Kin recognition and cannibalism in a subsocial spider

Evolution of cooperation and group living in spiders from subsocial family groups may be constrained by their cannibalistic nature. A tendency to avoid cannibalizing kin may facilitate tolerance among spiders and implies the ability to identify relatives. We investigated whether the subsocial spider Stegodyphus lineatus discriminates kin by recording cannibalism among juveniles in experiments during which amount of food and size difference among spiders in groups were varied. We hypothesized that family groups should be less cannibalistic than groups of mixed‐parental origin. Further, we tested whether food‐stress would influence cannibalism rates differently in kin and nonkin groups and the effect of relatedness on cannibalism within groups of spiders of variable size compared with those of homogenous size. In groups of six spiders, more spiders were cannibalized in nonsib groups than in sib groups under low food conditions. A tendency for nonkin biased cannibalism in starved spider pairs supported that kin recognition in S. lineatus is expressed when food is limited. Size variance of individuals within well‐fed groups of siblings and unrelated spiders had no influence on cannibalism rates. Apparently, both hunger and high density are important promoters of cannibalism. In addition to inclusive fitness benefits, we suggest that an ability to avoid cannibalizing kin will favour the evolution of cooperation and group living in phylogenetically pre‐adapted solitary species.     

Sexual size dimorphism predicts the frequency of sexual cannibalism within and among species of spiders

Sexual cannibalism varies widely among spiders, but no general evolutionary hypothesis has emerged to explain its distribution across taxa. Sexual size dimorphism (SSD) also varies widely among spiders and could affect the vulnerability of males to cannibalistic attacks by females. We tested for a relationship between SSD and sexual cannibalism within and among species of spiders, using a broad taxonomic data set. For most species, cannibalism was more likely when males were much smaller than females. In addition, using phylogenetically controlled and uncontrolled analyses, there was a strong positive relationship between average SSD of a species and the frequency of sexual cannibalism. This is the first evidence that the degree of size difference between males and females is related to the phylogenetic distribution of sexual cannibalism among a broad range of spiders.     

Sibling Cannibalism of Young Red Drum, Sciaenops ocellatus, in Relation to Size Disparity and Metabolic Rates

Cannibalism is a leading cause of young mortality in the red drum, Sciaenops ocellatus, culture, and size disparity is its primary causative factor. Cannibalism did not occur in young fish, graded by a difference of 0.5cm from 2.0 to 4.5cm in total length. There was a shift in the size distribution from unimodal (normal) when there was no cannibalism, to bimodal or trimodal when cannibalism occurred. The results suggested that the wider was the size disparity, the greater was the sibling cannibalism. Size disparity increased with growth and in turn with sibling cannibalism, which became insignificant when the young grew to a size of about 6cm. We present threshold levels of size disparity among siblings to prevent mutual cannibalism. We hypothesize that through sibling cannibalism a hierarchy may evolve in a red drum cohort, at which a stable population will form.     

Sex-dependent effects of sibling cannibalism on life history traits of the ladybird beetle Harmonia axyridis (Coleoptera: Coccinellidae)

To investigate the sex-dependent effects of sibling cannibalism on variations in life history traits, I analysed body size, weight and instar interval in relation to the occurrence of sibling cannibalism in the ladybird beetle Harmonia axyridis. Sibling cannibalism at the time of hatching significantly affected the body size and weight of adults. There was a 2.32% and 1.05% increase in the body size of males and females, respectively, and a 3.55% increase and a 2.30% decrease in their respective body weights. Sibling cannibalism also significantly shortened the total and larval instar intervals, by 4.24% in males and by 1.22% in females, mainly due to shortening of the first instar. These results suggest that the effects of sibling cannibalism on life history traits are sex-differentiated and are greater in males than in females. A simulation of aphid density indicated that shortening the instar interval affected larval survival; the aphid density when the larvae completed development was 39.71% and 10.52% larger for cannibalistic males and females, respectively, than for non-cannibals. These results suggest that sibling cannibalism promotes more rapid development and larger adult size, although the effect was more pronounced in males than in females. Faster development may be adaptive for resource tracking, and the large adult size may increase fecundity in females and mating success in males through female mate choice, both resulting in an increase in the fitness of cannibals.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 349–360.     

Refuge from cannibalism in complex-structured habitats: implications for the accumulation of invertebrate predators

Abstract.  1. Like many invertebrate predators, the wolf spider Pardosa littoralis Banks (Araneae: Lycosidae) accumulates in complex-structured habitats replete with leaf litter (thatch). Here we test the hypothesis that P. littoralis accumulates in complex habitats to gain refuge from cannibalism.
2. A laboratory experiment examined the effects of habitat complexity (thatch present or absent) and size-class pairing of conspecific spiders (large vs. small, small vs. small, and large vs. large) on the incidence of cannibalism. Spider survival was significantly higher (22%) in complex-structured habitats with thatch than in simple-structured habitats lacking thatch. Furthermore, cannibalism occurred more frequently in P. littoralis when the size of conspecifics was asymmetric (large vs. small spiders) than when spiders were of equal size. There was no interactive effect of habitat complexity and size-class pairing on spider survival.
3. A field experiment examined the effects of habitat complexity, conspecific density, and access to alternative prey on the prevalence of cannibalism in P. littoralis . Access to alternative prey significantly increased the number of spiders recovered from field enclosures, as did the presence of leaf litter thatch. That fewer spiders were recovered when thatch and alternative prey were absent suggests that cannibalism was most prevalent under these conditions.
4. Overall, results suggest that habitat complexity reduces agonistic interactions and cannibalism among wolf spiders, providing encouragement to pest managers that the structure of agricultural habitats can be managed to maximise densities of generalist predators for enhanced pest suppression.     

Nutritional value of cannibalism and the role of starvation and nutrient imbalance for cannibalistic tendencies in a generalist predator

1. Cannibalism is considered an adaptive foraging strategy for animals of various trophic positions, including carnivores. However, previous studies on wolf spiders have questioned the high nutritional value of cannibalism. We therefore analysed two different aspects of nutritional quality of conspecifics in the wolf spider Pardosaprativaga: their value for survival, growth and development; and the growth efficiency of feeding on conspecifics. We also measured the propensity for cannibalistic attacks and the consumption rate of conspecifics in an experiment where hunger level and nutrient balance were manipulated. In all experiments, cannibalism was compared with predation on fruit flies as control prey. 2. The growth experiment gave ambiguous results regarding the nutritional quality of conspecifics. Spiders on pure cannibalistic diets split into two distinct groups, one performing much better and the other much worse than spiders on fruit fly diets. We discuss the possibility that the population is dimorphic in its cannibalistic propensity, with the latter group of individuals showing a high level of inhibition against cannibalistic attacks in spite of a high nutritional value of cannibalism. 3. The food utilization experiment confirmed the high nutritional quality of conspecifics, as cannibalistic spiders had the same growth rate as spiders fed insect prey in spite of a much lower consumption rate. 4. Inhibition against cannibalistic attacks was demonstrated in medium-sized juveniles: only half of the spiders attacked a prescribed victim of 50% the size of their opponents, and the latency for those that did attack was more than half an hour, compared with a few minutes for spiders fed fruit flies. 5. Nutrient-imbalanced spiders utilized an alternative insect diet less efficiently than balanced spiders, whereas no difference was present in efficiency of utilizing conspecifics. This result indicates that spiders can remedy at least part of a nutrient imbalance through cannibalism. 6. As spiders can escape nutritional imbalance as well as restore energy reserves through cannibalism, we predicted both nutrient imbalance and hunger to stimulate cannibalism. This prediction was confirmed only with respect to hunger. Nutrient-imbalanced spiders had reduced cannibalistic consumption, perhaps due to lowered predatory aggressiveness as a result of bad condition.     

Maternal hatching synchronization in a subsocial burrower bug mitigates the risk of future sibling cannibalism

Sibling cannibalism—the killing and consumption of conspecifics within broods—carries a high risk of direct and inclusive fitness loss for parents and offspring. We reported previously that a unique vibrational behavior shown by the mother of the subsocial burrower bug, Adomerus rotundus (Heteroptera: Cydnidae), induced synchronous hatching. Maternal regulation may be one of the most effective mechanisms for preventing or limiting sibling cannibalism. Here, we tested the hypothesis that synchronous hatching induced by maternal vibration in A. rotundus prevents sibling cannibalism. Mothers and their mature egg masses were allocated to three groups: synchronous hatching by maternal vibration (SHmv), synchronous hatching by artificial vibration (SHav), and asynchronous hatching (AH). We then investigated the influence of each hatching strategy on the occurrence of sibling cannibalism of eggs and early‐instar nymphs in the laboratory. No difference in the proportion of eggs cannibalized was observed among the three groups. However, the proportion of nymphs cannibalized was higher in the AH group than in the SHmv group. The difference in the number of days to first molting within clutch was significantly higher in the AH group than in the SHmv group. Junior nymphs were sometimes eaten by senior nymphs. However, immediately after molting, senior nymphs were at a high risk of being eaten by junior nymphs. Our results indicate that synchronous hatching of A. rotundus is necessary to mitigate the risk of sibling cannibalism.     

Interdemic variation of cannibalism in a wolf spider (Pardosa monticola) inhabiting different habitat types

Abstract.  1. Cannibalism was investigated in the wolf spider Pardosa monticola (Clerck) using spiders collected from four populations with varying densities, inhabiting two different coastal dune habitat types. Sampled individuals were paired randomly and tested immediately for their cannibalism propensity.
2. The occurrence of cannibalism was found to be influenced by the size (cephalothorax width) of both the smaller and the larger spider of a pair. Larger size differences enhanced cannibalism.
3. Cannibalism rates were not significantly different in spiders from high-density compared with low-density populations. Cannibalism rates showed, however, large variability between habitat types, with higher rates in spiders from dune grasslands than from dune slacks. This is suggested to result from differences in prey availability throughout the growing season between both habitat types.
4. Different size classes of spiders did not use different microhabitats, indicating that microhabitat segregation as a cannibalism-avoidance behaviour is absent in this species.     

Family Affects Sibling Cannibalism in the Black Widow Spider,Latrodectus hesperus

Adaptive foraging tactics are shaped by genes, the environment and gene–environment interactions. Because of relatively high levels of agonism toward conspecifics, spiders have been a popular focus for behavioral–ecological examinations of conspecific predation, or cannibalism. Surprisingly, studies examining the underlying, proximate assumption that cannibalism in spiders is a heritable trait shaped by interactions between genes and the environment are virtually non‐existent. Here, we examine the influence of family on the expression of sibling cannibalism in the post‐hatching, group‐living phase of an otherwise solitary, web‐building spider, the North American black widow (Latrodectus hesperus). Our results showed significant levels of variation in cannibalistic propensity among 26 sibships, with some families cannibalizing full sibs within 2 d and other families waiting 3 wk before resorting to cannibalism. A similar family‐level effect was evident in measures of sibling cohabitation, voracity toward cricket prey, and development speed. Negative correlations between maternal egg sac investment and offspring cannibalism suggest that this family effect may stem, at least in part, from a maternal effect, although we were not able to directly test the prediction that cannibalism is most common from spiderlings in poor condition. Thus, we present novel data suggesting family effects seem to be responsible for cannibalism in L. hesperus spiderlings; however, future work will be required to disentangle the relative importance of shared genes and shared maternal environment. We discuss several mechanisms that could explain the persistence of family‐level variation in cannibalism, a trait that seems likely to be subject to strong directional selection.     

Cannibalism and kin recognition in Delena cancerides (Araneae: Sparassidae), a social huntsman spider

Social behaviour in spiders is rare: of the 39 000 species of spiders known, only 23 are considered to be cooperatively social. Delena cancerides is a social species of the huntsman spider that is endemic to Australia. This species is virtually unique among social spiders, having evolved social behaviour in the absence of a snare web. It is thought that this form of social behaviour in D. cancerides has evolved via the sub-social route, that is, the extension of an ancestrally occurring period of maternal care and the delayed dispersal of juveniles. Most social spiders show no aggression towards non-kin conspecifics, prompting suggestions that spiders cannot recognize kin; however, D. cancerides individuals are highly aggressive towards conspecifics introduced from outside their own colony. In order to determine whether selective aggression in D. cancerides has its basis in kin recognition, tolerance behaviour was assessed in the context of kinship and size. We observed that, in general, juveniles preferred to starve than engage in cannibalism of any conspecifics, related or not. However, where cannibalism did occur, non-kin were preferentially eaten, indicating that this species is clearly capable of kin recognition. Size thresholds were also established, below which juveniles are tolerated by adults and above which aggressive interactions leading to death occur. We conclude that kin recognition and juvenile dispersal explain the uncharacteristically high levels of genetic polymorphism in this species.     

Ontogenetic constraints and diet shifts in Perch (Perca fluviatilis): mechanisms and consequences for intra‐cohort cannibalism

1. In many populations, sufficient size variation to allow for cannibalism may develop not only among age cohorts but also within them. Here, we used data on resource dynamics, consumer body size distribution and gape size limitation to unravel mechanisms promoting cannibalism within cohorts of young‐of‐the‐year (YOY) perch (Perca fluviatilis). 2. Perch are strongly gape limited when feeding on large zooplankton during early ontogeny. As a consequence, only initially large fish were able to shift to feeding on abundant large invertebrates, necessary to sustain fast growth. 3. We suggest that a combination of high initial size variation and exclusive access to resources for individuals with an initial size advantage is a prerequisite for the development of a size distribution sufficient for intra‐cohort cannibalism to occur. 4. During the time when cannibalism was observed, growth of the largest individuals in YOY perch cohorts was faster than that of smaller individuals. However, the energy gain from cannibalism did not increase growth rate enough to reach a size necessary to feed on more abundant size classes of victims, and therefore, the effect of cannibalism on overall cohort density was minor. 5. In addition to a high energy gain from cannibalism allowing for fast growth, strong resource limitation and slow growth rates of small individuals (i.e. potential victims) are a prerequisite not only for the development of intra‐cohort cannibalism but also for its persistence.     

Variation in offspring size within a population of the web-building spiderAgelena limbata

Within a population of the web-building spiderAgelena limbata, the weight of the first instar nymphs ranged from 1.187 to 6.559 mg. Both intraclutch and interclutch variation were recorded. The mean weights were different among clutches and the coefficients of variation within a clutch ranged from 3.3 to 29.2%. Variation in the nymphal weight was certainly derived from variation in the egg weight because there was a high correlation between the two weights. Factors affecting interclutch variation in nymphal weight were examined by multiple regression analysis. Nymphal weight was positively correlated with the body size and food conditions of female parents, and negatively correlated with the clutch size. Among these three factors, the food conditions of female parents had the largest apparent effect on the interclutch variation. The results suggest that females with larger body size and more food produce larger offspring, and that there is a trade-off between offspring size and clutch size. Heavier nymphs had larger body size (carapace width) and may have larger energy reserves. Heavier nymphs survived experimental starvation for a significantly longer period.     

Non-Random Sibling Cannibalism in the Marine Gastropod Crepidula coquimbensis

Sibling cannibalism is commonly observed in marine species. For instance, intrabrood cannibalism has been documented in marine gastropods with direct development, suggesting a relationship between embryo behavior and the evolution of life history strategies. However, there has been little effort to document the factors driving sibling cannibalism in marine species. The kin selection theory suggests that the level of relatedness plays an important role in cannibalism patterns. We examined Crepidula coquimbensis, a marine gastropod that broods and encloses its brooded offspring in capsules. Encapsulated embryos show sibling cannibalism and high levels of intracapsular multiple paternity. Given these features, cannibalistic behavior may be driven by kin-relatedness. To test this hypothesis, we constructed artificial aggregations of embryos to mimic three levels of relatedness: high, medium and low. For each category of aggregation, the cannibalism rate and benefits (i.e. size at hatching of surviving offspring) were estimated. In addition, at the end of embryo development, we performed parentage analyses to determine if cannibalism was associated with the relatedness between cannibal and victim embryos. Our results show that the intensity of sibling cannibalism increased in aggregations characterized by the lowest level of relatedness. There were important benefits of cannibalism in terms of hatching cannibal size. In addition, cannibalism between embryos was not random: the variation in reproductive success between males increased over the course of the experiment and the effective number of fathers decreased. Altogether, these results suggest that polyandry may play an important role in the evolution of sibling cannibalism in C. coquimbensis and that kin selection may operate during early embryonic stages in this species.     

Sexual Selection in the Water Spider: Female Choice and Male–Male Competition

The water spider Argyroneta aquatica is the only spider spending its whole life under water, and one of the few spider species in which males are larger than females. Previous studies indicated that males can cannibalize females, which is uncommon among spiders. Here we aimed to further test for a potential influence of sexual selection on male body size. We examined the importance of female choice by testing whether females prefer the larger of two simultaneously presented males as mating partners. Further, we examined the influence of male–male competition by comparing the fighting behaviour between large and small males when alone or when together with a female, and we determined the outcome of fights. We found that females approach and choose large males as mating partners, despite the risk of male cannibalism. Additionally, males intensively compete for females, and large males clearly win against smaller ones. Hence sexual selection seems to be important for the evolution of the peculiar sexual size dimorphism of water spiders, as large size is beneficial for males in both the intra‐ and intersexual context. Previous studies have suggested an important role of natural selection in the sex‐specific body size of water spiders, but natural and sexual selection mechanisms apparently work in the same direction, favouring large male size.     

Territoriality mediates atypical size‐symmetric cannibalism in the Amblypygi Phrynus longipes

Cannibalism can have important demographic and ecological effects on populations. Typically, cannibalism is size‐structured, where larger individuals eat smaller conspecifics. Initial cursory observations of the whip spider, Phrynus longipes, however, suggested that cannibalism might not be size‐structured in this species, perhaps because cannibalism is often a by‐product of territory contests. We staged paired interactions and recorded latency to escalate to physical aggression or cannibalize to understand the dynamics of cannibalism and resource contests. We employed a multimodel comparative approach to tease apart the contest characteristics that best predicted cannibalism during behavioral trials between P. longipes opponents. We found that, while armament size symmetry predicted escalation of contests, cannibalism was best predicted by body mass asymmetry. Further, cannibalism was most likely to occur among individuals similar in armament, but dissimilar in body mass. This suggests a discrepancy between phenotypes that may have evolved to communicate resource holding potential (e.g., armaments which benefit individuals if dishonest), and body mass as a cue of resource holding potential.     

Sibling cannibalism in aphidophagous ladybirds: its impact on sex-dependent development and body weight

Abstract:  The effects of sibling egg cannibalism on sex-dependent development and adult body weight were determined by rearing simultaneously two groups of sibling larvae of ladybirds, viz. Propylea dissecta and Coccinella transversalis individually. The first group included cannibals (i.e. neonates, which took a sibling egg as the first meal and later fed on aphids, Aphis craccivora ) and the other included non-cannibals (reared exclusively on aphids). The cannibal larvae developed faster with heavier adults than non-cannibals. This expedited development and nutritional advantage (increased body weight) was greater in first instars indicating maximum benefit of sibling cannibalism to them. Sibling cannibalism was relatively more advantageous to male than to female ladybirds. Laboratory data predict that the larger size of adults in the field could possibly be the result of sibling egg cannibalism at the neonate level.     

Development of size structure in tiger salamanders: the role of intraspecific interference

Cannibalism affects patterns of density-dependent mortality and may regulate population size. In many cases, rates of cannibalism depend on size structure, the frequency distribution of body sizes in the population, because cannibals can often only capture and consume smaller individuals. Size differences within single-age groups can be caused by a variety of factors. In this research we tested the hypothesis that size variation among larval tiger salamanders is due, in part, to interference interactions among individuals of different sizes. We found that size variation was greater when we raised larvae in groups rather than in isolation. This increase in size variation was due more to a relative deceleration of growth among smaller individuals rather than acceleration among larger individuals. We also found that smaller larvae had lower feeding rates than larger larvae when in groups, but not when isolated. Including spatial structure to limit physical interactions did not affect the size specificity of feeding rate, although it reduced feeding rates overall. We argue that these results are consistent with the hypothesis that larger larvae interfere, probably indirectly, with the feeding behavior of small individuals and this contributes to increases in size variation over time. We hypothesize that this indirect interference is caused by a behavioral response of smaller larvae to the risk of predation (cannibalism) by larger individuals. Received: 18 May 1998 / Accepted: 29 April 1999     

Differentiated egg size of the cannibalistic salamander <Emphasis Type="Italic">Hynobius retardatus</Emphasis>

Larvae of the salamander, Hynobius retardatus, are carnivorous, and even though there are two morphs, a typical morph and a broad-headed or “cannibal” morph, both are cannibalistic. They also sometimes eat other large prey, for example larvae of the frog, Rana pirica. In natural habitats, use of both conspecific and R. pirica larvae as food may contribute more strongly to high survival and substantially to fitness when larval densities are higher, because early-stage H. retardatus larvae sometimes experience scarcity of their typical prey. In cannibalistic oviparous amphibians, larger individuals that developed from larger eggs can more efficiently catch and consume larger prey and thus their survival may be better than that of smaller individuals developed from smaller eggs. Populations might therefore diverge in respect of egg size in response to variation in the density of conspecific and R. pirica larvae in natural ponds, with eggs being larger when larval density is higher. I examined how variance in hatchling size correlated with the incidence of cannibalism, and whether increasing larval density in natural ponds correlated with increasing egg size. Variance in initial larval body size facilitated cannibalism, and egg size increased as larval density in the ponds increased. In ponds with high larval density, where cannibalism and large prey consumption is a critical factor in offspring fitness, the production of fewer clutches with larger eggs, and thus of fewer and larger offspring, results in greater maternal fitness. Variation among the mean egg size in populations is likely to represent a shift in optimum egg size across larval density gradients.     

Games among cannibals: competition to cannibalize and parent-offspring conflict lead to increased sibling cannibalism

Sibling cannibalism occurs in many species, yet understanding of sibling cannibalism as an adaptation currently lags behind understanding of other antagonistic interactions among siblings. Observed sibling cannibalism phenotypes likely reflect the interaction between competitive games among siblings and parent-offspring conflict. Using a game-theoretic approach, we derive optimal offspring cannibalism behaviour and parental modifiers that limit or facilitate cannibalism. The results are compared to contemporary frequency-independent analysis. With the addition of game interactions among siblings or parent-offspring co-evolution, our model predicts increased cannibalism (compared to the frequency-independent prediction), as offspring compete to eat siblings. When infertile eggs are present--strengthening competition--offspring risk eating viable siblings in order to gain access to infertile eggs, intensifying parent-offspring conflict. We use the results to make new predictions about the occurrence of sibling cannibalism. Additionally, we demonstrate the utility of trophic egg laying as a maternal mechanism to promote egg eating.     

Synchrony during aggression in adult male Atlantic spotted dolphins <Emphasis Type="Italic">(Stenella frontalis)</Emphasis>

Synchrony among Atlantic spotted dolphins (Stenella frontalis) is crucial for successfully overcoming bottlenose dolphins (Tursiops truncatus) during interspecies aggression (Cusick and Herzing 2014). The present study examined synchrony in adult Atlantic spotted dolphins during aggressive encounters with bottlenose dolphins. Across group size, aggressive behaviors increased preceding synchrony, peaked during synchrony, and decreased dramatically after synchrony. Although smaller groups (< 10 dolphins) became synchronous more frequently than larger groups (> 10 dolphins), larger groups remained synchronous longer; however, smaller groups exhibited greater aggressive behaviors during synchrony, suggesting that additional aggressive behaviors may be necessary to compensate for the smaller group size, whereas larger groups may be able to rely on synchrony with less aggression. Disorganized squawk bouts synchronized as physical synchrony began, but only if coupled with escalating aggression.