Prey Capture Behaviour in the Social Spider Anelosimus eximius (Araneae: Theridiidae): Responses to Prey Size and Type

Some species of web building spiders use different capture tactics for different prey types. The main factors influencing the attack behaviour are the ability of the insect to escape, the risks of injury to the spiders and prey size. This study evaluated the effects of size and prey type on prey capture behaviour of the social spider Anelosimus eximius as influenced by the number of spiders attracted by prey movements that did not bite until the immobilization (bystanders) and the number of spiders that contributed to prey immobilization (catchers). We carried out a two‐factor (prey size and type) experiment offering prey belonging to four orders: Diptera, Lepidoptera, Hymenoptera and Orthoptera, in a size gradient within each prey type. Both factors influenced the number of spiders recruited as bystanders, but only prey body size influenced the number of catchers in the subduing process. The possible advantages of the presence of bystanders around the interception site are discussed.     

Prey Capture and Feeding in the Social Spider Anelosimus eximius

Wild colonies of the social spider Anelosimus eximius (Araneae, Theridiidae) appear often to be food-limited and not all females come to reproduction. Using a limited number of marked females in an artificial colony, set up in the laboratory, this study attempts a first analysis of the participation in prey capture and ingestion. Marked females of the same age and experience were observed during the attack of prey insects, the ensuing transportation of the prey to the retreat, and the feeding session. No correlation was found between the time females spent hunting and the time they spent feeding. Females that laid eggs had fed longer and imbibed more nutrients, but had not hunted more than those females that did not reproduce. These, it is speculated, were denied access to the prey by the reproducing females.     

Coordination of Behavioral Sequences Between Individuals During Prey Capture in a Social Spider, Anelosimus eximius

The analysis of collaborative predation sequences performed by groups of 10 individuals (females) in a nonterritorial permanent-social spider, A. eximius, shows that prey-captures are organized in successive steps. Spiders begin by throwing sticky silk, which hinders the prey in the web; they then throw dry silk, which completes the immobilization of the prey. The third step is characterized by bites that paralyze the prey that will be then carried. A concordance test reveals a coordination of the individual's acts that explains the collaborative prey-capture efficiency. No individual specialization in one type of act has been shown. On the contrary, by using living preys or artificially dead vibrated preys, we show that all individuals have equipotential behaviors. Furthermore, each spider is able to adjust its behavior to the state of the prey. Individuals already involved in prey transportation can again display bites or sticky silk throwing if the prey is artificially vibrated. This mechanism, which corresponds to stimergic processes responsible for self-organized phenomena, already described in social insects, permits a coordination of individual acts without the recourse of direct communication. These results permit us to understand better how individuals coordinate their acts and lead us to support the hypothesis that the transition between solitary species and social species in spiders could have been sudden.     

Regulation of the number of spiders participating in collective prey transport in the social spider Anelosimus eximius (Araneae, Theridiidae)

In an experimental study, mechanisms by which cooperative prey transport is achieved in social spiders were clarified. Factors that could influence the number of individuals that participate in prey transport (prey mass, length and vibration) were investigated. Results show that two factors are fundamental: the vibrations and the prey length. Prey mass did not seem to influence spiders' participation. Thus, the single fact that individuals respond locally to environmental stimuli (intensity of vibration, available site on the prey) explains how spiders cooperate and efficiently capture a wide range of prey types without complex communication systems.     

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.     

Waxes of the social spider Anelosimus eximius (Araneae,Theridiidae): Abundance of novel n-propyl esters of long-chain methyl-branched fatty acids

The pentane extract of the social spider, Anelosimus eximius (Araneae, Theridiidae), contains hydrocarbons, fatty acids and their methyl esters, and a series of novel propyl esters of long-chain methyl-branched fatty acids. The propyl esters comprise almost three-fourths of the extract and consist predominantly of odd-numbered carbon chain components. Mass spectrometric analyses of the propyl esters, their methyl esters and cyanide derivatives showed that mono-, di- and trimethyl branched components with methyl branches on even numbered carbons predominate. The major components are propyl 4,20- and 4,30-dimethylhentriacontanoate and propyl 6,20- and 6,30-trimethylhentriacontanoate. The hydrocarbon fraction consists of n-, monomethyl- and dimethylalkanes, containing a relatively high proportion of even-numbered carbon chain components. The abundance of even-numbered carbon chain length alkanes and odd-numbered carbon chain length fatty acyl groups, along with abundant methyl-branches suggest that the propionyl-CoA and its carboxylated product, methylmalonyl-CoA, play important roles in the biosynthesis of these unique waxes. Arch. Insect Biochem. Physiol. 36:295–314, 1997. © 1997 Wiley-Liss, Inc.     

Natal Dispersal Patterns of a Subsocial Spider Anelosimus cf. jucundus (Theridiidae)

Species that alternate periods of solitary and social living may provide clues to the conditions that favor sociality. Social spiders probably originated from subsocial‐like ancestors, species in which siblings remain together for part of their life cycle but disperse prior to mating. Exploring the factors that lead to dispersal in subsocial species, but allow the development of large multigenerational colonies in social species, may provide insight into this transition. We studied the natal dispersal patterns of a subsocial spider, Anelosimus cf. jucundus, in Southeastern Arizona. In this population, spiders disperse from their natal nests in their penultimate and antepenultimate instars over a 3‐mo period. We tracked the natal dispersal of marked spiders at sites with clustered vs. isolated nests. We found that most spiders initially dispersed less than 5 m from their natal nests. Males and females, and spiders in patches with different densities of nests, dispersed similar distances. The fact that both sexes in a group dispersed, the lack of a sex difference in dispersal distance, and the relatively short distances dispersed are consistent with the hypothesis that natal dispersal results from resource competition within the natal nest, rather than inbreeding avoidance in competition for mates. Additionally, an increase in the average distance dispersed with time and with the number of spiders leaving a nest suggests that competition for nest sites in the vicinity of the natal nest may affect dispersal distances. The similar distances dispersed in patches with isolated vs. clustered nests, in contrast, suggest that competition among dispersers from different nests may not affect dispersal distances.     

Exploratory recruitment plasticity in a social spider (Anelosimus eximius)

Exploratory recruitment was investigated in an artificial experimental set-up on location in French Guyana. Groups of 200 freshly collected spiders of the neotropical social theridiid Anelosimus eximius were released on an open circular surface and offered a choice between two accessible shelters. Results indicated that a clear-cut collective decision was not always reached, unlike what we found using a different set-up in another set of experiments. Simulations were conducted using available information in order to explore the potential causes for this difference. Theoretical projections fit experimental data and strongly suggest that variability in the collective response results from a combination between modifications of the environment's properties and alteration of the recruitment procedure. Multiple variants of the theoretical set-up (including external bias) are investigated and emphasize plasticity in the collective response. New experimental studies are suggested and adaptative value of exploratory recruitment in social spiders is briefly discussed.     

The Comb-footed spider genera Theridion, Achaearanea and Anelosimus of Israel (Araneae: Theridiidae)

Israeli thendiid spiders of the genera Theridion, Achaearanea and Anelosimus have been revised. A relative richness in species is presented providing thereby updated information on the little known Mediterranean spider fauna. All type and non-type material previously described from the Middle East, deposited in several European collections has been re-examined, along with species from adjacent regions considered pertinent to the study undertaken. Altogether 21 species are recognized. Systematic, ecologic and all available zoogeographic information on taxa treated are discussed along with recent, pertaining literature. The presence of seven species formerly reported from Israel has been confirmed and the occurrence of another four species unknown hitherto from this region, has been proved. Some of these have never been adequately described or illustrated.
Ten new species are described: Theridion ochreolus, T. agaricographus, T hierwhonticus, T. jordanensis, T. negebensis, T. gekkonicus, T. dafnensis, T. vallisalinarum, T. pustiliferus and Anelosimus giladensis. The male of Theridion melanostictum is described for the first time. Keys, illustrations of diagnostic characters and records of distribution are provided for each species, all readily applicable also in adjacent countries. These may provide clues for better understanding of zoogeographic patterns of the Palearctic fauna, including those of the Old World Desert belt extending south and east of the Mediterranean region.     

Natural history of the non-solitary sheetweaving spider Anelosimus jucundus (Araneae: Theridiidae)

Field observations were made on one Panamanian population of Anelosimus jucundus. Almost all webs were found in a relatively exposed area, on small shrubs and composites. This is in contrast to A. eximius , a colonial congeneric which in our study site built its large webs along moist, fern-covered banks. Web structure varied with different plant substrates; for example, only some of the webs incorporated a clearly defined lower sheet. Webs were built by penultimate instar females, and perhaps by adult females and males. In some cases, several adult females inhabited a web, but they remained under separate leaf canopies; during census observations co-operative efforts were not noted. Prey consisted mainly of winged insects. Juvenile sex ratio was about equal. The degree of sociality demonstrated by A. jucundus is discussed.     

Revision and phylogenetic analysis of American ethicus and rupununi groups of Anelosimus (Araneae, Theridiidae)

Two species groups of the social spider genus Anelosimus are revised. The ethicus group contains six species found in South America, in an area ranging from the Guianas to southern Brazil and Argentina. Of these, A. rabus Levi, 1963, A. ethicus ( Keyserling, 1884 ), and A. nigrescens ( Keyserling, 1884 ) are redescribed, while A. nigrescens is removed from synonymy with A. ethicus . Three new species are described: A. misiones sp. nov., A. sumisolena sp. nov. and A. inhandava sp. nov. Anelosimus ethicus is reportedly either subsocial or solitary, while the behaviour of the other species in the group is unknown. The rupununi group contains two quasisocial species, A. rupununi Levi, 1956 and A. lorenzo Fowler & Levi, 1979, from the Caribbean and tropical South America. Both are redescribed here. A parsimony analysis of morphological characters provides support for the monophyly of both groups. In the phylogeny, subsociality optimizes to the base of Anelosimus , indicating that the common ancestor of the ethicus group was subsocial. Its members can thus be predicted to be subsocial, or secondarily solitary. Quasisociality arose de novo in the rupununi group, representing one of 6−7 independent origins in theridiids. Study of the biology of Anelosimus is important to advance our understanding of the evolution of sociality in spiders.     

Synchronized and rhythmical activity during the prey capture in the social spiderAnelosimus eximius (Araneae,Theridiidae)

Summary Anelosimus eximius is a social spider species of South America. Many individuals share the same web and participate in prey capture, taking some ten seconds to locate the prey in the silky structures. In the laboratory, we analyzed the movements of each spider which took part in the pursuit, and showed that they were both synchronized and rhythmical. Spiders alternate simultaneous periods of immobility (involving 100% of the attacking individuals) and activity (involving at least 70% of the spiders).The results are discussed with reference to the model developed by Goss and Deneubourg (1988) suggesting that autocatalysis may be the motor of certain synchronized and rhythmical activities in social arthropods.     

Carcass Decoration Changes Web Structure and Prey Capture Rate in an Orb-Web Spider, Cyclosa mulmeinensis (Araneae, Araneidae)

Cyclosa spiders attach prey carcasses as decorations to their webs, but the functions of the carcasses are unclear and controversial. We investigated distinctive features of these webs in the field and conducted prey-capture experiments in the lab using the orb-web spider Cyclosa mulmeinensis. Webs with attached decoration had a significantly narrower mesh width than those without decoration and a higher degree of vertical asymmetry in the web’s shape. In the laboratory, webs without decorations trapped significantly more prey, even though other features of the webs were nearly identical. These results suggest that web decorations do not attract prey in this species, but might play other roles such as blinding predators to the spider’s presence.     

Female body size, fecundity parameters and foundation of new colonies in Anelosimus jabaquara (Araneae, Theridiidae)

Summary. As in other social spider species, subadult Anelosimus jabaquara females found new colonies after solitary dispersal. Some individuals, however, usually remain and reproduce in their natal nests. To test the hypothesis that large females disperse more often than smaller ones, we compared the body size of A. jabaquara females that remained in their natal colonies with those that left to build solitary webs. We also compared clutch size, egg diameter, total egg volume and spiderling size in both conditions. Emigrating females were significantly larger and laid larger clutches. The smaller females that had not dispersed laid significantly larger eggs, although their total egg volume was lower. Spiderlings of solitary females were smaller and had a smaller range of size variation than those from colonies. We discuss the implications of these results in terms of costs and benefits of dispersion for spiders in different nutritional conditions.     

Madagascar: an unexpected hotspot of social Anelosimus spider diversity (Araneae: Theridiidae)

Abstract. The spider genus Anelosimus Simon, 1891 (Theridiidae) currently contains over forty described species, found worldwide in tropical to warm temperate areas. American Anelosimus are all social, a rare trait among spiders, but social behaviour has not been reported for Anelosimus species elsewhere. Old World Anelosimus are poorly known, both behaviourally and taxonomically, and no Anelosimus species have yet been described from sub-Saharan Africa or Madagascar. Based on a preliminary phylogenetic analysis we predicted sociality in an undescribed Madagascar species because it grouped among social New World species. An expedition to Madagascar then found no less than five undescribed periodic-social (subsocial) Anelosimus species in Périnet reserve. A sixth species from the same locality is known from museum specimens and the Anelosimus diversity of Périnet is comparable with the most diverse single locality in the Americas. Subsocial species play a key role in understanding the evolution of permanent sociality (quasisociality). This increased pool of available subsocial study species demonstrates the utility of phylogenies as predictors of traits in species thus far unstudied. Here, A. andasibe sp.n. , A. may Agnarsson sp.n. , A. nazariani sp.n. , A. sallee sp.n. , A. salut sp.n. and A. vondrona sp.n. are described. Anelosimus locketi Roberts, 1977 from Aldabra Atoll is a junior synonym of A. decaryi ( Fage, 1930 ) comb.n. from Madagascar. Preliminary data on the behaviour of the new species are given, indicating a level of sociality similar to the American A.‘arizona’¹. The phylogenetic analysis supports the monophyly of the Madagascar group and places it as sister to a clade containing the eximius lineage from the Americas, and a pair of undescribed Tanzanian species.     

Delayed juvenile dispersal benefits both mother and offspring in the cooperative spider Anelosimus studiosus (Araneae: Theridiidae)

Anelosimus studiosus juveniles usually remain in their natalwebs with their mothers until maturity, forming temporary coloniesin which individuals cooperate in web maintenance and preycapture. In a semi-natural environment, we experimentally removedjuveniles from their natal webs at mid-development. In thecontrol group, the juveniles were immediately replaced in theirnatal webs; in the experimental removal group, the juvenileswere not replaced and a sample of them were allowed to buildindividual webs. Colonies and solitary juveniles were exposedto natural prey densities and censused regularly for numbersand stages of spiders, and for prey capture. On average, juvenilesin colonies survived longer, developed faster, and had more resources per individual than did solitary juveniles. However,some of the solitary juveniles obtained more resources thanindividual juveniles in colonies. Mothers in the control groupsurvived longer and produced second broods earlier than mothersin the experimental removal group. Within the control group,older and larger colonies captured more and larger prey. Larger colonies had a lower coefficient of variation in prey capturedper juvenile. Overall, delayed juvenile dispersal benefitsboth juveniles and mothers.