The significance of relatedness and gene flow on population genetic structure in the subsocial spider Eresus cinnaberinus (Araneae: Eresidae)

Interdemic selection, inbreeding and highly structured populations have been invoked to explain the evolution of cooperative social behaviour in the otherwise solitary and cannibalistic spiders. The family Eresidae consists of species ranging from solitary and intermediate subsocial to species exhibiting fully cooperative social behaviour. In this study we, in a hierarchical analysis, investigated relatedness of putative family clusters, inbreeding and population genetic structure of the subsocial spider Eresus cinnaberinus. Five hierarchical levels of investigation ranging from large scale genetic structure (distances of 250 and 50 km level 1 and 2) over microgeographic structure (20 km² and 4 km², level 3 and 4) to a single hill transect of 200 m (level 5) were performed. The purpose of level 5 was two-fold: (1) to investigate the relatedness of putative family groups, and (2) to evaluate the influence of both family living and sampling design on higher level estimates. Relatedness estimates of putative family groups showed an average relatedness of R=0.26. There was no indication of inbreeding. In contrast to social spiders, genetic variation was abundant, H_e?0.10. The population genetic structure was intermediate between social and asocial spiders. Genetic variance increased continually across hierarchical levels. Family structured neighbourhoods biased differentiation estimates among level 5 samples (F_ST? 0.04) and level 3 and 4 samples (0.07ST<0.18), and apparent inbreeding among level 3 and 4 samples, F_IS>0, was caused by disjunct sampling from separate neighbourhoods. Larger scale samples were highly differentiated 0.12ST<0.26, depending on level and sampling design. Due to a distance effect family living did not influence estimates of the higher level 1. Although the dispersing sex among social spiders and the subsocial E. cinnebarinus differ, females versus males, female behaviour of both sociality classes lead to high genetic variance.     

Constraints affecting partial prey consumption by a crab spider,Diaea sp. indet. (Araneae: Thomisidae)

Summary The influence of feeding constraints on the feeding behaviour of Diaea sp. indet., was investigated. Diaea is a crab spider which ambushes its prey and practises extraintestinal digestion. A laboratory study was carried out using fruit flies, Drosophila immigrans, as prey. Diaea feeds from two sites on the prey — initially the head, followed by the posterior abdomen, with most of the prey's contents being extracted from the head. If additional prey are available, Diaea will, instead of switching to the posterior abdomen, catch a new prey item. The efficiency with which Diaea can extract food is influenced by changes which occur in the prey as a consequence of it being killed and fed on. Evaporative fluid loss from prey is an important constraint on food uptake because in influences the viscosity of the prey's contents. Regardless of whether a new prey item arrives, Diaea discards the prey item on which it is feeding before all of the available food has been extracted from it. The fluid content of the prey is not only part of the food the spider extracts, it is also a resource enabling efficient transfer of food from prey to predator. The value of the prey's fluid content as a resource decreases as a function of feeding time and as a consequence of the spider feeding on the prey.     

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.     

Prey capture by the crab spider Misumena calycina (Araneae: Thomisidae)

Summary Crab spiders Misumena calycina (L.) in pasture rose Rosa carolina flowers regularly attacked bumble bees, smaller bees, and syrphid flies that visited these flowers. Attacks reached a maximum rate of over 20/h during mid morning, but only 1.6% of the most important prey item, bumble bees, were captured. The next most important food source, the most frequently taken item, syrphid flies Toxomerus marginatus (Say), were captured in 39% of the attempts. Since these flies have a biomass only 1/60th that of bumble bees, they comprised a much less important food source than did bumble bees. Spiders would obtain over 7% more food by specializing on bumble bees than by attacking all insect visitors, and as much as 20% more food at certain times of the day. However, they did not show a tendency to specialize at any time.     

Factors influencing the evolution of social behaviour in Australian crab spiders (Araneae: Thomisidae)

The social Diaea are non-territorial, periodically-social spiders that do not weave a snare web, a factor considered to be important in spider sociality. Maternal care and heritable retreats are factors common to most group living animals, including social Diaea; suggesting that they are important factors in the evolution of spider sociality. A 4 year survey, along with field and laboratory experiments revealed that mother spiders provided crucial care in the form of a protective Eucalyptus leaf nest and large prey for their offspring. After the mother's death, the nest was inherited and expanded by the offspring. Larger groups built larger, more protective nests, but expended less individual effort doing so, and so survived better than smaller groups.     

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

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

Female‐limited colour polymorphism in the crab spider Synema globosum (Araneae: Thomisidae)

Conspicuous colour variation, caused by the influence of the environment on phenotype or by genetic differences among individuals, is frequently observed in nature. If genetic in origin, colour variation can facilitate the study of mechanisms that contribute to the maintenance of true polymorphisms. Here we describe, for the first time, the female‐limited colour polymorphism in the crab spider, Synema globosum. We looked for associations between life‐history traits and female colour morph, and identified potential agents of selection that could influence the maintenance of the polymorphism. Our results showed that the polymorphism is discrete and heritable, and that differences in colour among morphs are likely to be detectable by honeybees, birds, and conspecifics. We found limited evidence of differences among morphs in morphology and ecology, and found no differences in components of reproduction. Based on the lines of evidence obtained in this study, we suggest that selection exerted by prey, predators, and/or mates is likely to influence the maintenance of the polymorphism observed in S. globosum. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 368–383.     

Behavioural and molecular evidence for selective immigration and group regulation in the social huntsman spider,Delena cancerides

Movement among social groups interacts with the costs and benefits of group‐living in complex ways. Unlike most other social spiders, the social huntsman spider, Delena cancerides, appears to enter foreign colonies, discriminates kin from non‐kin, and has very limited dispersal options because their bark retreats are rare, making this species an interesting model organism with which to examine the role of inter‐colony movement on group‐living. We examined movement among field colonies of D. cancerides in three ways: (1) by tracking the dispersal and immigration of marked spiders into foreign colonies; (2) by recording resident spiders' behaviour toward introduced immigrants; and (3) by inferring intra‐colony relatedness and immigration patterns through allozyme electrophoresis. Of the marked spiders, only young juveniles moved into neighbouring colonies, whereas subadults and adults did not. Introduced juveniles were tolerated in foreign colonies, whereas introduced adult males and subadults were usually attacked by the resident adult female, unless she had similar sized subadult/adult offspring of her own. Allozyme profiles from unmanipulated field colonies showed that 47% of sampled colonies contained at least one immigrant and that average within colony relatedness was below 0.5. These data align with previous research on the costs and benefits of group‐living for D. cancerides, suggesting that spiders actively seek and regulate group membership based on interests of both the immigrant and the colony. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Microstructure of the silk spigots of the green crab spider Oxytate striatipes (Araneae: Thomisidae)

The genus Oxytate L. Koch, 1878 comprises a homogeneous group of nocturnal crab spiders that have silk apparatuses even though they do not spin webs to trap prey. We examined the microstructure of the silk spinning apparatus of the green crab spider Oxytate striatipes, using field emission scanning electron microscopy. The silk glands of the spider were classified into three types: ampullate, pyriform and aciniform. The spigots of these three types of silk gland occur in both sexes. Two pairs of major ampullate glands send secretory ductules to the anterior spinnerets, and another two pairs of minor ampullate glands supply the median spinnerets. In addition, the pyriform glands send ductules to the anterior spinnerets (45 pairs in females and 40 pairs in males), and the aciniform glands feed silk into the median (9–12 pairs in females and 7–10 pairs in males) and the posterior (30 pairs in both sexes) spinnerets. The spigot system of O. striatipes is simpler and more primitive than other wandering spiders: even the female spiders possess neither tubuliform glands for cocoon production nor triad spigots for web‐building.     

Mating structure and genetic relatedness among gynes in the primitively eusocial wasp Polistes snelleni (Hymenoptera: Vespidae)

Mating structure and genetic relatedness among gynes (potential reproductive females) of the paper wasp Polistes snelleni were investigated using DNA microsatellite markers. All colonies had one foundress inseminated by a single male, and no sign of inbreeding was detected. The mean genetic relatedness among gynes was 0.734 ± 0.028 (SE), which is not significantly different from the expected value of 0.75 for full sisters.     

Complete mitochodrial genome of the crab spider Ebrechtella tricuspidata (Araneae: Thomisidae): A novel tRNA rearrangement and phylogenetic implications for Araneae

Mitochondrial genomes are widely used for phylogenetic and phylogeographic analyses among arthropods, but there is a lack of sufficient mitochondrial genome sequence data for spiders. Herein, we sequenced and characterized the complete mitochondrial genome of a crab spider Ebrechtella tricuspidata (Araneae: Thomisidae). The circular mitochondrial genome is 14,352 bp long, including a standard set of 37 genes and an A + T-rich region. Nucleotide composition is highly biased toward A + T nucleotides (77.3%). A novel gene order rearrangement is detected by a tRNA (trnL1) translocation. Tandem repeats are not identified in the A + T-rich region. Most of the tRNAs are greatly reduced in size and cannot be folded into typical cloverleaf-shaped secondary structures. The phylogenetic analysis confirms that the mitochondrial genome sequences are useful in resolving higher-level relationship of Araneae. Overall, our data present in this study will elevate our knowledge on the architecture and evolution of spider mitochondrial genome.     

Securing paternity: mating plugs in the dwarf spider Oedothorax retusus (Araneae: Erigoninae)

One of the various male strategies to prevent or impede female remating is the production of a mating plug that covers the female genital opening or remains inside of the female genital tract after mating. Such structures have been described for many species in many animal taxa; however, in most cases, we know little or nothing about their specific adaptive value. Our investigations demonstrate that females of the dwarf spider species Oedothorax retusus (Westring, 1851) (Linyphiidae, Erigoninae) exhibit a substance on one or both of her paired genital openings only after copulation. We performed double-mating trials and forced the second male to mate into the previously used or unused spermathecal duct of the female by amputating one of his paired male gonopods (pedipalps). Furthermore, to investigate whether the duration of the first mating has an effect on the size and efficiency of the mating plug, we interrupted first matings after either 1 or 3 min, categorized plug size and recorded mating behaviour of subsequent males. The amount of secretion transferred was larger in long compared to short copulations. A long first copulation successfully prevented subsequent males from mating into the used ducts, whereas mating success after short first matings was similar to matings into unused copulatory ducts of the females. The present study demonstrates that a male O. retusus can prevent a rival from transferring sperm into the same spermatheca by applying a mating plug, but only if he mates for long enough.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 574–583.     

Spatial distribution of genetic relatedness in a moorland population of red grouse (Lagopus lagopus scoticus)

Several aspects of the ecology and biology of red grouse (Lagopus lagopus scoticus) could prevent the complete admixture of genes within and between populations. Male red grouse display a high degree of natal philopatry, are territorial, and show less aggression to kin man to non-kin. Such factors acting in combination predict limited male-mediated gene flow, which will promote social structure within a population by the formation of stable kin clusters, and facilitate a rapid rise in allelic coancestry and/or inbreeding. In this study we utilize hypervariable microsatellite polymorphisms to examine the extent of social affiliation between relatives in a moorland population of grouse from NE Scodand. Levels of genetic relatedness between individual male red grouse occupying territories at Glas Choille in die spring and autumn of 1995 were examined, and kin clusters delimited. Nine kin groups (mean size = 2.4 individuals) were identified prior to breeding in the spring, which increased to 11 kin groups (mean size =4.0 individuals) when territories were reformed in the autumn. The majority of tiiose individuals that were recruited into the adult population during the autumn already had a first-order male relative established, supporting the hypothesis that recruitment is facilitated by behavioural interactions among relatives. The demographic and population genetic consequences of philopatric recruitment and kin clustering are examined and discussed.     

Genital damage in the orb-web spider Argiope bruennichi (Araneae: Araneidae) increases paternity success

The morphology of male genitalia often suggests functions besidessperm transfer that may have evolved under natural or sexualselection. In several species of sexually cannibalistic spiders,males damage their paired genitalia during mating, limitingthem to one copulation per pedipalp. Using a triple-mating experiment,we tested if genital damage in the orb-web spider Argiope bruennichiincreases male fitness either through facilitating his escapefrom an aggressive female or by obstructing the female's inseminationducts against future copulation attempts from other males. Wefound no survival advantage for males damaging their pedipalps;however, copulations into a previously used insemination ductwere significantly shorter when the previous male had left partsof his genitalia inside the insemination duct. Because copulationduration determines paternity in this species, our result suggeststhat male genital damage in A. bruennichi is sexually selected.By breaking off parts of their intromittent organs inside avirgin female, males can reduce sperm competition and therebyincrease their paternity success.     

Characterization of microsatellite loci in the subsocial spider Stegodyphus lineatus (Araneae: Eresidae)

Stegodyphus lineatus spiders live in groups consisting of closely related individuals. There appears to be no discrimination against related individuals as mates but females mate multiply, despite the fact that matings are shown to carry a cost. We have developed eight polymorphic dinucleotide microsatellite markers that allow us to assess levels of heterozygosity and relatedness among individuals of this species. These molecular markers are likely to prove highly effective tools for estimating levels of inbreeding and thus allow us to test hypotheses about the relationships between social structure, mating strategies and inbreeding avoidance.     

The combined effects of temperature and diet on development and survival of a crab spider,Misumenops tricuspidatus (Fabricius) (Araneae: Thomisidae)

In the present study, the combined effects of temperature and diet on development and survival of a crab spider, Misumenops tricuspidatus (Fabricius) (Araneae: Thomisidae) in laboratory conditions were investigated. The experiments were carried out at five constant temperatures ranging from 15°C to 35°C on two kinds of diets, fruit flies (Drosophila melanogaster) and a mixed diet of fruit flies and dung flies. It was found that development rate of eggs increased with successive temperature increments, reaching a maximum at 30°C, then declined at 32°C and that eggs survived well between 20°C and 30°C (>70%), but no eggs survived to hatching at 35°C regardless of whether the spiders were fed on single or mixed diet. Juveniles completed development on both diets at all constant temperatures tested, but survival was low at the extreme temperatures. Juvenile development times decreased over successive temperature increments up to 30°C, then increased at 32°C. Females developed faster than males. Diet also influenced development time, survival and number of moults to reach maturity. Juveniles raised on the mixed diet composed of fruit flies and dung flies developed faster, survived better, and required fewer moults to reach maturity than on a diet composed of only fruit flies. Plots of development rates (reciprocal of mean development times) and survival rates (expressed as percentages) against constant temperatures indicated that M. tricuspidatus is well adapted to low temperatures, but detrimentally affected by high temperatures. Using linear regression, the lower development threshold (LDT) and the sum of effective temperatures (SET) for all life stages of M. tricuspidatus on each diet were estimated. LDT and SET varied among developmental stages and between diets.     

Patterns of diversification and genetic population structure of small mammals in Taiwan

Taiwan is a mountainous island off the coast of the Asian continent. The island is located on the continental shelf and rises to an elevation of nearly 4000 m. It became an island approximately 4 million years ago and has been connected to the continent more than once since its emergence. Therefore, the elevational zonation of two Taiwanese pairs of congeners in the rodent genera Apodemus and Niviventer can be explained by one of two competing hypotheses. One hypothesis assumes that speciation of the two congeners occurred in situ after an ancestral species migrated from the continent. In contrast, the second hypothesis argues the zonation resulted from separate incursion events during the connections. The phylogenetic analyses, which are based on electrophoretic allozyme data, reject the first of these hypotheses. Furthermore, genetic population structure and gene flow of three species of Taiwanese rodents (Apodemus semotus, Niviventer culturatus and Microtus kikuchii) are studied in relation to their patterns of elevational distribution. The genetic structure of A. semotus and N. culturatus, which have extensive elevational distribution, is rather homogeneous and no genie discontinuity or cline in allele frequency was detected. Gene flow among subpopulations for these two species is rather substantial; therefore, the potential isolating effect imposed by deep river valleys is minimal. Conversely, isolation of populations of M. kikuchii on different mountaintops is quite complete as mirrored by the high level of genie differentiation and low gene flow.     

Foraging behavior of the communal spider,Philoponella republicana (Araneae: Uloboridae)

The communal orb-weaving spider, Philoponella republicana,was observed in the subtropical moist forest of Southeast Peru. These spiders live in colonies of conspecifics whose individual orbs are connected by silk. The wrapping of a prey prior to feeding is a large component of the prey capture process because P. republicanahas no venom with which to kill an insect. Wrapping time was the only aspect of prey capture that was strongly correlated with the size of the insect captured. Occasionally we observed several individuals working together to wrap a prey item. These joint efforts were more frequent on prey larger than the capturing spider. Although group captures accounted for only 5.5% of captures, they represented 14.7% of the biomass obtained. A comparison of the relationship between wrapping time and prey size for solitary and group efforts suggested that, by working together, the spiders reduced their total handling time. In most cases only one spider fed on the captured prey.     

Where sociality and relatedness diverge: the genetic basis for hierarchical social organization in African elephants

Hierarchical properties characterize elephant fission–fusion social organization whereby stable groups of individuals coalesce into higher order groups or split in a predictable manner. This hierarchical complexity is rare among animals and, as such, an examination of the factors driving its emergence offers unique insight into the evolution of social behaviour. Investigation of the genetic basis for such social affiliation demonstrates that while the majority of core social groups (second-tier affiliates) are significantly related, this is not exclusively the case. As such, direct benefits received through membership of these groups appear to be salient to their formation and maintenance. Further analysis revealed that the majority of groups in the two higher social echelons (third and fourth tiers) are typically not significantly related. The majority of third-tier members are matrilocal, carrying the same mtDNA control region haplotype, while matrilocality among fourth-tier groups was slightly less than expected at random. Comparison of results to those from a less disturbed population suggests that human depredation, leading to social disruption, altered the genetic underpinning of social relations in the study population. These results suggest that inclusive fitness benefits may crystallize elephant hierarchical social structuring along genetic lines when populations are undisturbed. However, indirect benefits are not critical to the formation and maintenance of second-, third- or fourth-tier level bonds, indicating the importance of direct benefits in the emergence of complex, hierarchical social relations among elephants. Future directions and conservation implications are discussed.     

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.