. Here, a substantial male investment in offspring results in a lower potential reproductive rate in males than in females, and access to males limits female reproductive success rather than vice versa. Thus, males are choosier than females and females compete more intensely over mates than do males. Broods from preferred matings were superior at escaping predation, when either males or females were allowed to choose a partner. However, only 'choosing' females benefited in terms of faster-growing offspring. Our results have important implications for mate-choice research: here we show that even the more competitive and less choosy sex may contribute significantly to sexual selection through mate choice.     

Plesiomorphic Escape Decisions in Cryptic Horned Lizards (Phrynosoma) Having Highly Derived Antipredatory Defenses

Escape theory predicts that the probability of fleeing and flight initiation distance (predator–prey distance when escape begins) increase as predation risk increases and decrease as escape cost increases. These factors may apply even to highly cryptic species that sometimes must flee. Horned lizards (Phrynosoma) rely on crypsis because of coloration, flattened body form, and lateral fringe scales that reduce detectability. At close range they sometimes squirt blood‐containing noxious substances and defend themselves with cranial spines. These antipredatory traits are highly derived, but little is known about the escape behavior of horned lizards. Of particular interest is whether their escape decisions bear the same relationships to predation risk and opportunity costs of escaping as in typical prey lacking such derived defenses. We investigated the effects of repeated attack and direction of predator turning on P. cornutum and of opportunity cost of fleeing during a social encounter in P. modestum. Flight initiation distance was greater for the second of two successive approaches and probability of fleeing decreased as distance between the turning predator and prey increased, but was greater when the predator turned toward than away from a lizard. Flight initiation distance was shorter during social encounters than when lizards were solitary. For all variables studied, risk assessment by horned lizards conforms to the predictions of escape theory and is similar to that in other prey despite their specialized defenses. Our findings show that these specialized, derived defenses coexist with a taxonomically widespread, plesiomorphic method of making escape decisions. They suggest that escape theory based on costs and benefits, as intended, applies very generally, even to highly cryptic prey that have specialized defense mechanisms.     

Behavioral and physiological female responses to male sex ratio bias in a pond-breeding amphibian

Introduction

The phenomenon of sexual conflict has been well documented, and in populations with biased operational sex ratios the consequences for the rarer sex can be severe. Females are typically a limited resource and males often evolve aggressive mating behaviors, which can improve individual fitness for the male while negatively impacting female condition and fitness. In response, females can adjust their behavior to minimize exposure to aggressive mating tactics or minimize the costs of mating harassment. While male-male competition is common in amphibian mating systems, little is known about the consequences or responses of females. The red-spotted newt (Notophthalmus viridescens) is a common pond-breeding amphibian with a complex, well-studied mating system where males aggressively court females. Breeding populations across much of its range have male-biased sex ratios and we predicted that female newts would have behavioral mechanisms to mitigate mating pressure from males. We conducted four experiments examining the costs and behavioral responses of female N. viridescens exposed to a male-biased environment.

Results

In field enclosures, we found that female newts exposed to a male-biased environment during the five-month breeding season ended with lower body condition compared to those in a female-biased environment. Shorter-term exposure to a male-biased environment for five weeks caused a decrease in circulating total leukocyte and lymphocyte abundance in blood, which suggests females experienced physiological stress. In behavioral experiments, we found that females were more agitated in the presence of male chemical cues and females in a male-biased environment spent more time in refuge than those in a female-biased environment.

Conclusions

Our results indicate that male-biased conditions can incur costs to females of decreased condition and potentially increased risk of infection. However, we found that females can also alter their behavior and microhabitat use under a male-biased sex ratio. Consistent with surveys showing reduced detection probabilities for females, our research suggests that females avoid male encounters using edge and substrate habitat. Our work illustrates the integrated suite of impacts that sexual conflict can have on the structure and ecology of a population.     

Influence of Some Potential Predation Risk Factors and Interaction between Predation Risk and Cost of Fleeing on Escape by the Lizard Sceloporus virgatus

Escape theory predicts that flight initiation distance (predator–prey distance when escape begins) increases as predation risk increases and decreases as cost of fleeing increases. Scant information is available about the effects of some putative predation risk factors and about interaction between simultaneously operating risk and cost of fleeing factors on flight initiation distance and distance fled. By simulating an approaching predator, I studied the effects of body temperature (BT), distance to nearest refuge, and eye contact with a predator, as well as simultaneous effects of predator approach speed and female presence/absence on escape behavior by a small ectothermic vertebrate, the lizard Sceloporus virgatus. Flight initiation distance decreased as BT increased, presumably because running speed increases as BT increases, facilitating escape. Distance to nearest refuge was unrelated to BT or flight initiation distance. Substrate temperature was only marginally related, and air temperature was not related to flight initiation distance. Eye contact did not affect flight initiation during indirect approaches that bypassed lizards by a minimum of 1 m, but an effect of eye contact found in other studies during direct approach might occur. Predator approach speed and presence of a female interactively affected flight initiation distance, which increased as speed increased and decreased when a female was present. In the presence of a female, flight initiation distance was far shorter than when no female was present. The high cost of forgoing a mating opportunity accounts for the interaction because the difference between female presence and absence is greater when risk is greater.     

Sexual conflict and the tragedy of the commons

It is widely understood that the costs and benefits of mating can affect the fecundity and survival of individuals. Sexual conflict may have profound consequences for populations as a result of the negative effects it causes males and females to have on one another's fitness. Here we present a model describing the evolution of sexual conflict, in which males inflict a direct cost on female fitness. We show that these costs can drive the entire population to extinction. To males, females are an essential but finite resource over which they have to compete. Population extinction owing to sexual conflict can therefore be seen as an evolutionary tragedy of the commons. Our model shows that a positive feedback between harassment and the operational sex ratio is responsible for the demise of females and, thus, for population extinction. We further show that the evolution of female resistance to counter harassment can prevent a tragedy of the commons. Our findings not only demonstrate that sexual conflict can drive a population to extinction but also highlight how simple mechanisms, such as harassment costs to males and females and the coevolution between harassment and resistance, can help avert a tragedy of the commons caused by sexual conflict.     

The effects of age at mating on female life-history traits in a seed beetle

Age at first reproduction is an important component of lifehistory across taxa and can ultimately affect fitness. Becausegenetic interests of males and females over reproductive decisionscommonly differ, theory predicts that conflict may arise overthe temporal distribution of matings. To determine the potentialfor such sexual conflict, we studied the direct costs and benefitsassociated with mating at different times for females, usingseed beetles (Acanthoscelides obtectus) as a model system. Virginfemales were resistant to male mating attempts at a very earlyage but subsequently reduced their resistance. Although we foundno difference in life span or mortality rates between femalesmated early in life and those mated later, females that matedearly in life suffered a 12% reduction in lifetime fecundity.Thus, there are direct costs associated with mating early inlife for females. Yet, males mate even with newly hatched females.We suggest that these data indicate a potential for sexual conflictover the timing of first mating and that female resistance tomating, at least in part, may represent a female strategy aimedat delaying mating to a later time in life.     

No Effect of Male Courtship Intensity on Female Remating in the Butterfly <Emphasis Type="Italic">Pieris napi</Emphasis>

In Pieris napi, female fitness increases with number of matings, but wild females mate at an unexpectedly low rate. From a sexual conflict perspective this could be because males manipulate female remating, or alternatively, because wild females experience costs associated with remating which are not applicable under laboratory conditions. To get an indication which sex controls remating and/or the different sexes’ relative costs and benefits of remating, we here test whether female mating frequency is affected by male courtship intensity. We found no effect on female mating frequency or lifespan. This indicates that (i) females control remating and their optimal mating frequency is lower compared to males, or (ii) males can manipulate female remating. We argue that both these alternatives may apply simultaneously to P. napiand that they are inseparable.     

Sexual selection on spontaneous mutations strengthens the between‐sex genetic correlation for fitness

A proposed benefit to sexual selection is that it promotes purging of deleterious mutations from populations. For this benefit to be realized, sexual selection, which is usually stronger on males, must purge mutations deleterious to both sexes. Here, we experimentally test the hypothesis that sexual selection on males purges deleterious mutations that affect both male and female fitness. We measured male and female fitness in two panels of spontaneous mutation‐accumulation lines of the fly, Drosophila serrata, each established from a common ancestor. One panel of mutation accumulation lines limited both natural and sexual selection (LS lines), whereas the other panel limited natural selection, but allowed sexual selection to operate (SS lines). Although mutation accumulation caused a significant reduction in male and female fitness in both the LS and SS lines, sexual selection had no detectable effect on the extent of the fitness reduction. Similarly, despite evidence of mutational variance for fitness in males and females of both treatments, sexual selection had no significant impact on the amount of mutational genetic variance for fitness. However, sexual selection did reshape the between‐sex correlation for fitness: significantly strengthening it in the SS lines. After 25 generations, the between‐sex correlation for fitness was positive but considerably less than one in the LS lines, suggesting that, although most mutations had sexually concordant fitness effects, sex‐limited, and/or sex‐biased mutations contributed substantially to the mutational variance. In the SS lines this correlation was strong and could not be distinguished from unity. Individual‐based simulations that mimick the experimental setup reveal two conditions that may drive our results: (1) a modest‐to‐large fraction of mutations have sex‐limited (or highly sex‐biased) fitness effects, and (2) the average fitness effect of sex‐limited mutations is larger than the average fitness effect of mutations that affect both sexes similarly.     

Sex and Weapons: Contrasting Sexual Dimorphisms in Weaponry and Aggression in Snapping Shrimp

Sexual dimorphisms in weaponry and aggression are common in species in which one sex (usually males) competes for access to mates or resources necessary for reproduction – sexually dimorphic weaponry and aggression, in other words, are frequently the result of intrasexual selection. In snapping shrimp, the major chela (snapping claw) can be a deadly weapon, and males of many species have larger chelae than females, a pattern readily interpreted as resulting from intrasexual selection. Thus, males might be expected to show more sex‐specific aggression than females, and be more aggressive overall. We tested these predictions in two species of snapping shrimp in a territorial defense context. Neither of these predictions was supported: in both species, females, but not males, engaged in sex‐specific aggression and females were more aggressive than males overall. These contrasting sexual dimorphisms – larger weaponry in males but higher aggression in females – highlight the importance of considering the function of weaponry and aggression in contexts other than direct competitions over mates. In addition, species differences in the degree of sexual dimorphism in chela size were due to differences in female, not male, chela size, and the species with greater sexual dimorphism in weaponry was significantly less aggressive overall; also, while paired and solitary males did not differ in residual chela size, for the species with greater sexual dimorphism, females carrying embryos had smaller residual chela sizes. These results suggest that understanding the sexual dimorphisms in weaponry and aggression in snapping shrimp requires understanding the relative costs and benefits of both in females as well as males.     

Sex allocation and nestling survival in a dimorphic raptor: does size matter?

Fisher's theory predicts equal sex ratios at the end of parentalcare if the costs and benefits associated with raising eachsex of offspring are equal. In raptors, which display variousdegrees of reversed sexual size dimorphism (RSD; females thelarger sex), sex ratios biased in favor of smaller males areonly infrequently reported. This suggests that offspring ofeach sex may confer different fitness advantages to parents.We examined the relative returns associated with raising eachsex of offspring of the brown falcon Falco berigora, a medium-sizedfalcon exhibiting RSD (males approximately 75% of female bodymass) and subsequent sex ratios. Female nestlings hatched eitherfirst or second did not receive more food nor did they hatchfrom larger eggs or remain dependent on parents for longer periodsthan male offspring from these hatch orders. Together with previousstudies this result indicates that even in markedly dimorphicspecies, the required investment to raise the larger sex islikely to be less than that predicted by body size differencesalone. Moreover, among last-hatched nestlings, both sexes faceda reduced food allocation and suffered a slower growth rateand thus final body size, with a concurrent increased probabilityof mortality. For last-hatched females the reduction in foodallocation was more marked, with complete mortality of all last-hatchedfemale nestlings monitored in this study. Once independent,males of any size but only larger females are likely to be recruitedinto the breeding population. The sex-biased food allocationamong last-hatched offspring favoring males thus reflects therelative returns to parents in raising a small member of eachsex.