Experimental manipulations of dispersal in the subsocial spider,Theridion pictum

Summary Theridion pictum (Walckenaer) is a subsocial spider, that is, the mother actively takes care of her offspring for an extended period of time after they emerge. It is demonstrated that spiderling dispersal is affected by the availability of food in the maternal web. Experimental manipulations indicate that in those webs where less food was provided, spiderlings dispersed significantly earlier and in significantly greater numbers. These results illustrate the importance of food availability as a permissive factor in the evolution of social spiders from subsocial ancestors. Food availability also affects the pattern of sex-biased philopatry and dispersal. Although both sexes are more likely to remain in the maternal web when more food is available, females are relatively more likely to do so. When less food is available, females are relatively more likely to disperse than are males.     

The selective advantages of cannibalism in a Neotropical mosquito

We investigated the benefits of larval cannibalism in the Neotropical mosquito Trichoprosopon digitatum. The clutch size of the mosquito in the field was strongly correlated with adult female size, indicating a fitness advantage to being large. In controlled laboratory experiments, we compared the survivorship and eventual adult sizes of larvae that were given the opportunity to cannibalise conspecifics throughout their lifetimes with the survivorship and adult sizes of larvae that were prevented from cannibalising. Since the benefits of cannibalism are likely to depend on the context in which it occurs, the experiment was conducted at two levels of alternative food availability. When food availability was high most larvae survived to adulthood, females cannibalised more than males and there was no measurable advantage to cannibalism in terms of survival rate, emergence time or adult size. Larvae were significantly more cannibalistic when food availability was lower, although under these conditions no larvae survived to adulthood. Nevertheless, under low food an important fitness benefit to cannibalism was revealed: individuals which had the opportunity to cannibalise survived significantly longer as larvae than those which did not. This increased longevity is likely to provide an important advantage to mosquito larvae when they wait for the input of unpredictable food sources. Received: 7 October 1995/Accepted after revision: 13 April 1996     

Factors influencing cannibalism in the wolf spider Pardosa agrestis (Araneae, Lycosidae)

Cannibalistic tendencies are well known in spiders and may be a significant factor influencing population size. The wolf spider, Pardosa agrestis, is the dominant non-web-building spider in a wide range of central European agricultural habitats. Preliminary field observations indicated an extended reproductive period, which results in a very wide size distribution of juvenile instars. We hypothesised that if cannibalism is enhanced by differences in size, especially during periods when prey is scarce, these populations might be susceptible to cannibalism in an ecologically significant way. Laboratory studies were conducted on juvenile P. agrestis in arenas. We analysed the following specific aspects of cannibalism: (1) the effect of the weight ratio between the opponents; (2) the effect of weight per se, and (3) the role of hunger level in determining cannibalistic tendencies of spiders. The role of weight and hunger were analysed in separate experiments, in both cases by controlling for the other variable. The results showed that cannibalism was strongly positively correlated with both weight ratio and hunger, but absolute size/age of an individual could not predict the occurrence of a cannibalistic event. These experiments generated the plausible hypothesis that cannibalism might be an important phenomenon in the regulation of real populations, which should be tested specifically in future field experiments. Received: 15 December 1997 / Accepted after revision: 10 October 1998     

Cooperative prey capture by young subsocial spiders: II. Behavioral mechanism

Subsocial spiders demonstrate an intermediate stage in the evolution of permanent sociality. Cooperative hunting is an important attribute of their sociality, but has not been documented in subsocial arthropods. After cannibalizing their mother, young of the subsocial spider Amaurobius ferox (Araneae, Amaurobiidae) remain together for several instars and feed communally. We monitored the collective prey capture behavior of the spiderlings. All the clutches showed collective capturing sequence (latency–orientation–moving–touching–seizing–feeding) toward the prey that was 10 times more massive than each individual. The first three individuals that exhibited attacking behavior were responsible for 90% of the total number of attacks, while 68% of the individuals within the group never exhibited attacks during the first 10 min following the introduction of prey into the communal web. First arriving individuals at the prey most often seized the antennae and legs of the prey, which probably facilitate access to the prey for subsequent individuals. The spiderlings that arrived later occupied more likely the abdomen and thorax, which contain more nutrition than the extremities occupied earlier. The individual apportionment of collective hunting behavior suggests a coordinated teamwork among individuals.     

Kin recognition by juvenile predatory mites: prior association or phenotype matching?

Kin recognition, the biased treatment of conspecific individuals based on genetic relatedness, is a widespread phenomenon in animals. The most common mechanisms mediating kin recognition are prior association (familiarity) and phenotype matching. Recognition based on prior association allows identifying familiar individuals. Recognition based on phenotype matching is an extension of prior-association-based recognition and allows identifying familiar and unfamiliar individuals due to a shared phenotypic trait. I investigated which of the two mechanisms is used by cannibalistic juvenile predatory mites, Phytoseiulus persimilis. Protonymphs that were associated with either siblings or non-kin discriminated familiar and unfamiliar larvae and preferentially cannibalized the latter irrespective of genetic relatedness. In contrast, despite previous association with either siblings or non-kin, protonymphs did not discriminate unfamiliar sibling and unfamiliar non-kin larvae. Association in the larval stage therefore mediated kin recognition based on familiarity, but not phenotype matching in cannibalistic P. persimilis protonymphs. Furthermore, in the presence of a familiar prey individual, sibling cannibalism occurred significantly sooner than non-kin cannibalism. This quick sibling cannibalism may have been the consequence of preferential association of siblings and/or may indicate the occurrence of an alternative cannibal phenotype. I discuss the adaptive significance of prior-association-based recognition for P. persimilis juveniles and emphasize the ability of P. persimilis to use multiple recognition mechanisms in dependence of the ontogeny and the ecological context.     

Social spider defense against kleptoparasitism

Summary Because of the large amount of webbing they provide, social spider colonies often host other satellite spider species referred to as kleptoparasites or food stealers. Such kleptoparasites may take advantage of increased prey capture rates associated with large spider aggregations. This study investigated the relationship between a cooperatively social spider species, Anelosimus eximius (Araneae: Theridiidae), which lives in the undergrowth of tropical rainforests in Peru, and its kleptoparasite, Argyrodes ululans (Araneae: Theridiidae), which specializes in foraging in An. eximius webs. Although large aggregates of spiders may be more attractive to kleptoparasites, the benefits of group defense may offset this cost. Natural colonies were observed, and enclosed field colonies containing fixed numbers of host spiders were manipulated in order to determine whether kleptoparasite success is affected by the number of social spiders that are available for defense. Prey was less likely to be stolen by Ar. ululans when a greater number of host An. eximius spiders were involved in prey capture. When hosts detected a kleptoparasite earlier and chased it more often, prey was more likely to be successfully defended. Ar. ululans was more successful in stealing small prey items in all colonies and gave up more readily on very large prey (> 11 mm). I conclude that communal living and group defense in An. eximius confer protection from the kleptoparasite Ar. ululans.     

Parent-offspring aggression in moorhens

Summary Colonial orb-weaving spiders from Mexico were studied to test predictions of risksensitive foraging theory: 1. group foraging increases prey capture/individual, and reduces prey variance; 2. spiders should be expected to exhibit risk-averse behavior (forage in groups) when the average level of prey exceeds individual needs, and exhibit risk-prone behavior (forage solitarily) when prey are searce. Laboratory and field studies show that group foraging increases capture efficiency and reduces variability in prey captured per spider. In desert/mesquite grassland habitat, where prey availability is low, M. atascadero forage solitarily in most cases. In tropical rainforest/agriculture sites, M. increassata forage in large colonies of thousands of webs. In intermediate habitats, M. spinipes forages solitarily or in groups, depending on prey availability. Over a range of sites with varying levels of prey, M. spinipes shifts from a risk-prone to a risk-averse group foraging strategy as prey increases.Group foraging behavior observed in colonial Metepeira fits the predictions of risk-sensitive foraging models. These findings explain why spiders tend to group webs together only in areas of superabundant prey. The role of risk-sensitivity in the evolution of coloniality in spiders is discussed.     

Age and sexual differences in the exploitation of two anthropogenic food resources for an opportunistic seabird

The availability of food resources has been suggested as a major factor in the substantial increase in reproductive output, survival, recruitment and, ultimately, population growth rates in most organisms. In fact, the artificial increase in food availability resulting from human activities has been suggested as a factor in the substantial increase in population size of several seabirds in recent decades. In the present study, our primary aim was to estimate the importance of the main natural prey and two alternative feeding resources, fishery discards and the invasive American crayfish Procambarus clarkii, for an opportunistic seabird, the Audouin’s gull Larus audouinii. We also assessed the influence of age and sex in the use of those three types of food. For this purpose, we compared the analyses of δ¹⁵N and δ¹³C in blood of male and female adults of known age and chicks with those in their potential prey. Our results reveal sex-related and age-related differences in the consumption of fish discards, small pelagic fish and American crayfish. Differences in the diet of males and females and also between adults and chicks could be related to different nutritional requirements. Age differences were probably related to their different foraging proficiency and the tendency of young breeders to opportunistically exploit anthropogenically derived food. This study illustrates the importance of considering the age and sex of individuals to obtain feasible dietary information and to understand how the exploitation of food of human origin could affect population growth.     

Space use in lions, <Emphasis Type="Italic">Panthera leo</Emphasis>, in the Selous Game Reserve: social and ecological factors

Habitat qualities, such as food supply or access to refuges, often influence home-range size. Furthermore, such qualitative differences usually lead to conspecific competition over space, which can be an important factor in determining the distribution of individuals within populations. In carnivores, patterns of resource dispersion are hypothesized to determine home range-size and group size. But in contests over space (or other resources), larger groups usually dominate smaller ones, and group size should therefore also affect home-range size. Here I describe the space use of lions, Panthera leo, in the Selous Game Reserve, Tanzania, and ask whether space use is related to pride size, habitat, or relatedness. Home ranges varied in size, but size showed no correlation to number of adult females in the pride or to habitat type. Lions exhibited a significant preference for riverine and short-grass habitat, and a significant avoidance of acacia woodland. Habitat preference ratios largely reflected prey availability in each habitat. Outer areas, as well as core areas of home ranges, were often used by two or more prides. Overlaps showed no correlation to relatedness among prides or habitat type. Thus, whereas home-range sizes and overlaps were determined by factors that could not be revealed by demographic factors or analyses of habitat composition or genetic structure, lion space use within each home range seemed driven mostly by prey availability, which mainly varies with habitat type.     

The behavioral ecology of cannibalism in cane toads (<Emphasis Type="Italic">Bufo marinus</Emphasis>)

Laboratory studies show that predatory cane toads (Bufo marinus) exhibit specialized toe-luring behavior that attracts smaller conspecifics, but field surveys of toad diet rarely record cannibalism. Our data resolve this paradox, showing that cannibalism is common under specific ecological conditions. In the wet–dry tropics of Australia, desiccation risk constrains recently metamorphosed toads to the edges of the natal pond. Juvenile toads large enough to consume their smaller conspecifics switch to a primarily cannibalistic diet (67% of prey biomass in stomachs of larger toads). Cannibalistic attack was triggered by prey movement, and (perhaps as an adaptive response to this threat) small (edible-sized) toads were virtually immobile at night (when cannibals were active). Smaller metamorphs were consumed more frequently than were larger conspecifics. The switch from insectivory to cannibalism reflects the high dry season densities of small conspecifics (in turn, due to desiccation-imposed constraints to dispersal) and the scarcity of alternative (insect) prey during dry weather. Our study pond (102 m in circumference) supported >400 juvenile toads, which consumed many metamorphs over the course of our study. Toads appear to be low-quality food items for other toads; in laboratory trials, juvenile toads that fed only on conspecifics grew less rapidly than those that ate an equivalent mass of insects. This effect was not due to parotoid gland toxins per se. Thus, cane toads switch to intensive cannibalism only when seasonal precipitation regimes increase encounter rates between large and small toads, while simultaneously reducing the availability of alternative prey.     

Using stable isotopes to reveal shifts in prey consumption by generalist predators.

The effectiveness of generalist predators in biological control may be diminished if increased availability of alternative prey causes individual predators to decrease their consumption of crop pests. Farming practices that enhance densities of microbidetritivores in the detrital food web can lead to increased densities of generalist predators that feed on pest species. The ability to predict the net biocontrol impact of increased predator densities depends upon knowing the extent to which individual predators may shift to detrital prey and feed less on crop pests when prey of the detritus-based food web are more abundant. We addressed this question by comparing ratios of stable isotopes of carbon (delta13C) and nitrogen (delta15N) in generalist ground predators and two types of prey (crop pests and microbidetritivores) in replicated 8 x 8 m cucurbit gardens subjected to one of two treatments: a detrital subsidy or no addition of detritus (control). Small sheet-web spiders (Linyphiidae) and small wolf spiders (Lycosidae) had delta13C values similar to those of Collembola in both the detrital and control treatments, indicating that small spiders belong primarily to the detrital food web. In control plots the larger generalist predators had delta13C values similar to those of the major insect pests, consistent with their known effectiveness as biocontrol agents. Adding detritus may have caused delta13C of one species of large wolf spider to shift toward that of the microbi-detritivores, although evidence is equivocal. In contrast, another large wolf spider displayed no shift in delta13C in the detrital treatment. Thus, stable isotopes revealed which generalist predators will likely continue to feed on pest species in the presence of greater densities of alternative prey.     

Mate quality,not aggressive spillover,explains sexual cannibalism in a size-dimorphic spider

Sexual cannibalism particularly before mating is costly for the male victim but also for the female aggressor if she risks remaining unmated. The aggressive spillover hypothesis explains the persistence of this behavior as a maladaptive side effect of positive selection on aggressiveness in a foraging context. The hypothesis predicts that the occurrence of sexual cannibalism is explained by female aggressiveness but is not related to male phenotype or behavioral type. An alternative hypothesis invokes sexual selection and makes the opposite prediction namely that sexual cannibalism is an expression of female choice and should hence mainly target males of low quality. We tested the above hypotheses on a sexually dimorphic nephilid spider Nephilengys livida, known for male monopolization of females via genital damage, female genital plugging, and mate guarding, by staging mating trials during which we recorded mating behaviors and occurrences of pre- and postcopulatory cannibalism. We did not restrict assessment of aggressiveness to the mating and foraging context but also included aggression against same sex conspecifics. To assess female personalities, i.e., consistent individual differences in behavior including aggressiveness, we repeatedly tested them for intra-sex aggression, voracity towards prey, locomotory activity, and boldness. Females exhibited consistent differences in intra-sex aggressiveness, latency to attack prey, and boldness. Aggressive females had shorter latencies to attack prey and were more active than non-aggressive ones. In contrast to the predictions of the aggressive spillover hypothesis, females that were aggressive towards prey and towards other females were not more likely to attack a male than non-aggressive females. In support of the mate choice hypothesis, less aggressive males were more likely attacked and cannibalized than more aggressive ones. This hints at sexual selection for aggressiveness in males and raises the question of mechanisms that maintain variation in male aggressiveness.     

Kleptoparasites: a twofold cost of group living for the colonial spider, Metepeira incrassata (Araneae, Araneidae)

Several species of kleptoparasitic and araneophagic spiders (Araneae: Family Theridiidae, Subfamily Argyrodinae) are found in colonial webs of the orb-weaving spider Metepeira incrassata (Araneae, Araneidae) from Mexico, where they steal food and/or prey upon their spider hosts. Census data from natural M. incrassata colonies reveal that the incidence of these species increases with colony size. This pattern may reflect the presence of several other orb-weaving spiders, each with their own kleptoparasitic species, invading larger M. incrassata colonies. As the number of these associated spiders increases, so does the density and number of Argyrodinae species in M. incrassata colonies, suggesting that associated spiders might reduce their own kleptoparasite load by building their webs within M. incrassata colonies. This represents a twofold cost to M. incrassata, as a field enclosure experiment revealed that a primarily kleptoparasitic species (Argyrodes elevatus) may reduce prey available to their hosts, but a kleptoparasitic/araneophagic species (Neospintharus concisus) inflicts high mortality upon M. incrassata. However, the cost of kleptoparasitism and predation by these species may be offset in part for M. incrassata individuals in large colonies by certain defensive mechanisms inherent in groups, i.e., “attack-abatement” and “selfish herd” effects. We conclude that increased occurrence of kleptoparasitic and/or predatory Argyrodinae spiders is a consequence of colonial web building and is an important potential cost of group living for colonial web-building spiders.     

Trade-off between pre- and postcopulatory sexual cannibalism in a wolf spider (Araneae,Lycosidae)

Sexual cannibalism can occur before, during or after mating. Relatively few experimental studies have examined why there is variation in the timing of sexual cannibalism. We examined the latency and number of attacks required for female spiders to capture male spiders pre- vs. postcopulation. We also examined the effects of female mating status and hunger level on the occurrence of pre- and postcopulatory cannibalism, which reflects the contribution of both relative capture success and female motivation to cannibalize males. Precopulatory cannibalism occurred after a shorter interval and required fewer chases and physical interactions for the female to successfully capture the male than was the case for postcopulatory cannibalism. Virgin females were more likely to engage in postcopulatory rather than precopulatory cannibalism and mated females vice versa. Those virgin females that did engage in precopulatory cannibalism had significantly lower body condition than virgin females engaging in postcopulatory cannibalism. While precopulatory cannibalism occurred more quickly and required fewer attacks by females, it comes at a potential cost of not mating with males. Hence, females are more likely to engage in precopulatory cannibalism if they have already mated or, if virgins, if they have low body condition. These results indicate that the decision of when to cannibalize males is dynamic and depends upon the relative value of a male as a mate versus a meal.     

Effects of combining an intraguild predator with a cannibalistic intermediate predator on a species-level trophic cascade

A greater diversity of natural enemies can in some cases disrupt prey suppression, particularly when natural enemies engage in intraguild predation, where natural enemies compete with and prey upon each other. However, empirical studies have often demonstrated enhanced prey suppression despite intraguild predation. A recent theoretical study proposed the hypothesis that, when the intermediate predator is cannibalistic, intraguild predation can reduce cannibalism within the intermediate predator population, leading to little change in intermediate predator mortality and thus enhanced prey suppression. The goal of this study was to examine this hypothesis empirically. Two summer-long field enclosure experiments were conducted in cotton fields. We investigated the effects of adding an intraguild predator, Zelus renardii, on (1) the abundance of a cannibalistic intermediate predator, Geocoris pallens, (2) the abundance of a herbivore, Lygus hesperus, and (3) cotton plant performance. G. pallens adult abundance did not increase, even when food availability was high and natural enemies were absent, suggesting that density-dependent cannibalism imposes an upper limit on its densities. Furthermore, although Z. renardii is an intraguild predator of G. pallens, G. pallens long-term densities were unaffected by Z. renardii. In the presence of the intermediate predator, the addition of the intraguild predator Z. renardii enhanced suppression of L. hesperus, and there were suggestions that Z. renardii and G. pallens partitioned the L. hesperus population. Effects of herbivore suppression cascaded to the plant level, improving plant performance. In conclusion, we provide empirical support for the hypothesis that the addition of an intraguild predator may enhance prey suppression if the intermediate predator expresses density-dependent cannibalism. Intraguild predation and cannibalism co-occur in many communities; thus their joint effects may be broadly important in shaping predator effects on herbivores and plant performance.     

Prior encounters with the opposite sex affect male and female mating behavior in a wolf spider (Araneae,Lycosidae)

Mate availability can vary widely in nature depending upon population density and sex ratio and can affect the ability of individuals to be selective in mate choice. We tested the effects of prior encounters with the opposite sex (i.e., exposure to the opposite sex either with or without mating) on subsequent mating behavior in two experiments that manipulated mate availability for both males and females in the wolf spider, Hogna helluo. The probability of mating in the experimental trial depended upon whether the prior encounter involved mating or not, and males and females responded in opposite directions. Exposure without mating resulted in a higher subsequent frequency of mating for females and a lower subsequent frequency of mating for males, while prior mating experience resulted in a lower frequency of female remating and a higher frequency of male remating. Prior exposure without mating did not affect female aggression. However, mated females engaged in precopulatory cannibalism more frequently than virgins. Mated males escaped postcopulatory cannibalism more frequently than virgins. Our results show that males respond to exposure without mating in the expected manner. However, prior mating (1 week earlier) had unexpected effects on males, which may be due to mated males being of higher quality. There were little or no effects of the size of the prior exposure individual or mate on subsequent mating behaviors. Further research is needed to determine why different species use different degrees of prior information in mate choice.     

Food availability,intrusion pressure and territory size: an experimental study of Anna's hummingbirds (Calypte anna)

Theoretical considerations implicate food availability and intrusion pressure as important determinants of territory size, but empirical studies have led to contradictory conclusions about cause-and-effect relationships among these three variables. To investigate this problem, we provided patches of electronically controlled artificial flowers, which were defended by male Calypte anna. Food availability was experimentally manipulated, and intrusion rate and territory size were calculated from behavioral observations of the territory owner. Changes in both food availability and intrusion rate were found to be significantly correlated with changes in territory size under certain conditions. Intrusion rate, which was influenced by food availability, was negatively associated with territory size so long as food availability was high. This association persisted even after possible effects of food availability were controlled statistically. Food availability was negatively correlated with territory size only when intrusion rates were high and after owners had been defending territories for 3 days. As food availability and intrusion rate increased, owners increasingly restricted their defense to the patch itself; partial regressions revealed a significant association for intrusion rate but not food availability. When intrusion rate was low and food availability varied from low to high levels, no relationship was observed between food availability and territory size, apparently because of opposing influences of food abundance on territory size. Correspondence to: P.W. Ewald     

Age-related shifts in the diet composition of southern elephant seals expand overall foraging niche

Southern elephant seals are important apex predators in a highly variable and unpredictable marine environment. In the presence of resource limitation, foraging behaviours evolve to reduce intra-specific competition increasing a species’ overall probability of successful foraging. We examined the diet of 141 (aged 1–3 years) juvenile southern elephant seals to test the hypotheses that differences between ages, sexes and seasons in diet structure occur. We described prey species composition for common squid and fish species and the mean size of cephalopod prey items for these age groups. Three cephalopod species dominated the stomach samples, Alluroteuthis antarcticus, Histioteuthis eltaninae and Slosarczykovia circumantarcticus. We found age-related differences in both species composition and size of larger prey species that probably relate to ontogenetic changes in diving ability and haul-out behaviour and prey availability. These changes in foraging behaviour and diet are hypothesised to reduce intra-specific food competition concomitant with the increase in foraging niche of growing juveniles.     

Interspecific comparisons of growth and diet among late larvae of three co-occurring clupeoid species in the Kii Channel,Japan

Growth and diet were compared among larvae of Japanese anchovy Engraulis japonicus, Japanese sardine Sardinops melanostictus and Pacific round herring Etrumeus teres. Compositions of prey items of the three species in the same month showed greater similarity than for the same species in different months. Prey size as well as prey taxa of the three species overlapped considerably with one another. Therefore, interspecific prey competition is likely in the case of limited food availability. The most abundant species tended to change from anchovy to round herring in early winter, from round herring to sardine in late winter and from sardine to anchovy in early spring, indicating a temporal segregation in use of the nursery grounds. Similar seasonal changes in growth rates were observed for the three species. Although interspecific prey competition is likely, the temporal segregation and similar temporal changes in growth rates could favor their coexistence.     

Trade-offs in foraging success and predation risk with spatial position in colonial spiders

Summary Colonial web-building spiders respond to trade-offs between selective forces relative to spatial position within colonies and thus provide support for the selfish herd theory. The size distribution of spiders within colonies of Metepeira incrassata, a colonial orb-weaver (Araneae: Araneidae) from tropical Mexico is nonrandom; larger (mature) spiders and females guarding eggsacs are more prevalent in the center, whereas more small (immature) spiders are found on the periphery. Experimental field studies with spiders of selected size classes show that larger spiders actively and aggressively seek protected positions in the center of the colony webbing, even though prey availability and capture rates are significantly higher on the periphery. Attacks by predatory wasps, other spiders, and hummingbirds are more frequent on the periphery than in the core of the colony. Reproductive females on the periphery are at greater risk because they are captured more often than smaller spiders, and if their egg sacs consequently remain unguarded, chances of cocoon parasitism are increased. As a result, spiders in the core of the colony have greater reproductive success, producing more egg sacs with greater hatching frequency. Colonial spiders thus appear to be making a trade-off between foraging and protection from predation and show a spatial organization predicted by the selfish herd theory. The influence of such trade-offs on individual fitness and the structure of colonies is discussed. Offprint requests to: G.W. Uetz