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.     

Relatedness and genetic structure in a socially polymorphic population of the spider Anelosimus studiosus

The evolution of sociality remains a challenge in evolutionary biology and a central question is whether association between kin is a critical factor favouring the evolution of cooperation. This study examines genetic structure of Anelosimus studiosus, a spider exhibiting polymorphic social behaviour. Two phenotypes have been identified: an ‘asocial’ phenotype with solitary female nests and a ‘social’ phenotype with multi‐female/communal nests. To address the questions of whether these phenotypes are differentiated populations and whether cooperative individuals are closely related, we used microsatellites to analyse individuals from both communal and solitary nests. We found no evidence of differentiation between social and solitary samples, implying high rates of interbreeding. This is consistent with the hypothesis that these phenotypes coexist as a behavioural polymorphism within populations. Pairwise relatedness coefficients were used to test whether cooperating individuals are more closely related than expected by chance. Pairwise relatedness of females sharing communal webs averaged 0.25, the level expected for half‐siblings and significantly more closely related than random pairs from the population. Solitary females collected at similar distances to the communal spider pairs were also more closely related than expected by chance (mean relatedness = 0.18), but less related than social pairs. These results imply that low dispersal contributes to increase likelihood of interaction between kin, but relatedness between social pairs is not explained by spatial structure alone. We propose that these phenotypes represent stages in the evolution of sociality, where viscous population structure creates opportunities for kin selection and cooperation is favoured under certain environmental conditions.     

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.     

Extended parental care and delayed dispersal: northern, tropical, and southern passerines compared

Using modern comparative methods, we found that both time toindependence and time with parents were significantly longerin southern hemisphere and tropical birds than in northern hemisphereones. These differences held even after removing Australianpasserines or cooperatively breeding species, and they do notdepend on habitat, diet, or migration pattern. In southern hemisphereand tropical regions, both cooperative breeding and non-cooperativeparents continue to feed their young for a similar length oftime, but cooperative breeders allow them to stay longer intheir natal territory after they become nutritionally independent.Nevertheless, the young of non-cooperative species stay longerwith their parents than do the young of non-cooperative speciesin the temperate northern hemisphere. The fact that extendedperiods of post-fledging parental care are widespread amongpasserines provides further empirical support for the view thatlife histories of southern and tropical birds are ‘slow,’with small clutches, extended parental care, and long lifespan;parents take care of fewer young for longer. These results supportrecent theoretical models that predict that high adult survivaland low turnover of territory owners generally favor natal philopatry.We suggest that the reasons why some species (with or withoutcooperative breeding) exhibit natal philopatry and others donot lie in the balance between productivity and survival ofadults and of retained or dispersing offspring.     

The phylogeny of the social Anelosimus spiders (Araneae: Theridiidae) inferred from six molecular loci and morphology

We use fragments of three nuclear genes (Histone 3, 18SrDNA, and 28SrDNA) and three mitochondrial genes (16SrDNA, ND1, and COI) totalling approximately 4.5kb, in addition to morphological data, to estimate the phylogenetic relationships among Anelosimus spiders, well known for their sociality. The analysis includes 67 individuals representing 23 of the 53 currently recognized Anelosimus species and all species groups previously recognized by morphological evidence. We analyse the data using Bayesian, maximum likelihood, and parsimony methods, considering the genes individually as well as combined (mitochondrial, nuclear, and both combined) in addition to a 'total evidence' analysis including morphology. Most of the data partitions are congruent in agreeing on several fundamental aspects of the phylogeny, and the combined molecular data yield a tree broadly similar to an existing morphological hypothesis. We argue that such congruence among data partitions is an important indicator of support that may go undetected by standard robustness estimators. Our results strongly support Anelosimus monophyly, and the monophyly of the recently revised American 'eximius lineage', although slightly altered by excluding A. pacificus. There was consistent support for the scattering of American Anelosimus species in three clades suggesting intercontinental dispersal. Several recently described species are reconstructed as monophyletic, supporting taxonomic decisions based on morphology and behaviour in this taxonomically difficult group. Corroborating previous results from morphology, the molecular data suggest that social species are scattered across the genus and thus that sociality has evolved multiple times, a significant finding for exploring the causes and consequences of social evolution in this group of organisms.     

The spider genus Enoplognatha (Araneae: Theridiidae) in Israel

Israeli theridiid spiders of the genus Enoplognatha are revised. Five species, four of them new, each represented by the two sexes, are recognized, and their affinities with similar species from adjacent regions are discussed. All species previously described from this region have been re-examined. The occurrence of E. mandibularis , formerly reported from Israel, has been confirmed, while the record of E. ovata has proved erroneous. A key to the species is provided.     

Morphological phylogeny of cobweb spiders and their relatives (Araneae, Araneoidea, Theridiidae)

This paper offers the first cladistic analysis of a wide selection of theridiid genera based on morphological data. The analysis treats 53 theridiid taxa representing 32 genera (Achaearanea, Anelosimus, Ameridion, Argyrodes, Ariamnes, Carniella, Cerocida, Chrysso, Coleosoma, Dipoena, Emertonella, Enoplognatha, Episinus, Euryopis, Faiditus, Kochiura, Latrodectus, Neospintharus, Nesticodes, Pholcomma, Phoroncidia, Rhomphaea, Robertus, Selkirkiella, Spintharus, Steatoda, Stemmops, Theridion, Theridula, Thymoites, Thwaitesia, Tidarren) and eight outgroup taxa representing the families Nesticidae (Eidmanella and Nesticus), Synotaxidae (Synotaxus, two species), Pimoidae (Pimoa), Linyphiidae (Linyphia), Tetragnathidae (Tetragnatha) and Araneidae (Argiope). The parsimony analysis of 242 morphological and behavioural characters found a single, most parsimonious tree. The monophyly of theridiids and their sister relationship with nesticids is strongly supported. The recent resurrection of Ariamnes and Rhomphaea from Argyrodes made the latter paraphyletic. However, Ariamnes and Rhomphaea are characterized by an array of characters, and Argyrodes still contains dramatically distinct clades for which names are available: Faiditus (removed from synonymy ? RS) and Neospintharus (RS). These revalidations provide a classification with greater information content and utility. These three genera, along with Ariamnes, Rhomphaea and Spheropistha, comprise the subfamily Argyrodinae. The monophyly and composition of the subfamilies Hadrotarsinae, Spintharinae, Pholcommatinae, Latrodectinae and Theridiinae are discussed. Theridion is paraphyletic and in need of revision. Anelosimus as currently circumscribed is paraphyletic, a problem resolved by revalidating Selkirkiella (RS) and Kochiura (RS). Numerous new combinations are established. The results suggest the monophyletic origin of both kleptoparasitism and araneophagy in the lineage leading to Argyrodinae, negating hypotheses that either arose from the other. Sociality evolved multiple times within the family, accounting for as much as one fourth of the origins of social behaviour among all spiders. No losses of sociality are implied. The hypothesis of maternal care as the pathway to sociality receives support. Evolution of theridiid webs is complex, with multiple modifications and loss of the basic theridiid cobweb. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141 , 447–626.     

Population structure and dispersal in a patchy landscape: nuclear and mitochondrial markers reveal area effects in the spider Theridion californicum (Araneae: Theridiidae)

Changes in land use have a major effect on patterns of biodiversity. However, few studies have examined the demographic and genetic shifts associated with a return to semi‐natural habitat following extended periods of human disturbance. Here we examine patterns of population structure in a spider restricted to the Pacific coastal strip of North America that exhibits an exuberant colour polymorphism. We use mitochondrial DNA and AFLP markers to examine genetic structure and estimate gene flow. The results show contrasting, gender‐specific patterns between these markers that suggest limited dispersal, combined with area effects most likely caused by expansion from refugial habitat patches following land‐management changes in a region of the San Francisco East Bay. Colour‐morph frequencies are not correlated with this complex genetic structure. Thus, unlike the classical area effects that were based on colour morphs, we demonstrate in T. californicum signals of historical contingency at neutral loci but not at the Colour locus, where traces of past events have been obliterated by balancing selection. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 600–620.     

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.     

Delayed dispersal and the costs and benefits of different routes to independent breeding in a cooperatively breeding bird

Why sexually mature individuals stay in groups as nonreproductive subordinates is central to the evolution of sociality and cooperative breeding. To understand such delayed dispersal, its costs and benefits need to be compared with those of permanently leaving to float through the population. However, comprehensive comparisons, especially regarding differences in future breeding opportunities, are rare. Moreover, extraterritorial prospecting by philopatric individuals has generally been ignored, even though the factors underlying this route to independent breeding may differ from those of strict philopatry or floating. We use a comprehensive predictive framework to explore how various costs, benefits and intrinsic, environmental and social factors explain philopatry, prospecting, and floating in Seychelles warblers (Acrocephalus sechellensis). Not only floaters more likely obtained an independent breeding position before the next season than strictly philopatric individuals, but also suffered higher mortality. Prospecting yielded similar benefits to floating but lower mortality costs, suggesting that it is overall more beneficial than floating and strict philopatry. While prospecting is probably individual‐driven, although limited by resource availability, floating likely results from eviction by unrelated breeders. Such differences in proximate and ultimate factors underlying each route to independent breeding highlight the need for simultaneous consideration when studying the evolution of delayed dispersal.     

Fitness costs and benefits of antipredator behavior mediated by chemotactile cues in the wolf spider Pardosa milvina (Araneae: Lycosidae)

Animals may exhibit a variety of defensive behaviors in thepresence of indirect predator cues. Such behavior offers immediatefitness benefits but may also incur substantial foraging andreproductive costs. We measured shifts in space use (verticalclimbing) by the wolf spider Pardosa milvina induced by chemotactilecues (silk and excreta) from a co-occurring predatory wolfspider Hogna helluo. We then measured foraging and reproductive costs, as well as survival benefits, of this behavior. For 2weeks, we maintained mated adult female Pardosa in plasticcontainers with one of three treated peat moss substrates:a container previously occupied by a conspecific for 3 days,a container previously occupied by an adult Hogna for 3 days,and a container devoid of either cue (control). We measuredprey capture efficiency, body condition, egg sac production,egg sac weight, and egg number for individuals in each treatment.We also counted the number of Pardosa that survived and exhibitedclimbing behavior in the presence of a live Hogna with andwithout silk and excreta cues. Pardosa climbed container wallssignificantly more often in the presence of Hogna silk andexcreta relative to other treatments. Pardosa exposed to Hognacues coupled with live Hogna survived significantly longerthan spiders that had no predator cues available. Pardosa placedin containers with Hogna cues, but no Hogna, lost weight morequickly, ate fewer prey, were in poorer body condition, producedlighter egg sacs, and produced fewer eggs than spiders in controlor conspecific treatments.     

Diapause and seasonal life cycle strategy in the house spider, Achaearanea tepidariorum (Araneae, Theridiidae)

Abstract In order to elucidate the mechanism regulating its seasonal life cycle, the photoperiodic response of Achaearanea tepidariorum has been analysed. Nymphal development was faster in long-day and slower in short-day photoperiods. The combined action of low temperature, poor food supply and short daylength induced diapause at an earlier developmental stage than short days alone. Thus, photoperiod is a primary factor inducing nymphal diapause, but the diapausing instar is influenced by both temperature and food supply. Hibernating nymphs became unresponsive to photoperiod in late December. After hibernation, however, sensitivity was restored and the nymphs remained sensitive to photoperiod throughout their life. This spider could also enter an imaginal or reproductive diapause. Photoperiod was again a primary inducing factor and temperature modified the photoperiodic response to some extent. The induction of the reproductive diapause was almost temperature-compensated whereas development was not. So the involvement of a photoperiodic counter system was suggested. Irrespective of whether the nymph had experienced diapause or not, the imaginal diapause was induced in response to a short-day photoperiod after adult moult. Based on these observations, the seasonal life cycle and the adaptive significance of nymphal and imaginal diapause are discussed.     

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.     

Costs and benefits of facultative aggregating behaviour in the orb-spinning spider Gasteracantha minax Thorell (Araneae: Araneidae)

Abstract The potential costs and benefits of foraging in aggregations are examined for the orb-spinning spider Gasteracantha minax. Web-site tenacity is low in this species; individuals frequently move among sites, thereby joining aggregations of different sizes. Female spiders in aggregations suffered lower predation rates and attracted more males than their solitary counterparts. However, aggregated eggsacs, probably produced by females in aggregations, experienced higher rates of parasitism than solitary eggsacs. We found no evidence of higher prey capture success rates among spiders in aggregations. However, we demonstrate a novel way in which spiders can increase their foraging efficiency by decreasing silk investment. A spider spinning a web within an existing aggregation can attach the support threads of its web to those of other webs, thereby exploiting the silk produced by other spiders.