A Continuum of Behavioral Plasticity in Urban and Desert Black Widows

Behavioral plasticity marks an individual's ability to modulate behavior across functional contexts. Behavioral syndromes, on the other hand, appear as consistent individual variation in behavior that is both repeatable for individuals within a functional context (e.g., consistent voracity toward prey) and correlated across contexts (e.g., high voracity toward prey and high levels of boldness toward enemies). Thus, adaptive plasticity and syndromes represent two extremes of a behavioral plasticity continuum upon which most behavioral phenotypes fall. We tested for both adaptive plasticity and behavioral syndromes in the western black widow spider, Latrodectus hesperus. We measured behavior in three contexts: startle, startle + prey, and startle + mate, and found (1) classic behaviorally plastic responses to predation risk, (2) high repeatability of behavior within contexts, and (3) evidence of a correlation between startle + prey and startle + mate contexts, indicative of a behavioral syndrome. As relative behavioral plasticity may vary across populations, we also compared urban and desert populations to test whether spiders from these habitats exhibit different behaviors and/or behavioral syndromes. While we found that urban males used in mating trials courted urban females significantly more than desert females, we found no other differences in the behavior of urban and desert black widows. Thus, black widows, regardless of habitat, are characterized by both context‐specific behavioral plasticity and across‐context correlations, presenting a phenotypic complexity that is likely exhibited, to varying degrees, by most organisms.     

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.     

Differences in aggression,activity and boldness between native and introduced populations of an invasive crayfish

Aggressiveness, along with foraging voracity and boldness, are key behavioral mechanisms underlying the competitive displacement and invasion success of exotic species. However, do aggressiveness, voracity and boldness of the invader depend on the presence of an ecologically similar native competitor in the invaded community? We conducted four behavioral assays to compare aggression, foraging voracity, threat response and boldness to forage under predation risk of multiple populations of exotic signal crayfish Pacifastacus leniusculus across its native and invaded range with and without a native congener, the Shasta crayfish P. fortis. We predicted that signal crayfish from the invaded range and sympatric with a native congener (IRS) should be more aggressive to outcompete a close competitor than populations from the native range (NR) or invaded range and allopatric to a native congener (IRA). Furthermore, we predicted that IRS populations of signal crayfish should be more voracious, but less bold to forage under predation risk since native predators and prey likely possess appropriate behavioral responses to the invader. Contrary to our predictions, results indicated that IRA signal crayfish were more aggressive towards conspecifics and more voracious and active foragers, yet also bolder to forage under predation risk in comparison to NR and IRS populations, which did not differ in behavior. Higher aggression/voracity/boldness was positively correlated with prey consumption rates, and hence potential impacts on prey. We suggest that the positive correlations between aggression/voracity/boldness are the result of an overall aggression syndrome. Results of stream surveys indicated that IRA streams have significantly lower prey biomass than in IRS streams, which may drive invading signal crayfish to be more aggressive/voracious/bold to acquire resources to establish a population.     

Temperature Mediates Shifts in Individual Aggressiveness,Activity Level,and Social Behavior in a Spider

Although in recent years behavioral syndromes have received a wealth of attention, how traits within syndromes respond to changing environments is not well resolved. Here, we test the effects of temperature on a suite of behavioral traits in the spider Anelosimus studiosus to determine (1) whether there are shifts in individuals’ social tendency, activity level, and foraging behavior in response to temperature, (2) if these traits shift are in the direction predicted by within‐population axes of trait covariance, and (3) whether the effects of temperature differ among individuals. In previous work, we documented a behavioral syndrome in A. studiosus where increased tolerance of conspecifics is correlated with decreased activity level and aggressiveness toward prey. Furthermore, there are distinct among‐population differences in behavior, where individuals from warm sites tend to be more aggressive and active than individuals from cold sites. Our data here reveal that at warmer temperatures A. studiosus exhibit diminished tolerance of conspecifics, increased activity levels, shorter latencies of attack, and increased tendencies to attack multiple prey items. Furthermore, we found that individual differences in behavior were consistent across temperature regimes for the majority of behavioral traits considered here: social tendency, activity level, and latency of attack. These findings are consistent with the hypothesis that these behaviors are linked together by shared genetic underpinnings (e.g., metabolic differences) and shift non‐independently in response to contemporary abiotic environment (i.e., temperature). Furthermore, our data suggest that temperature itself could be responsible for the among‐population variation in social structure in A. studiosus.     

Challenging the Aggressive Spillover Hypothesis: Is Pre‐Copulatory Sexual Cannibalism a Part of a Behavioural Syndrome?

Pre‐copulatory cannibalism – females devouring males during courtship – may bring no benefit to either sex. The ‘aggressive spillover hypothesis’ (ASH) posits that pre‐copulatory cannibalism represents a spillover of female aggressiveness from the juvenile foraging context, when aggressiveness is advantageous, to the adult mating context, when aggressiveness may be non‐adaptive or maladaptive. The ASH suggests that individuals exhibit limited plasticity in aggressive behaviours because they are genetically canalised for indiscriminate aggressiveness towards prey and conspecifics, including males. Hence, a tendency to employ pre‐copulatory cannibalism is a part of the female aggression syndrome, an assertion generally accepted in the personality field. We here re‐evaluate the previous findings in the light of personality criteria, which we propose for ASH validation: between‐individual differences, repeatability and heritability in tendency for pre‐copulatory attacks (and pre‐copulatory cannibalism) and voracity towards prey, and their correlation. To re‐evaluate ASH and to allow for additional or alternative explanations, we ask whether pre‐copulatory cannibalism depends on female hunger, mating status, size and/or male quality. Finally, we ask whether cannibalistic females have a reduced reproductive success as predicted by the ASH. While repeatability and heritability in voracity towards prey and its correlation with the tendency to engage in pre‐copulatory cannibalism were found in certain systems, we lack any evidence for repeatability and heritability in pre‐copulatory cannibalistic attempts and for its maladaptiveness. Rather than only resorting to the ASH, foraging and mate choice hypotheses may also explain pre‐copulatory cannibalism. We suggest clarifying the use of the terms sexual cannibalism (effect) and female aggressiveness or tendency to attack and devour males (cause), and argue that male strategies to avoid cannibalism should be considered. We propose testing the ASH as the explanation for pre‐copulatory cannibalism in those cases where female tendency to devour males correlates with actual pre‐copulatory cannibalism and when all the above criteria are fulfilled. Finally, we propose future directions for studying the ASH.     

Daring, risk assessment and body condition interactions in steppe buzzards Buteo buteo vulpinus

Trading-off food and safety is considered a predominant component of foraging behavior. We examined whether predators' response to potential prey is state-dependent, and suggest that risk of injury while pursuing the prey affects the predators' foraging behavior in a similar manner as predation risk affects prey foraging behavior. We examined how the daring of steppe buzzards Buteo buteo vulpinus in pursuing their prey varied with body condition. We measured the time it took juvenile buzzards to attack a mouse placed in a Bal-Chatri cage trap and related it to their body condition. We found that buzzards in poorer body condition tended to hesitate less before pursuing their prey than did buzzards in better body condition. We suggest that behavioral decisions of buzzards are state-dependant and are influenced by the risk of injury involved in pursuing their prey.     

Advantages of social foraging in crab spiders: Groups capture more and larger prey despite the absence of a web

Among group‐living spiders, subsocial representatives in the family of crab spiders (Thomisidae) are a special case, as they build protective communal leaf nests instead of extensive communal capture webs. It could thus be inferred that antipredator benefits (e.g., enhanced protection in larger nests) rather than foraging‐related advantages (e.g., capture of more and larger prey) promote sociality in this family. Nonetheless, subsocial crab spiders do share prey, and if this behaviour does not reflect mere food scramble but has a cooperative character, crab spiders may offer insights into the evolution of social foraging applicable to many other cooperative predators that hunt without traps. Here, we performed a comparative laboratory feeding experiment on three of the four subsocial crab spider species—Australomisidia ergandros, Australomisidia socialis and Xysticus bimaculatus—to determine if crab spiders derive advantages from foraging in groups. In particular, we tested artificially composed groups of five sibling spiderlings vs. single siblings in terms of prey capture success and prey size preference. Across species, groups had higher prey capture success (measured in terms of capture rates and capture latency) and were more likely to attack large, sharable prey—dynamics leading to reduced food competition among group members in favour of living and foraging in groups. Within groups, we further compared prey extraction efficiency among the three applied social foraging tactics: producing, scrounging and feeding alone. In A. ergandros, individuals were exceptionally efficient when using the non‐cooperative scrounger tactic, which entails feeding on the prey provided by others. Thus, our multispecies comparison confirms foraging advantages in maintaining a cooperative lifestyle for crab spiders, but also demonstrates the relevance of research into exploitation of cooperative foraging in this family.     

Previous foraging success influences web building in the spider Stegodyphus lineatus (Eresidae)

Stegodyphus lineatus (Eresidae) is a desert spider that buildsan aerial capture web on bushes in the Negev desert in southernIsrael. Web building for spiders is costly in energy, time,and risk of predation. Spiders should trade-off these costswith the benefits in terms of prey capture. We tested the hypothesisthat the previous foraging success of the spider influencesthe effort invested in foraging. Specifically, we asked whetheran increase in food intake causes spiders to reduce web renewalactivity and web size. Alternatively, time constraints on foragingand development, resulting from a short growing season, couldinduce spiders to continue foraging even when supplemented withprey. The cost of web building was measured as time and massloss. To build an average size web (about 150 cm²), we calculatedthat a spider requires 6 h and that spiders lose 3%-7% of their weight.In field experiments, spiders responded differently to food supplementationin 2 different years. In 1994, they improved their condition comparedto individuals whose webs were removed to reduce foraging opportunitiesand compared to control spiders. In 1995, spiders tested earlier inthe season than the previous year did not improve their conditionin response to prey supplementation. Nonetheless, in both years, food-supplementedspiders built significantly smaller webs than food-deprived andcontrol spiders. This result was confirmed in a laboratory experiment whereprey intake was controlled. We conclude that for S. lineatus immediateforaging risks outweigh the potential time constraints on foraging.     

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.     

Link between Aggressiveness and Shyness in the Spider <Emphasis Type="Italic">Philodromus albidus</Emphasis> (Araneae,Philodromidae): State Dependency over Stability

Behavioral syndromes, seen as correlations among two or more functionally different behaviors, are well documented in many different animal taxa. They may not be present automatically, however, and their consistency within populations and individuals also varies among studies. Here, we studied a behavioral syndrome comprising foraging aggressiveness and boldness/shyness and its time consistency in the cursorial spider Philodromus albidus. We measured foraging aggressiveness as the number of prey killed per 2-hour period. Boldness/shyness was assessed as a latency of initiating exploration in a novel environment. We found the presence of the behavioral syndrome in P. albidus, as bold individuals were also more aggressive than were shy individuals. The syndrome was consistent through time within the population but not consistent for individuals. We further discuss the possible causes of the pattern.     

Genetically-based variation between two spider populations in foraging behavior

Summary Optimal foraging theory is based on the assumption that at least some aspects of foraging behavior are genetically determined (Pyke et al. 1977; Kamil and Sargent 1980; Pyke 1984). Nonetheless, very few studies have examined the role of genetics in foraging behavior. Here, we report on geographical differences in the foraging behavior of a spider (Agelenopsis aperta) and investigate whether these differences are genetically determined. Field studies were conducted on two different populations of A. aperta: one residing in a desert riparian habitat, and the other in a desert grassland habitat. Data from the spiders' natural encounters with prey demonstrated that grassland spiders exhibited a higher frequency of attack than riparian spiders towards 13 of 15 prey types, including crickets and ants. Grassland spiders also had shorter latencies to attack 12 of 15 prey types, including crickets and ants, than riparian spiders. Subsequently, we reared grassland and riparian spiders under controlled conditions in the laboratory and observed their interactions with prey to determine whether the populational differences we found in the field could be genetic. Again, grassland spiders showed a shorter latency to attack prey (crickets, ants) than riparian spiders. These latencies were not significantly affected by the hunger state or age of the spiders. Finally, we reared a second generation (F2) of grassland and riparian spiders in the laboratory and observed their interactions with prey to determine whether the populational differences in the previous generation were due to genetic effects or maternal effects. As before, grassland spiders exhibited a shorter latency to attack prey (crickets) than riparian spiders. We conclude that the foraging differences we observed between these two populations of A. aperta are genetically determined. These differences probably have resulted from either natural selection acting directly on attack frequency and the latency to attack prey, or natural selection acting on traits which are genetically correlated with these aspects of foraging behavior.     

Task specialization in two social spiders,Stegodyphus sarasinorum (Eresidae) and Anelosimus eximius (Theridiidae)

Understanding the social organization of group‐living organisms is crucial for the comprehension of the underlying selective mechanisms involved in the evolution of cooperation. Division of labour and caste formation is restricted to eusocial organisms, but behavioural asymmetries and reproductive skew is common in other group‐living animals. Permanently, social spiders form highly related groups with reproductive skew and communal brood care. We investigated task differentiation in nonreproductive tasks in two permanently and independently derived social spider species asking the following questions: Do individual spiders vary consistently in their propensity to engage in prey attack? Are individual spiders' propensities to engage in web maintenance behaviour influenced by their previous engagement in prey attack? Interestingly, we found that both species showed some degree of task specialization, but in distinctly different ways: Stegodyphus sarasinorum showed behavioural asymmetries at the individual level, that is, individual spiders that had attacked prey once were more likely to attack prey again, independent of their body size or hunger level. In contrast, Anelosimus eximius showed no individual specialization, but showed differentiation according to instar, where adult and subadult females were more likely to engage in prey attack than were juveniles. We found no evidence for division of labour between prey attack and web maintenance. Different solutions to achieve task differentiation in prey attack for the two species studied here suggest an adaptive value of task specialization in foraging for social spiders.     

Differences in prey personality mediate trophic cascades

Functional trait approaches in ecology chiefly assume the mean trait value of a population adequately predicts the outcome of species interactions. Yet this assumption ignores substantial trait variation among individuals within a population, which can have a profound effect on community structure and function. We explored individual trait variation through the lens of animal personality to test whether among‐individual variation in prey behavior mediates trophic interactions. We quantified the structure of personalities within a population of generalist grasshoppers and examined, through a number of field and laboratory‐based experiments, how personality types could impact tri‐trophic interactions in a food chain. Unlike other studies of this nature, we used spatial habitat domains to evaluate how personality types mechanistically map to behaviors relevant in predator–prey dynamics and found shy and bold individuals differed in both their habitat use and foraging strategy under predation risk by a sit‐and‐wait spider predator. In the field‐based mesocosm portion of our study, we found experimental populations of personality types differed in their trophic impact, demonstrating that prey personality can mediate trophic cascades. We found no differences in respiration rates or body size between personality types used in the mesocosm experiment, indicating relative differences in trophic impact were not due to variation in prey physiology but rather variation in behavioral strategies. Our work demonstrates how embracing the complexity of individual trait variation can offer mechanistically richer understanding of the processes underlying trophic interactions.     

The Effect of Feeding History on Prey Capture Behaviour in the Orbweb Spider Argiope keyserlingi Karsch (Araneae: Araneidae)

The prey capture behaviour of the orb-web spider Argiope keyserlingi Karsch was examined experimentally by subjecting spiders to two different feeding regimes (food deprived and food satiated) and three types of prey: Drosophila, blowflies (Lucilia cuprina) and bees (Apis mellifera). The attack behaviour of the spiders was influenced by both their foraging history and the type of prey. Food deprived spiders attacked Drosophila and bees more frequently than food satiated spiders, and food satiated spiders travelled more slowly to any of the prey types than food deprived spiders. Furthermore, Drosophila were never wrapped in silk but only grasped with the chelicerae, whereas both blowflies and bees were always wrapped. This provides experimental confirmation that feeding history affects the decision of orb-web spiders to accept or reject any given prey.     

How does the presence of a conspecific individual change the behavioral game that a predator plays with its prey?

Behavioral games predators play among themselves may have profound effects on behavioral games predators play with their prey. We studied the behavioral game between predators and prey within the framework of social foraging among predators. We tested how conspecific interactions among predators (little egret) change the predator–prey behavioral game and foraging success. To do so, we examined foraging behavior of egrets alone and in pairs (male and female) in a specially designed aviary consisting of three equally spaced pools with identical initial prey (comet goldfish) densities. Each pool was comprised of a risky microhabitat, rich with food, and a safe microhabitat with no food, forcing the fish to trade off food and safety. When faced with two versus one egret, we found that fish significantly reduced activity in the risky habitat. Egrets in pairs suffered reduced foraging success (negative intraspecific density dependence) and responded to fish behavior and to their conspecific by changing their visiting regime at the different pools—having shorter, more frequent visits. The time egret spent on each visit allowed them to match their long-term capture success rate across the environment to their capture success rate in the pool, which satisfies one aspect of optimality. Overall, egrets in pairs allocated more time for foraging and changed their foraging tactics to focus more on fish under cover and fish ‘peeping’ out from their shelter. These results suggest that both prey and predator show behavioral flexibility and can adjust to changing conditions as needed in this foraging game.     

Interpopulation Variations in Behavioral Syndromes of a Jumping Spider from Insecticide‐Treated and Insecticide‐Free Orchards

Variations in environmental conditions can influence behavioral syndromes (correlated tendencies in behaviors), and understanding the factors that shape trait covariation is particularly relevant when species are challenged by environmental changes. We investigated how behavioral syndromes varied at extremes of a gradient of anthropogenic disturbance, using apple orchards with different histories of insecticidal applications as a model system. Eris militaris (Araneae: Salticidae) jumping spiders were sampled from an insecticide‐free orchard and an insecticide‐treated orchard from Southern Québec. Spiders were tested for activity, aggression, boldness, and voracity under standardized conditions. Behavioral syndrome structure was compared between the two populations using Bayesian multiresponse models and structural equation modeling. Syndrome structure differed significantly between the two populations. The insecticide‐free population showed evidence of a syndrome involving all measured traits, while only aggression, boldness and voracity were correlated in the insecticide‐treated population. The insecticide‐free population showed negative correlations between active and voracious behavioral types vs. aggressive and bold types while the insecticide‐treated population showed a negative correlation between aggression‐boldness and voracity. This research is a first step in investigating the impact of anthropogenic disturbances on behavioral syndromes and demonstrates that behavioral syndromes may vary with respect to insecticidal applications.     

Prey‐specific foraging tactics in a web‐building spider

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Taking a trip to the shelf: Behavioral decisions are mediated by the proximity to foraging habitats in the black‐legged kittiwake

For marine top predators like seabirds, the oceans represent a multitude of habitats regarding oceanographic conditions and food availability. Worldwide, these marine habitats are being altered by changes in climate and increased anthropogenic impact. This is causing a growing concern on how seabird populations might adapt to these changes. Understanding how seabird populations respond to fluctuating environmental conditions and to what extent behavioral flexibility can buffer variations in food availability can help predict how seabirds may cope with changes in the marine environment. Such knowledge is important to implement proper long‐term conservation measures intended to protect marine predators. We explored behavioral flexibility in choice of foraging habitat of chick‐rearing black‐legged kittiwakes Rissa tridactyla during multiple years. By comparing foraging behavior of individuals from two colonies with large differences in oceanographic conditions and distances to predictable feeding areas at the Norwegian shelf break, we investigated how foraging decisions are related to intrinsic and extrinsic factors. We found that proximity to the shelf break determined which factors drove the decision to forage there. At the colony near the shelf break, time of departure from the colony and wind speed were most important in driving the choice of habitat. At the colony farther from the shelf break, the decision to forage there was driven by adult body condition. Birds furthermore adjusted foraging behavior metrics according to time of the day, weather conditions, body condition, and the age of the chicks. The study shows that kittiwakes have high degree of flexibility in their behavioral response to a variable marine environment, which might help them buffer changes in prey distribution around the colonies. The flexibility is, however, dependent on the availability of foraging habitats near the colony.     

Foraging speed in staging flocks of semipalmated sandpipers: evidence for scramble competition

Foraging speed is a key determinant of fitness affecting both foraging success and predator attack survival. In a scramble for food, for instance, evolutionary stable strategy models predict that speed should increase with competitor density and decrease when the risk of attack by predators increases. Foraging speed should also decrease in richer food patches where the level of competition is reduced. I tested these predictions in fall staging flocks of semipalmated sandpipers (Calidris pusilla) foraging for an evasive prey. Capture rate of these prey decreased with sandpiper density as the presence of competitors reduced the availability of resources for those behind. Foraging speed was evaluated indirectly by measuring the time needed to cross fixed boundaries on mudflats over 6 years. As predicted, foraging speed increased with sandpiper density and decreased with food density, but, unexpectedly, increased closer to obstructive cover where predation risk was deemed higher. When foraging closer to cover, from where predators launch surprise attacks, the increase in foraging speed may compensate for an increase in false alarms that interrupted foraging. While foraging in denser flocks decreases foraging success, joining such flocks may also increase safety against predators. In semipalmated sandpipers that occupy an intermediate position in the food chain, foraging behavior is influenced simultaneously by the evasive responses of their prey and by the risk of attack from their own predators.     

Foraging Benefits in a Colour Polymorphic Neotropical Orb Web Spider

Conspicuous body colouration in sedentary predators such as orb web spiders seems paradoxical as potential prey can see and avoid the webs. Several studies have demonstrated that rather than deterring prey, the colours act as sensory traps for flower‐seeking insects. In chromatically polymorphic species, the existence of more than one colour morph may lead to differing levels of prey attraction. To explore these issues, we studied a neotropical orb web spider, Verrucosa arenata, which shows colour polymorphism with predominantly white or yellow abdomen colours. We asked whether a particular morph is dominant in the population, and whether a particular morph is associated with enhanced foraging success and body condition. Here we showed that although yellow morphs attracted more prey, white morphs were in better body condition. We showed that model prey such as honeybees are able to discriminate between the morphs. We discuss these findings in relation to the functional significance of bright body colouration and colour polymorphism in spiders.