Males of the orb-web spider Argiope bruennichi sacrifice themselves to unrelated females

Costs of inbreeding can lead to total reproductive failure and inbreeding avoidance is, therefore, common. In classical sex roles with no paternal care, the selective pressure to avoid inbreeding is mostly on the female, which carries the higher costs. In some orb-web spiders, this situation is very different because females are polyandrous and males are monogynous or at most bigynous. Additionally, females of many entelegyne orb weavers are thought to bias paternity post-copulatorily towards a desired mate. This increases the selective pressure on males to adjust their investment in a mating with regard to the compatibility to a female.Here, we examine whether genetic relatedness influences mating behaviour in the orb-web spider Argiope bruennichi. We mated either a sibling or a non-sibling male to a female in single copulation trials and compared copulation duration, cannibalism rate and female fecundity.Our experiment revealed that males prolonged their copulation duration and were cannibalized more frequently when mating with a non-sibling female. Males mating with a sibling female were more likely to escape cannibalism by copulating briefly, thus presumably increasing their chances of re-mating with a more compatible female. This suggests that males can adaptively adjust their investment relating to the compatibility of a female.     

Determinants of natural mating success in the cannibalistic orb-web spider Argiope bruennichi

Monogynous mating systems (low male mating rates) occur in various taxa and have evolved several times independently in spiders. Monogyny is associated with remarkable male mating strategies and predicted to evolve under a male-biased sex ratio. While male reproductive strategies are well documented and male mating rates are easy to quantify, especially in sexually cannibalistic species, female reproductive strategies, the optimal female mating rate, and the factors that affect the evolution of female mating rates are still unclear. In this study, we examined natural female mating rates and tested the assumption of a male-biased sex ratio and female polyandry in a natural population of Argiope bruennichi in which we controlled female mating status prior to observations. We predicted variation in female mating frequencies as a result of spatial and temporal heterogeneity in the distribution of mature females and males. Females had a low average mating rate of 1.3 and the majority copulated only once. Polyandry did not entirely result from a male-biased sex-ratio but closely matched the rate of male bigamy. Male activity and the probability of polyandry correlated with factors affecting pheromone presence such as virgin females' density. We conclude that a strong sex ratio bias and high female mating rates are not necessary components of monogynous mating systems as long as males protect their paternity effectively and certain frequencies of bigyny stabilise the mating system.     

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.     

Function of bright coloration in the wasp spider Argiope bruennichi (Araneae: Araneidae)

There are two major competing explanations for the counter-intuitive presence of bright coloration in certain orb-web spiders. Bright coloration could lure insect prey to the web vicinity, increasing the spider's foraging success. Alternatively, the markings could function as disruptive camouflage, making it difficult for the insect prey to distinguish spiders from background colour variation. We measured the prey capture rates of wasp spiders, Argiope bruennichi, that were blacked out, shielded from view using a leaf fragment, or left naturally coloured. Naturally coloured spiders caught over twice the number of prey as did either blacked-out or leaf-shielded spiders, and almost three times as many orthopteran prey. Spectrophotometer measurements suggest that the bright yellow bands on the spider's abdomen are visible to insect prey, but not the banding on the legs, which could disguise the spider's outline. Thus, our results provide strong support for the hypothesis that bright coloration in the wasp spider acts as a visual lure for insect prey and weak support for the hypothesis that the arrangement of the banding pattern across the spider's body disguises the presence of the spider on the web.     

Novel molecular and mechanical properties of egg case silk from wasp spider, Argiope bruennichi

Araneoid spiders use specialized abdominal glands to produce up to seven different protein-based silks/glues that have various mechanical properties. To date, the fibroin sequences encoding egg case fibers have not been fully determined. To gain further understanding of a recently reported spider silk protein gene family, several novel strategies were utilized in this study to isolate two full-length cDNAs of egg case silk proteins, cylindrical silk protein 1 (CySp1, 9.1 kb) and cylindrical silk protein 2 (CySp2, 9.8 kb), from the wasp spider, Argiope bruennichi. Northern blotting analysis demonstrated that CySp1 and CySp2 are selectively expressed in the cylindrical glands. The amino acid composition of raw egg case silk was closely consistent with the deduced amino acid composition based on the sequences of CySp1 and CySp2, which supports the assertion that CySp1 and CySp2 represent two major components of egg case silk. CySp1 and CySp2 are primarily composed of remarkable homogeneous assemble repeats that are 180 residues in length and consist of several complex subrepeats, and they contain highly homologous C-termini and markedly different N-termini. Our results suggest a possible link between CySp1 and CySp2. In addition, comparisons of stress/strain curves for dragline and egg case silk from Argiope bruennichi showed obvious differences in ultimate strength and extensibility, and similarities in toughness.     

Intramolecular homologous recombination event occurred in the spider egg case silk gene CySp2 of wasp spider Argiope bruennichi

To gain further understanding of egg case silk proteins gene family, Zhao et al. (2006) isolated two full-length cDNAs for egg case silk proteins, cylindrical silk protein 1 (CySp1) and cylindrical silk protein 2 (CySp2), from the wasp spider, Argiope bruennichi. CySp2 was reported to contain no apparent Signal peptide sequences, and the CySp1-CySp2 complex, which would possess a signal peptide, would be transported across the endoplasmic reticulum and secreted to the Golgi. According to a report by Hayashi, genomic DNA sequencing is one approach that can be successfully utilized to retrieve 5′ ends of silk genes; using this method, we retrieved the 5’ end of CySp1. We found that CySp2 contained a typical signal peptide similar to that found in CySp1; thus, due to technical limitations, an artificial error had occurred in the CySp2 sequence reported by Zhao et al.     

Family-specific chemical profiles provide potential kin recognition cues in the sexually cannibalistic spider Argiope bruennichi

Kin recognition, the ability to detect relatives, is important for cooperation, altruism and also inbreeding avoidance. A large body of research on kin recognition mechanisms exists for vertebrates and insects, while little is known for other arthropod taxa. In spiders, nepotism has been reported in social and solitary species. However, there are very few examples of kin discrimination in a mating context, one coming from the orb-weaver Argiope bruennichi. Owing to effective mating plugs and high rates of sexual cannibalism, both sexes of A. bruennichi are limited to a maximum of two copulations. Males surviving their first copulation can either re-mate with the current female (monopolizing paternity) or leave and search for another. Mating experiments have shown that males readily mate with sisters but are more likely to leave after one short copulation as compared with unrelated females, allowing them to search for another mate. Here, we ask whether the observed behaviour is based on chemical cues. We detected family-specific cuticular profiles that qualify as kin recognition cues. Moreover, correlations in the relative amounts of some of the detected substances between sexes within families indicate that kin recognition is likely based on subsets of cuticular substances, rather than entire profiles.     

Web damage and feeding experience influence web site tenacity in the orb-web spider Argiope keyserlingi Karsch

For orb-web spiders, the decision to remain at a particular site or to relocate elsewhere will ultimately depend on the quality of the site. In the past, the quality of the site has been measured in terms of prey availability; spiders experiencing large numbers of prey items are less likely to relocate. However, regular web damage caused by larger nonprey animals may also contribute to the quality of a particular site. Laboratory experiments revealed that the frequency and extent of web relocation by the orb-web spider Argiope keyserlingi was influenced by the feeding regime and the rate of nonprey web damage. Daily movement patterns were influenced by web damage, and these movements were in the direction away from the source of damage. However, the cumulative distance moved during the 7 days of the experiment was influenced by the frequency with which spiders were fed. Spiders that were not given prey moved further than spiders that obtained prey. These data indicate that spiders respond to web damage on a daily basis, but the cumulative movement of spiders over a longer period is influenced mostly by the history of prey ingestion rate. Copyright 2000 The Association for the Study of Animal Behaviour.     

Organization of the spinnerets and spigots in the orb web spider,Argiope bruennichi (Araneae: Araneidae)

Although the basic taxonomic characteristics usually remain unchanged, some spinning apparatuses undergo consistent adaptive variations. As the presence of additional protuberances known as nubbins and tartipores have caused disagreements regarding some Araneidae spiders, more detailed definitions on the cuticular structures have recently been proposed. Reflecting this definition, microstructural organization of silk spinning apparatuses in the orb web spider Argiope bruennichi were reconsidered using field emission scanning electron microscopy. Among the seven kinds of functional spigots in females, it was revealed that two types (major ampullates and pyrifoms) are located on anterior spinnerets and another five types are distributed on median (minor ampullates, tubuliforms and aciniforms) or posterior (tubuliforms, flagelliforms, aggregates and aciniforms) spinnerets, respectively. In addition to functional spigots, cuticular remnants of the nubbins and the tartipores were found on the spinning fields, but the number of tartipores on each spinneret varied among individuals based on maturity. Nevertheless, three kinds of cuticular protuberances of ampullate silk glands were clearly visible at both the anterior and median spinnerets.     

Relative strengths of top-down and bottom-up forces in a tropical forest community

We tested integrative bottom-up and top-down trophic cascade hypotheses with manipulative experiments in a tropical wet forest, using the ant-plant Piper cenocladum and its associated arthropod community. We examined enhanced nutrients and light along with predator and herbivore exclusions as sources of variation in the relative biomass of plants, their herbivores (via rates of herbivory), and resident predaceous ants. The combined manipulations of secondary consumers, primary consumers, and plant resources allowed us to examine some of the direct and indirect effects on each trophic level and to determine the relative contributions of bottom-up and top-down cascades to the structure of the community. We found that enhanced plant resources (nutrients and light) had direct positive effects on plant biomass. However, we found no evidence of indirect (cascading through the herbivores) effects of plant biomass on predators or top predators. In contrast, ants had indirect effects on plant biomass by decreasing herbivory on the plants. This top-down cascade occurred whether or not plant resources were enriched, conditions which are expected to modify top-down forces. Received: 9 August 1998 / Accepted: 1 December 1998     

Effects of social information on life history and mating tactics of males in the orb‐web spider Argiope bruennichi

Informed mating decisions are often based on social cues providing information about prospective mating opportunities. Social information early in life can trigger developmental modifications and influence later mating decisions. A high adaptive value of such adjustments is particularly obvious in systems where potential mating rates are extremely limited and have to be carried out in a short time window. Males of the sexually cannibalistic spider Argiope bruennichi can achieve maximally two copulations which they can use for one (monogyny) or two females (bigyny). The choice between these male mating tactics should rely on female availability that males might assess through volatile sex pheromones emitted by virgin females. We predict that in response to those female cues, males of A. bruennichi should mature earlier and at a smaller body size and favor a bigynous mating tactic in comparison with controls. We sampled spiders from two areas close to the Southern and Northern species range to account for differences in mate quality and seasonality. In a fully factorial design, half of the subadult males from both areas obtained silk cues of females, while the other half remained without female exposure. Adult males were subjected to no‐choice mating tests and could either monopolize the female or leave her (bigyny). We found that Southern males matured later and at a larger size than Northern males. Regardless of their origin, males also shortened the subadult stage in response to female cues, which, however, had no effects on male body mass. Contrary to our prediction, the frequencies of mating tactics were unaffected by the treatment. We conclude that while social cues during late development elicit adaptive life history adjustments, they are less important for the adjustment of mating decisions. We suggest that male tactics mostly rely on local information at the time of mate search.     

Spatial, bottom-up, and top-down effects on the abundance of a leaf miner

Many plant and animal species have higher densities at the centre of their distribution, with a gradual decline in abundance towards the edge of the range, though reasons for this pattern is not well known. We examined the abundance of the leaf miner Cameraria sp. nova over the range of its host plant Quercus myrtifolia in Florida and addressed how bottom-up and top-down factors varied over its whole distribution. Leaf miner densities, plant quality and natural enemy effects on mine survivorship were evaluated in 40 sites and spatially structured models were used to determine the effects of spatial location on the abundance of Cameraria and effects of both bottom-up (tannin concentration, foliar nitrogen, soil nitrogen, and leaf area) and top-down factors (larval parasitism and predation) on abundance and survivorship. Cameraria mines were, on average, three times more abundant on edge/coastal sites compared to centre/inland sites and did not support the hypothesis of higher abundance on the centre of the distribution. Differences in plant quality, larval parasitism and successful emergence of mines on edge versus central sites might be partially responsible for this finding. A trend surface equation with latitude and longitude combined explained almost 52% of the variation in Cameraria density and a trend surface map also revealed peaks of Cameraria abundance on the edges of the plant distribution. Correlograms also indicated a significant spatial structure of Cameraria as mines were positively spatially autocorrelated at small distances (≈122 km). Partial regression analyses indicated that 69% of the variation in Cameraria abundance was explained by the effects of latitude, longitude, elevation and percentage of foliar nitrogen. Our results indicated that variation in Cameraria abundance was mostly explained by spatial position and significant effects of bottom-up and top-down factors were not detected in our large-scale study.     

Eco‐genomic analysis of the poleward range expansion of the wasp spider Argiope bruennichi shows rapid adaptation and genomic admixture

Poleward range expansions are commonly attributed to global change, but could alternatively be driven by rapid evolutionary adaptation. A well‐documented example of a range expansion during the past decades is provided by the European wasp spider Argiope bruennichi. Using ecological niche modeling, thermal tolerance experiments and a genome‐wide analysis of gene expression divergence, we show that invasive populations have adapted to novel climatic conditions in the course of their expansion. Their climatic niche shift is mirrored in an increased cold tolerance and a population‐specific and functionally differentiated gene expression response. We generated an Argiope reference genome sequence and used population genome resequencing to assess genomic changes associated with the new climatic adaptations. We find clear genetic differentiation and a significant admixture with alleles from East Asian populations in the invasive Northern European populations. Population genetic modeling suggests that at least some of these introgressing alleles have contributed to the new adaptations during the expansion. Our results thus confirm the notion that range expansions are not a simple consequence of climate change, but are accompanied by fast genetic changes and adaptations that may be fuelled through admixture between long separated lineages.     

Ecosystem alteration modifies the relative strengths of bottom-up and top-down forces in a herbivore population

1. Ecosystem alterations can affect the abundance, distribution and diversity of plants and animals, and thus potentially change the relative strength of bottom-up (the plant resource) and top-down (natural enemies) trophic forces acting on herbivore populations. 2. The hypothesis that alterations of the forest ecosystem associated with precommercial thinning have contributed to the increased severity of outbreaks of Neodiprion abietis (Harris), a sawfly defoliator, through the reduction of trophic forces acting on N. abietis larvae, was tested using exclusion techniques. 3. The relative contributions to N. abietis larval mortality of bottom-up and top-down forces both increased with increasing levels of defoliation and were both reduced by thinning. The reduction of bottom-up and top-down forces caused a 58% mean increase in N. abietis larval survival in thinned compared with untreated stands, which is less than would be expected by the sum of the effects of thinning on each source of mortality. Evidence indicates that the partly compensatory, partly additive nature of the mortality associated with trophic forces in the system under study is responsible for this discrepancy. 4. To our knowledge, this is the first study to show the impact of ecosystem alterations on the balance between bottom-up and top-down forces acting on an eruptive herbivore population along a gradient of host-plant defoliation, and how this can lead to increased outbreak severity. It is stressed that accurate estimates of the relative contributions of bottom-up and top-down forces to mortality cannot be obtained if the additive or compensatory nature of the mortality associated with these trophic forces is overlooked.     

横纹金蛛捕食经验对其丝纤维力学行为与性能的影响

蛛网是蜘蛛的捕食工具,蛛网网丝的结构与性能不仅影响蜘蛛的捕食效率,也关系着蜘蛛的捕食投入。本文利用单纤维电子强力仪研究横纹金蛛(Argiope bruennichi)室内捕食面包虫(Tenebrio molitor)时上前与返回捕食拖丝的力学性能以及捕食经验对圆网半径丝修补前后的力学性能的影响。结果表明,与上前捕食拖丝相比,返回捕食拖丝减小了弹性区的投入,增加了屈服区和加强区的投入,且返回时捕食拖丝更具柔韧性。整体而言,与初始半径丝相比,在未喂食面包虫的条件下,网的半径修补丝增加了力学性能的投入;而在喂食面包虫的条件下,网的半径修补丝减少了力学性能的投入。所测试丝样中出现了两种类型的蛛丝力学行为:一种为典型的蛛丝力学行为;另外一种为似黏流性材料的力学行为,其反映的是满足蛛丝耗散猎物或自身下降时动能的另外一种力学性能的策略。本研究表明蜘蛛能根据其捕食经验遵循Cost-Benefit原则对蜘蛛丝的力学性能进行调节,从而调整捕食投入。     

The molecular structures of major ampullate silk proteins of the wasp spider,Argiope bruennichi: A second blueprint for synthesizing de novo silk

The dragline silk of orb-weaving spiders possesses extremely high tensile strength and elasticity. To date, full-length sequences of only two genes encoding major ampullate silk protein (MaSp) in Latrodectus hesperus have been determined. In order to further understand this gene family, we utilized in this study a variety of strategies to isolate full-length MaSp1 and MaSp2 cDNAs in the wasp spider Argiope bruennichi. A. bruennichi MaSp1 and MaSp2 are primarily composed of remarkably homogeneous ensemble repeats containing several complex motifs, and both have highly conserved C-termini and N-termini. Two novel amino acid motifs, GGF and SGR, were found in MaSp1 and MaSp2, respectively. Amino acid composition analysis of silk, luminal contents and predicted sequences indicates that MaSp1 and MaSp2 are two major components of major ampullate glands and that the ratio of MaSp1 to MaSp2 is approximately 3:2 in dragline silk. Furthermore, both the MaSp1:MaSp2 ratio and the conserved termini are closely linked with the production of high quality synthetic fibers. Our results make an important contribution to our understanding of major ampullate silk protein structure and provide a second blueprint for creating new composite silk which mimics natural spider dragline silk.     

Relocation, reproduction and remaining alive in the orb-web spider

When mortality is high, animals run a risk if they wait to accumulate resources for improved reproduction so they may trade-off the time of reproduction with number and size of offspring. Animals may attempt to improve food acquisition by relocation, even in 'sit and wait' predators. We examine these factors in an isolated population of an orb-web spider Zygiella x-notata . The population was monitored for 200 days from first egg laying until all adults had died. Large females produced their first clutch earlier than did small females and there was a positive correlation between female size and the number and size of eggs produced. Many females, presumably without eggs, abandoned their web site and relocated their web position. This is presumed because female Zygiella typically guard their eggs. In total, c . 25% of females reproduced but those that relocated were less likely to do so, and if they did, they produced the clutch at a later date than those that remained. When the date of lay was controlled there was no effect of relocation on egg number but relocated females produced smaller eggs. The data are consistent with the idea that females in resource-poor sites are more likely to relocate. Relocation seems to be a gamble to find a more productive site but one that achieves only a late clutch of small eggs and few achieve that.     

悦目金蛛卵袋的结构与组成

采用SEM和氨基酸自动分析仪对悦目金蛛(Argiope amoena)卵袋的结构、纤维组成和覆盖层的氨基酸组成进行了观察研究.结果表明悦目金蛛卵袋呈封闭的倒三角形,是由多种丝腺纺出的丝纤维形成的多个覆盖层构成的,从外向内分别为:金黄色大壶状腺框架丝,灰绿色、浅棕色、白色外覆盖层和棕色内覆盖层,其中外覆盖层是由6根大直径(约4.8-6.8 μm) 柱状腺丝纤维和一些小直径(约0.26-0.76 μm)纤维混合成的基本纤维束单元所折叠成的片层丝被构成的,小直径丝纤维的数量从灰绿色丝被(外)到白色丝被(内)逐渐增加,而内覆盖层仅仅是由6根大直径(约5.20 μm)柱状腺丝纤维构成的基本纤维束单元折叠而成.卵袋外覆盖层丝(白色绒状纤维除外)的甘氨酸、丙氨酸和丝氨酸等氨基酸的百分含量分别介于卵袋内覆盖层丝和葡萄状腺丝相对应氨基酸的百分含量之间,这一数据支持Foradori等关于卵袋小直径丝纤维是由葡萄状腺纺出的推论.悦目金蛛能调节卵袋层丝的颜色、直径、覆盖层丝纤维和氨基酸的组成,使卵袋获得各方面最优化的功能,为卵或若蛛提供支持和保护.