Female nursery web spiders (Pisaurina mira) benefit from consuming their mate

The often coincidental involvement of cooperation and conflict in animal reproduction is epitomized by sexual cannibalism, which can lead to obvious male costs while simultaneously providing direct benefits to developing offspring. Male nursery web spiders (Pisaurina mira) avoid postcopulatory sexual cannibalism by wrapping females with silk. Here, we test the hypothesis that this silk wrapping results in a loss of consumption cost for females. In specific, we hypothesize that females lose out on potential fitness benefits associated with cannibalizing their mating partners. To test this, we paired females with males that were experimentally manipulated to prevent the silk wrapping of females, thereby increasing the likelihood of sexual cannibalism. Females either did not kill their mate, and thus consumed nothing, or did kill their mate. If females killed their mating partner, we allowed them to consume the male, consume nothing, or consume a cricket. We found no effect of female or male body sizes on the likelihood of females killing their mate. While our treatments did not affect the number of offspring females produced, females that consumed either a male or a cricket produced egg sacs faster than females that consumed nothing, suggesting a benefit of increased postcopulatory food consumption. Further, only females that ate a male had heavier and longer lived offspring, suggesting a benefit of sexual cannibalism specifically. Our results support the hypothesis that females can receive fitness benefits associated with sexual cannibalism.     

Eat or Not to Eat: Reversed Sexual Cannibalism as a Male Foraging Strategy in the Spider Micaria sociabilis (Araneae: Gnaphosidae)

Reversed sexual cannibalism represents an unusual situation in which a male kills and consumes a female. We examined this rare phenomenon in the spider Micaria sociabilis, whose males were observed to regularly cannibalise old females. In this study, we investigated male motivation for such behaviour in the light of ecological conditions such as mate availability and prey availability. We found that male cannibalism is not affected by short‐term starvation but rather by male feeding history during the ontogenetic development in combination with prey availability during the adult stage. Males from the summer generation reached bigger sizes than males from the spring generation and females from both generations. They developed in the period with exceptionally high prey occurrence, but when they reached the adult stage, the prey availability decreased. In this period, we observed the highest frequency of cannibalism, however, only when the sex ratio was female biased. Reversed sexual cannibalism in M. sociabilis seems to represent an advantageous male foraging strategy, which is affected by prey availability and male feeding history, tuned by sex ratio and directed towards females of inferior quality.     

Sexual cannibalism in Nephila plumipes as a consequence of female life history strategies

The evolution of sexual cannibalism has been modelled as both an adaptive and nonadaptive female strategy. Recent evidence from several species suggests a connection between female foraging and sexual cannibalism, but the precise benefits for females have remained obscure. Here, we investigate the difference between cannibalistic and noncannibalistic female Nephila plumipes by removing the potential nutritional benefit of cannibalism. Courting and mating males that were killed by a female were immediately removed so that the female could not consume them. Nevertheless, cannibalistic females gained more mass from maturation to oviposition and produced larger first clutches than noncannibalistic females, although cannibalistic females matured at a smaller size and mass than noncannibalistic females. In juvenile instars, mass gain was generally smaller in females that moulted in a good condition but intermoult intervals were shorter. However, the time from maturity to oviposition was not shorter in females that matured in a good condition. Male behaviour did not differ according to the risk of cannibalism. We suggest that sexual cannibalism in N. plumipes is a side‐effect of an increased foraging vigour of females that matured at a smaller size and body mass. Selection pressure on males to avoid cannibalism may be weak because of limited mating opportunities.     

Does Female Personality Determine Mate Choice Through Sexual Cannibalism?

Animal personalities (e.g. consistent across‐context behavioural differences between individuals) can lead to differences in mate choice. However, evidence for this link remains limited. Pre‐mating sexual cannibalism can be a behavioural syndrome (i.e. a suboptimal personality) in which adaptive female aggression towards heterospecific prey spills over on non‐adaptive aggression towards courting males, independently of the female mating or feeding status (i.e. the ‘aggressive spillover hypothesis’, ASH). On the other hand, sexual cannibalism can also be a form of mate choice by which females selectively kill or mate with males depending on the male phenotype. We introduce the hypothesis that the most aggressive females in the population will not only attack males more frequently, but will be less likely to impose sexual selection on males through sexual cannibalism. Assuming that in a field common garden experiment in which females were fed ad libitum the rate of weight gain by a female may reflect her voracity or aggressiveness, we show that in the cannibalistic burrowing wolf spider Lycosa hispanica (formerly L. tarantula), voracity towards heterospecific prey predicts a female's tendency towards sexual cannibalism. Unmated females with higher weight gains were more cannibalistic and attacked males regardless of the male phenotype. On the other hand, females that were less voracious tended to be less cannibalistic, and when they did kill a male, they were selective, killing males in poorer condition and mating with those in better condition. Our results demonstrate that females with different phenotypes (growth rates) differently imposed selection on male condition, tentatively supporting the hypothesis that female aggression levels can spill over on sexual selection through sexual cannibalism.     

Seasonal aspects of sexual cannibalism in the praying mantis (<Emphasis Type="Italic">Mantis religiosa</Emphasis>)

According to the adaptive foraging hypothesis of sexual cannibalism, females face a trade-off between mating and consuming a courting male. Because male and prey availability can change seasonally, sexual cannibalism may change with season. However, we are not aware of any work examining how sexual cannibalism in insects relates to the time of season. Here, we examined the seasonal pattern of sexual cannibalism and reproductive behaviour in the sexually cannibalistic praying mantis (Mantis religiosa). We repeatedly collected the last instars of praying mantises from the field and brought them up under natural weather and photoperiod, but standardised feeding and socioecological conditions. After the females reached sexual maturity, we allowed all of the females to mate during two mating trials. In comparison to female praying mantises maturing later in the season, early-maturing females were larger but of poorer body condition on the day of a mating trial (20 days after the adult moult). During the first round of mating trials, early-maturing virgin females cannibalised males more frequently than their late-maturing counterparts. In contrast, late-maturing females that mated in the first round of mating trials were more likely than early-maturing, nonvirgin females to be cannibalistic in the second round of mating trials. The latency time until copulation was correlated with a risk of sexual cannibalism and was longer in early-maturing females. Our study suggests that the date of the last (adult) moult plays an important role in the occurrence of sexual cannibalism.     

Female hunger can explain variation in cannibalistic behavior despite male sacrifice in redback spiders

Male Australian redback spiders (Latrodtctus hasselti Thorell:Theriidae) place their abdomens directly over their mate's mouthpartsduring copulation, increasing the likelihood of sexual cannibalism.Male sacrifice may be adaptive because cannibalized males increasetheir paternity relative to those that are not eaten. Despitemale sacrifice behavior, however, up to 50% of laboratory matingsmay end without sexual cannibalism. Here, I report a similarpattern in the field, where males were not cannibalized in 35%of observed matings (6/17). I examined variation in female cannibalisticbehavior by evaluating the following three hypotheses for theoccurrence of cannibalism from the female perspective: (1) themistaken identity hypothesis proposes that females sometimescannibalize males because they mistake them for prey, (2) themate rejection hypothesis predicts that females cannibalizemales who are unacceptable as mates, and (3) the feeding opportunismhypothesis predicts that hungry females are more likely to becannibalistic Field observations refuted die first two hypotheses:females recognized males as potential mates (i.e., nonprey),and cannibalized and noncannibalized males were not phenotypicallydifferent. The feeding opportunism hypothesis was supported.In staged field matings, cannibalistic females were hungrierthan their noncannibaUstic counter-parts. Moreover, a logisticregression analysis indicated that hunger was a significantpredictor of cannibalism. Because redback males are below thetypical prey size that females accept, well-fed females areless likely to consume their mates, despite the vulnerable matingposture. These results indicate that, although males facilitatesexual cannibalism, their fate may depend on the female's physicalcondition.     

Kin‐bias,breeding site selection and female fitness in a cannibalistic Neotropical frog

Resource availability influences sexual selection within populations and determines whether behaviours such as territoriality or resource sharing are adaptive. In Thoropa taophora, a frog endemic to the Atlantic Coastal Rainforest of Brazil, males compete for and defend limited breeding sites while females often share breeding sites with other females; however, sharing breeding sites may involve costs due to cannibalism by conspecific tadpoles. We studied a breeding population of T. taophora to determine (i) whether this species exhibits polygynous mating involving female choice for territorial males and limited breeding resources; (ii) whether limited breeding resources create the potential for male–male cooperation in defence of neighbouring territories; and (iii) whether females sharing breeding sites exhibit kin‐biased breeding site choice, possibly driven by fitness losses due to cannibalism among offspring of females sharing sites. We used microsatellites to reconstruct parentage and quantify relatedness at eight breeding sites in our focal population, where these sites are scarce, and in a second population, where sites are abundant. We found that at localities where the appropriate sites for reproduction are spatially limited, the mating system for this species is polygynous, with typically two females sharing a breeding site with a male. We also found that females exhibit negative kin‐bias in their choice of breeding sites, potentially to maximize their inclusive fitness by avoiding tadpole cannibalism of highly related kin. Our results indicate that male territorial defence and female site sharing are likely important components of this mating system, and we propose that kinship‐dependent avoidance in mating strategies may be more general than previously realized.     

A test of sexual cannibalism models, using a sit-and-wait predator

Sexual cannibalism, the capture and consumption of one sex by the other resulting from sexual interactions, provides an intriguing set of conditional fitness payoffs. Usually these payoffs will only benefit the cannibal, but under certain circumstances advantages might accrue to the cannibalized individual as well. Here I test three models of sexual cannibalism with data from a sit-and-wait predator, the crab spider Misumena vatia , whose lifestyle differs from the species for which these models were generated. The models include both precopulatory mechanisms (no gametes exchanged) and postcopulatory ones (gametes exchanged) perceived to generate adult fitness payoffs and situations in which cannibalism is not adaptive in the adult stage (aggressive spillover). In M. vatia , females sometimes cannibalize males before mating can occur. Precopulatory cannibalism is unlikely to reward female M. vatia significantly because of the diminutive size of the males, but it could be part of a broader syndrome that provides the females with large food payoffs earlier in the life cycle. Although the frequency of cannibalism is low (3.8–7.6%), this level suffices to be an important selective factor, a point accentuated by the extremely cautious behaviour of the males toward the females. I then compare the traits associated with these models to the species for which they were designed and certain others for which adequate data also exist. None of the sets of responses to these traits closely resembles each other in the different species. These results suggest that sexual cannibalism has developed in response to a heterogeneous collection of lifestyles and phylogenetic constraints.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 427–437.     

Sexual cannibalism in the fishing spider and a model for the evolution of sexual cannibalism based on genetic constraints

Several hypotheses have been proposed for the evolution of sexual cannibalism by females. Newman and Elgar (1991) suggested that sexual cannibalism prior to mating by virgin female spiders may have evolved as a result of female foraging considerations. According to this model, an adult female's decision to mate or cannibalize a courting male should be based on an assessment of the male's value as a meal versus his value as a mate. The current study provides an empirical test of the assumptions and predictions of this model in the sexually cannibalistic fishing spider. Adult females were subjected to different food treatments, and exposed to adult males in the laboratory. However, only one of the assumptions of the model and none of its five predictions were upheld. We failed to find any effects of female foraging, female mating status, female size, male size or time of the season on females' behaviour towards courting males. Females behaved stereotypically, and many females were left unmated despite numerous mating opportunities. We also demonstrate costs of sexual cannibalism in a natural population. We propose that the act of sexual cannibalism in the fishing spider is non-adaptive, and develop a model for the evolution of premating sexual cannibalism in spiders based on genetic constraints. According to this hypothesis, sexual cannibalism by adult females may have evolved as an indirect result of selection for high and non-discriminate aggression during previous ontogenetic stages. Genetic covariance between different components of aggressive behaviour may constrain the degree to which (1) juvenile and adult aggression and/or (2) aggression towards conspecifics and heterospecifics can vary independently. We briefly review the support for our model, and suggest several critical tests that may be used to assess the assumptions and predictions of the model.     

The Role of Body Size in Mating Interactions of the Sexually Cannibalistic Fishing Spider Dolomedes triton

Some arachnids display extreme sexual size dimorphism (SSD) with adult females being several times larger than adult males. One explanation for SSD in species that exhibit pre‐copulatory sexual cannibalism (female attack, kill and consumption of the male prior to mating) is that smaller males may be less likely victims of predatory attacks by females. However, in some sexually cannibalistic species SSD is relatively moderate (i.e. males are similar in size to females) suggesting benefits of large male body size. Here, I report the results of an experiment designed to explore the ramifications of body size in mating interactions of the sexually cannibalistic, North American fishing spider (Dolomedes triton). Results suggest that male size does not influence courtship behavior, the likelihood of being attacked, or the male's ability to secure a mounting. However, large males were superior at gaining copulations once mounted. Sexual cannibalism may also be predicated on female size. Female condition (mass/cephalothorax area) did not explain any of these behaviors from the copulatory sequence, however, females with a smaller cephalothorax area were more likely to attack courting males. Finally, analysis of the ratio of female size to male size showed that when SSD is weak males are more likely to escape attacks and mate successfully. Results are discussed in light of several hypotheses for sexual cannibalism, and the benefits of large male body size illustrated here are put forth as potential explanations for the relatively moderate extent of SSD found in this sexually cannibalistic species.     

Avoidance of post-coital cannibalism in the brachyurid paddle crab Ovalipes catharus

Sexual cannibalism was examined experimentally in the New Zealand paddle crab Ovalipes catharus, where it is the female that risks cannibalism during and after sexual activity. In this species copulation only occurs when the female is soft-shelled after moulting and she is most vulnerable to cannibalism. Male paddle crabs protect and copulate with the females until the females are no longer sexually receptive. Males appear able to identify a female with whom they have recently copulated. After a brief separation, significantly fewer females were cannibalized in cases where a female was returned to her original sexual partner than in those that had exchanged partners. The asymmetry, where females gain some protection during moulting but males receive no such advantage, may, at least partially, explain the skewed sex ratios which are sometimes found in samples of O. catharus from nature.     

Cohabitation of juvenile females with mature males promotes sexual cannibalism in fishing spiders

Precopulatory sexual cannibalism, or female predation of a potentialmate before mating, illustrates an extreme intersexual conflict.Unlike sexual cannibalism occurring after mating, precopulatorysexual cannibalism cannot be construed as a male strategy. Thus,research on the adaptive significance of this phenomenon hasfocused on female benefits. In the present study, I test theidea that precopulatory sexual cannibalism represents an adaptivefemale trade-off between the material costs and benefits ofmating with a male (forgoing food, securing sperm) and preyingon a male (forgoing sperm, securing food). I pay particularattention to the rarely tested prediction that precopulatorysexual cannibalism by virgin females should increase as eachfemale's expectation of future mating opportunities increases.I use the phenomenon of cohabitation between adult males andjuvenile females nearing sexual maturity as a means to manipulatefemale expectation of future mate availability. Results indicatethat feeding on a male has significant positive effects on severalmeasures of female fecundity. However, the likelihood of precopulatoryattacks was not explained by a female's recent feeding history.Finally, as predicted, juvenile female cohabitation with maturemales (expectation of future mating opportunities) heightensthe prevalence of precopulatory attacks by virgin females, suggestingthat juvenile experience can alter a female's propensity forsexual cannibalism. This is the first study to suggest thatjuvenile experience can alter a female's propensity for sexualcannibalism. This finding emphasizes the point that studiesof sexual selection and mating systems need to consider theeffects of juvenile experience on adult behavior.     

Does a single meal affect female reproductive output in the sexually cannibalistic praying mantid Iris oratoria?

1. One explanation of the evolution of sexual cannibalism, the female’s consumption of a male during or following courtship or copulation, is that this behaviour increases the female’s fitness. This study tests the assumption that a single meal increases female reproductive output significantly in the sexually cannibalistic praying mantid Iris oratoria L. 2. In 38 mating trials, seven of the females cannibalised the males. In order to augment the number of females that fed, an additional nine females were each fed one cricket nymph at the end of the mating trial. 3. Three measures of female reproductive output – the occurrence of oviposition, the mass of the first ootheca, and the number of eggs in the first ootheca – increased significantly with female feeding condition, which was a reflection of food consumed before the mating trial. Females that copulated later in the season tended to lay lighter oothecae. 4. The females’ consumption of a meal during the mating trial, either a conspecific male or a cricket, did not influence any measure of reproductive output significantly, although possible effects upon subsequent oothecae cannot be ruled out. 5. If, as the present study suggests, a single meal provides a negligible or delayed benefit to female reproductive output, the evolution of sexual cannibalism might lie in alternative explanations, which include possible fitness benefits to cannibalistic females in the nymphal stage or possible paternity benefits to the cannibalised males.     

Filial Cannibalism in the Paternal Mouthbrooding Cardinalfish Apogon lineatus: Egg Production by the Female as the Nutrition Source for the Mouthbrooding Male

We investigated about filial cannibalism by Apogon lineatus based on 25 mouthbrooding males sampled in Tokyo Bay, Japan. Eggs were found in the stomachs of all mouthbrooding males, indicating that Apogon lineatus performs partial brood cannibalism. The number of eggs in the stomachs ranged from 1015 to 9384, corresponding to 30% on average of the entire brood. This study revealed that mouthbrooding males could gain considerable energy by partial brood cannibalism during brood periods, that is, females would produce surplus eggs as the nutrition sources for starving males.     

Male adaptations to minimize sexual cannibalism during reproduction in the funnel‐web spider Hololena curta

Males of many spider species risk being attacked and cannibalized while searching for, courting, and mating with conspecific females. However, there are exceptions. We show that the funnel‐web spider, Hololena curta, has 3 adaptations that minimize risk to males during courtship and mating, and enhance reproductive success. First, males detected chemical or tactile signals associated with webs of virgin females, and differentiated them from webs of mated females, enabling males to increase encounter rates with virgin females and avoid aggressive mated females. Second, males produced stereotyped vibrational signals during courting which induced female quiescence and suppressed female aggression. Third, when touched by males, sexually receptive females entered a cataleptic state, allowing males to safely approach and copulate. Because males can mate multiple times and the sex ratio in natural populations of H. curta is female biased, overall reproductive output is likely increased by males of this species avoiding sexual cannibalism.     

Male mating behaviour reduces the risk of sexual cannibalism in an Australian praying mantid

In cases where sexual cannibalism represents a sexual conflict, we should expect to find male traits that reduce the risk of cannibalism. In fact, a wide variety of such traits have been proposed, including elaborate courtship displays, cautious approach behaviours, and opportunistic mating whilst a female is feeding. However, there is very little direct evidence that these behaviours actually reduce the risk of sexual cannibalism for males, and the evidence that does exist comes mainly from spider studies. In this study of Pseudomantis albofimbriata praying mantids we found evidence of various male behaviours that reduced the risk of sexual cannibalism. Most males attempted to avoid detection entirely and others moved in a slow and stealthy manner. We also observed five cases of opportunistic mating, but saw no evidence of any male courtship routine. It seems that mounting from the rear of a female or slowly approaching her from the front, allows many males to go unnoticed and, therefore, substantially reduces the risk of sexual cannibalism in this mating system. Interestingly, we could not identify any female traits that males use to assess the risk of sexual cannibalism. It may be that P. albofimbriata males use alternative means to assess these risks. The presence of these behavioural strategies by males suggests a net cost of being consumed, and that sexual cannibalism is likely to be a female strategy in this system.     

A Test for Male Parental Care in a Fundulid, the Bluefin Killifish, Lucania goodei

Under field conditions, breeding male bluefin killifish, Lucania goodei, have been observed aggressively defending territories from other breeding males, non-breeding females, and minnows (mainly Notropis harperi). We performed an aquarium experiment to test whether male aggression serves to protect newly deposited eggs from predation. We allowed a male and a female to spawn in a yarn mop, removed the female, and exposed the eggs to one of four treatments (spawning male present, two minnows present, spawning mal+two minnows present, no adult fish present). Mops were censused daily for seven days. Egg predation rates were highest in the male+minnows and male only treatments. Egg predation rates in the male+minnows treatment did not differ from the predicted predation rate (sum of male only and minnows only treatments). Hence, there is no evidence for male parental care in L. goodei. In addition, we compared the egg predation rates (filial cannibalism) between males of 3 different color morphs and found no evidence for differential egg cannibalism.     

Female fecundity and offspring survival are not increased through sexual cannibalism in the spider Larinioides sclopetarius

Many hypotheses explaining the evolution and maintenance of sexual cannibalism incorporate the nutritional aspect of the consumption of males. Most studies have focused on a fecundity advantage through consumption of a male; however, recent studies have raised the intriguing possibility that consumption of a male may also affect offspring quality. In particular, recent studies suggest prolonged survival for offspring from sexually cannibalistic females. Here, we measured the protein and lipid content of males compared to insect prey (crickets), quantified female nutrient intake of both prey types and finally assessed how sexual cannibalism affects female fecundity and spiderling quality in the orb‐web spider Larinioides sclopetarius. We found no evidence that sexual cannibalism increased fecundity when compared to a female control group fed a cricket. Contrary to previous studies, spiderlings from females fed a male showed reduced survival under food deprivation compared to spiderlings from the control group. Offspring from females fed a male also tended to begin web construction sooner. The low lipid content of males compared to crickets may have reduced offspring survival duration. Whether additional proteins obtained through consumption of a male translate to enhanced silk production in offspring requires further investigation.     

Female remating propensity contingent on sexual cannibalism in sagebrush crickets, Cyphoderris strepitans: a mechanism of cryptic female choice

Male sagebrush crickets (Cyphoderris strepitans) permit femalesto engage in an unusual form of sexual cannibalism during copulation:females feed on males' fleshy hind wings and ingest hemolymphoozing from the wounds they inflict. These wounds are not fatal,and normally only a portion of the hind wings are eaten at anyone mating, so that mated males are not precluded from matingagain. As a result, nonvirgin males have fewer material resources tooffer females than do virgin males, such that females shouldbe selected to preferentially mate with high-investment virginmales. We tested the hypothesis that female mating preferencesfavor males capable of supplying females with the highest materialinvestment. Our results indicate that both female diet and opportunitiesfor sexual cannibalism influence female mating behavior. Femalesmaintained on a low-nutrient diet mounted males significantlysooner than females maintained on a high-nutrient diet, indicatingthat a female's overall nutrient intake may determine her propensityto mate. In addition, females were significantly more reluctantto mount and mate with males whose hind wings had been surgicallyremoved and thus were incapable of providing females with awing meal. Finally, females initially mated to dewinged malesremated with winged males significantly sooner than femalesallowed to feed freely during their initial mating, resultingin cryptic female choice of investing males.     

Sexual cannibalism, competition, and size dimorphism in the orb-weaving spider Nephila plumipes Latreille (Araneae: Araneoidea)

The degree and direction of sexual dimorphism varies widely,but in several taxa of orb-weaving spiders, including Nephila,males may be less than one-tenth the size of females. This differenceis commonly attributed to selection through precopulation sexualcannibalism: females may either fail to detect very small males,or ignore them as potential prey items. However, there is oftenthe potential for male-male competition in these species becauseseveral males can be found on the web of a single female. Weinvestigated experimentally the effects of sexual cannibalismand male-male competition on male body size and hence sexualdimorphism in the Australian golden orb-weaver (Nephila plumipes).Small males were less likely to be detected and cannibalizedthan larger males. However, larger males excluded small malesfrom the central hub of the web, where mating takes place. Theconflicting effects of sexual cannibalism and male-male competitionmay be responsible for the relatively large variation in malebody size in this species.