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Spontaneous male death during copulation in an orb-weaving spider.

Males of some cannibalistic species of spiders and insects appear to sacrifice themselves by allowing the female to eat them, and the adaptive significance of such drastic terminal reproductive investment has recently been demonstrated for a spider. Typically, the female has to kill the male, but it has been suggested that males of some species in the cannibalistic orb-weaving spider genus Argiope may die in copula without female 'collaboration'. Here, we provide the first experimental evidence to our knowledge of programmed sudden death after onset of copulation in males of the spider Argiope aurantia. Our observations reveal that males invariably die during the insertion of their second pedipalp, regardless of whether they mate with newly moulted, defenceless females or with older mature females that often attack them. We determined experimentally that the death of males is triggered immediately upon insertion of the second palp, when males become unresponsive, and heartbeat ceases within minutes of insertion. We discuss the possible adaptive significance of programmed death during copulation, and argue that male death has evolved in a context other than sexual cannibalism.

Animals↗

Spider signals: are web decorations visible to birds and bees?

We are becoming increasingly aware of animal communication outside the range of human sensitivity. Web decorations are silk structures used by orb-web spiders to deceive prey and predators. However, despite the level of interest in these structures, their visibility to prey and predators has never, to our knowledge, been objectively assessed. Here, we use spectrophotometric analyses to show that the decorations of all five tested spider species are visible to honey bees and birds over short and long distances. Furthermore, the discoid decorations of one species may provide some protection against arthropod predators. However, these decorations are inefficient at camouflaging the spider against birds, despite the overlap between the spider's body and web decoration.

Animal Communication↗

Oldest true orb-weaving spider (Araneae: Araneidae).

The aerial orb web woven by spiders of the family Araneidae typifies these organisms to laypersons and scientists alike. Here we describe the oldest fossil species of this family, which is preserved in amber from Alava, Spain and represents the first record of Araneidae from the Lower Cretaceous. The fossils provide direct evidence that all three major orb web weaving families: Araneidae, Tetragnathidae and Uloboridae had evolved by this time, confirming the antiquity of the use of this remarkable structure as a prey capture strategy by spiders. Given the complex and stereotyped movements that all orb weavers use to construct their webs, there is little question regarding their common origin, which must have occurred in the Jurassic or earlier. Thus, various forms of this formidable prey capture mechanism were already in place by the time of the explosive Cretaceous co-radiation of angiosperms and their flying insect pollinators. This permitted a similar co-radiation of spider predators with their flying insect prey, presumably without the need for a 'catch-up lag phase' for the spiders.

Animals↗

Hatching responses of subsocial spitting spiders to predation risk.

The carrying of eggs often renders parents vulnerable to predators due to increased conspicuousness or decreased mobility. Nonetheless, egg-carrying parents can escape from the predators to which they are vulnerable. Previous studies have demonstrated heavy predation by spider-eating jumping spiders (Portia labiata) on egg-carrying spitting spider (Scytodes pallida) females, but little predation on eggless females. If the timing of hatching is phenotypically plastic, then both S. pallida females and their eggs could reduce the risk of predation by hatching early. Hence, this study examines the hatching responses of S. pallida to chemical cues from P. labiata, both in the laboratory and in the field, and addresses the following questions. (i) Do cues from predatory P. labiata influence the hatching traits of S. pallida? (ii) Are the olfactory cues from predators sufficient for predator detection by S. pallida ? (iii) Are hatching responses to predatory P. labiata controlled by egg-carrying S. pallida females, or directly by their embryos? The study provides evidence of hatching as a life-history switch point, which shows an adaptive plasticity in response to predation risk in egg-carrying S. pallida. Egg-carrying S. pallida females, but not unattended eggs, adjust egg-hatching time (the interval between oviposition and hatching) in response to the threat of predation on both the female and her eggs by P. labiata. In the presence of P. labiata, eggs that are carried by females hatch sooner; the hatchlings of these eggs are therefore smaller than hatchlings born in the absence of P. labiata. Chemical cues that are released from the draglines of P. labiata are sufficient to elicit changes in the egg-hatching traits of S. pallida. Hatching early in response to this predator may benefit both females and their offspring. To my knowledge, this is the first direct experimental study to demonstrate predator-induced hatching plasticity in spiders and, in particular, in animals with parental care.

Adaptation, Physiological↗

Spiders that decorate their webs at higher frequency intercept more prey and grow faster.

Many orb-weaving spiders decorate their webs with extra, bright white, ultraviolet light reflecting silk. Previous studies suggest that these decorations increase a spider's foraging efficiency by improving web attractiveness, which is known as the prey-attraction hypothesis. One assumption of this hypothesis is that individuals which decorate their webs at a higher frequency are expected to have a higher growth rate. Using a decoration-building orb-weaving spider, Argiope versicolor, I show a strong positive relationship between the growth rate in terms of weight gain and the frequency of decoration-building, as well as the rate of insect interception. This is the first study to reveal a fitness consequence of decorating behaviour in spiders.

Analysis of Variance↗

The age and evolution of sociality in Stegodyphus spiders: a molecular phylogenetic perspective.

Social, cooperative breeding behaviour is rare in spiders and generally characterized by inbreeding, skewed sex ratios and high rates of colony turnover, processes that when combined may reduce genetic variation and lower individual fitness quickly. On these grounds, social spider species have been suggested to be unstable in evolutionary time, and hence sociality a rare phenomenon in spiders. Based on a partial molecular phylogeny of the genus Stegodyphus, we address the hypothesis that social spiders in this genus are evolutionary transient. We estimate the age of the three social species, test whether they represent an ancestral or derived state and assess diversification relative to subsocial congeners. Intraspecific sequence divergence was high in all of the social species, lending no support for the idea that they are young, transient species. The age of the social lineages, constant lineage branching and the likelihood that social species are independently derived suggest that either the social species are 'caught in sociality' or they have evolved into cryptic species.

Animals↗

Wolbachia and other endosymbiont infections in spiders.

Maternally inherited endosymbiotic bacteria, such as Wolbachia, Rickettsia and Spiroplasma, have been shown to have wide-ranging effects on the reproduction of their hosts. We present data on the presence of each of these sorts of bacteria in spiders, a group for which there are currently few data, but where such infections could explain many observed reproductive characteristics, such as sex ratio skew. The Wolbachia and Spiroplasma variants that we find in spiders belong to the same clades previously found to infect other arthropods, but many of the rickettsias belong to two, novel, hitherto spider-specific bacterial lineages. We find evidence for coexistence of different bacterial types within species, and in some cases, within individuals. We suggest that spiders present a useful opportunity for studying the effect of these sorts of bacteria on the evolution of host traits, such as those that are under sexual selection.

Animal Diseases↗

Expression of Pax group III genes suggests a single-segmental periodicity for opisthosomal segment patterning in the spider Cupiennius salei.

Pair-rule patterning forms a key step for segmentation in insects. The expression patterns of pair-rule gene orthologs in representatives of other arthropod groups imply that these genes were segmentation genes in the last common ancestor of the various arthropod groups, but almost nothing is known about the underlying mechanism in noninsect arthropods. Here, we cloned and analyzed members of the Pax group III genes from the spider Cupiennius salei. Pax group III genes comprise genes like the Drosophila genes paired, gooseberry, and gooseberry-neuro, as well as the vertebrate Pax 3 and Pax 7 genes. We recovered three Pax group III genes from the spider C. salei, Cs-pairberry-1, Cs-pairberry-2, and Cs-pairberry-3, and show that the combined expression of the three spider genes mimics the patterns in insects, suggesting an ancestral role for Pax group III genes in segmentation, neurogenesis, and appendage formation in arthropods. One of the genes, pairberry-3, is expressed in a segmental periodicity before overt morphological segmentation is visible, suggesting a single segmental periodicity for opisthosomal segment pattering in the spider. Comparisons among arthropods suggest that the underlying mechanisms for pair-rule gene orthologs are more diverged than the ones for the segment-polarity genes. We argue that there may be a correlation between the lower variation in patterns of segment-polarity genes and the phylotypic stage. The segment-polarity genes are required to define the segment borders of the embryo at the germ-band stage, the arthropod phylotypic stage. Pair-rule gene orthologs act more upstream and may display more variation in their action.

Animals↗

Use of spider silk fibres as an innovative material in a biocompatible artificial nerve conduit.

Defects of peripheral nerves still represent a challenge for surgical nerve reconstruction. Recent studies concentrated on replacement by artificial nerve conduits from different synthetic or biological materials. In our study, we describe for the first time the use of spider silk fibres as a new material in nerve tissue engineering. Schwann cells (SC) were cultivated on spider silk fibres. Cells adhered quickly on the fibres compared to polydioxanone monofilaments (PDS). SC survival and proliferation was normal in Live/Dead assays. The silk fibres were ensheathed completely with cells. We developed composite nerve grafts of acellularized veins, spider silk fibres and SC diluted in matrigel. These artificial nerve grafts could be cultivated in vitro for one week. Histological analysis showed that the cells were vital and formed distinct columns along the silk fibres. In conclusion, our results show that artificial nerve grafts can be constructed successfully from spider silk, acellularized veins and SC mixed with matrigel.

Animals↗

Extreme diversity, conservation, and convergence of spider silk fibroin sequences.

Spiders (Araneae) spin high-performance silks from liquid fibroin proteins. Fibroin sequences from basal spider lineages reveal mosaics of amino acid motifs that differ radically from previously described spider silk sequences. The silk fibers of Araneae are constructed from many protein designs. Yet, the repetitive sequences of fibroins from orb-weaving spiders have been maintained, presumably by stabilizing selection, over 125 million years of evolutionary history. The retention of these conserved motifs since the Mesozoic and their convergent evolution in other structural superproteins imply that these sequences are central to understanding the exceptional mechanical properties of orb weaver silks.

Amino Acid Motifs↗

Distribution and function of GABAB receptors in spider peripheral mechanosensilla.

The mechanosensilla in spider exoskeleton are innervated by bipolar neurons with their cell bodies close to the cuticle and dendrites attached to it. Numerous efferent fibers synapse with peripheral parts of the mechanosensory neurons, with glial cells surrounding the neurons, and with each other. Most of these efferent fibers are immunoreactive to gamma-aminobutyric acid (GABA), and the sensory neurons respond to agonists of ionotropic GABA receptors with a rapid and complete inhibition. In contrast, little is known about metabotropic GABAB receptors that may mediate long-term effects. We investigated the distribution of GABAB receptors on spider leg mechanosensilla using specific antibodies against 2 proteins needed to form functional receptors and an antibody that labels the synaptic vesicles on presynaptic sites. Both anti-GABAB receptor antibodies labeled the distal parts of the sensory cell bodies and dendrites but anti-GABABR1 immunoreactivity was also found in the axons and proximal parts of the cell bodies and some glial cells. The fine efferent fibers that branch on top of the sensory neurons did not show GABAB receptor immunoreactivity but were densely labeled with anti-synapsin and indicated synaptic vesicles on presynaptic locations to the GABAB receptors. Intracellular recordings from sensory neurons innervating the slit sensilla of the spider legs revealed that application of GABAB receptor agonists attenuated voltage-activated Ca2+ current and enhanced voltage-activated outward K+ current, providing 2 possible mechanisms for controlling the neurons' excitability. These findings support the hypothesis that GABAB receptors are present in the spider mechanosensilla where their activation may modulate information transmission.

Animals↗

Multisensory cues and multimodal communication in spiders: insights from video/audio playback studies.

Spiders perceive the world using multiple sensory modes, including vibration, vision, and chemical senses, for prey detection and communication. These sensory modes are used in many communication contexts, either individually or in multimodal signaling. Selection for effective signaler-receiver communication and species discrimination is especially strong for these predatory and potentially cannibalistic arthropods, resulting in the evolution of considerable diversity in signaling behaviors. In this paper, we review sensory mechanisms involved in spider signaling and present an overview of recent work done on wolf spiders (Lycosidae) that use multimodal communication (simultaneous visual and vibratory signals) in sexual signals during courtship. The relative importance of visual and vibratory signaling modes, and the use of multiple modes varies among closely related species in the genus Schizocosa, providing a model system for investigating multisensory guidance of complex behavior. Here we examine previous and current research on responses of female spiders to components of male courtship behavior, using several experimental techniques including cue isolation (single sensory modes), video/audio digitization and playback, and cue-conflict (mixed conspecific/heterospecific components) to tease apart elements of multimodal signaling.

Animal Communication↗

Recruitment of cell groups through Delta/Notch signalling during spider neurogenesis.

Early neurogenesis in the spider is characterised by a stereotyped pattern of sequential recruitment of neural cells from the neuroectoderm, comparable with neuroblast formation in Drosophila: However, in contrast to Drosophila, where single cells delaminate from the neuroectoderm, groups of cells adopt the neural fate and invaginate into the spider embryo. This raises the question of whether Delta/Notch signalling is involved in this process, as this system normally leads to a singling out of individual cells through lateral inhibition. I have therefore cloned homologues of Delta and Notch from the spider Cupiennius salei and studied their expression and function. The genes are indeed expressed during the formation of neural cells in the ventral neuroectoderm. Loss of function of either gene leads to an upregulation of the proneural genes and an altered morphology of the neuroectoderm that is comparable with Delta and Notch mutant phenotypes in Drosophila: Thus, although Delta/Notch signalling appears to be used in the same way as in Drosophila, the lateral inhibition process produces clusters of invaginating cells, rather than single cells. Intriguingly, neuroectodermal cells that are not invaginating seem to become neural cells at a later stage, while the epidermal cells are derived from lateral regions that overgrow the neuroectoderm. In this respect, the neuroectodermal region of the spider is more similar to the neural plate of vertebrates, than to the neuroectoderm of Drosophila:

Amino Acid Sequence↗

Spider peripheral mechanosensory neurons are directly innervated and modulated by octopaminergic efferents.

Octopamine is a chemical relative of noradrenaline providing analogous neurohumoral control of diverse invertebrate physiological processes. There is also evidence for direct octopaminergic innervation of some insect peripheral tissues. Here, we show that spider peripheral mechanoreceptors are innervated by octopamine-containing efferents. The mechanosensory neurons have octopamine receptors colocalized with synapsin labeling in the efferent fibers. In addition, octopamine enhances the electrical response of the sensory neurons to mechanical stimulation. Spider peripheral mechanosensilla receive extensive efferent innervation. Many efferent fibers in the legs of Cupiennius salei are GABAergic, providing inhibitory control of sensory neurons, but there is also evidence for other neurotransmitters. We used antibody labeling to show that some efferents contain octopamine and that octopamine receptors are concentrated on the axon hillocks and proximal soma regions of all mechanosensory neurons in the spider leg. Synaptic vesicles in efferent neurons were concentrated in similar areas. Octopamine, or its precursor tyramine, increased responses of mechanically stimulated filiform (trichobothria) leg hairs. This effect was blocked by the octopamine antagonist phentolamine. The octopamine-induced modulation was mimicked by 8-Br-cAMP, a cAMP analog, and blocked by Rp-cAMPS, a protein kinase A inhibitor, indicating that spider octopamine receptors activate adenylate cyclase and increase cAMP concentration. Frequency response analysis showed that octopamine increased the sensitivity of the trichobothria neurons over a broad frequency range. Thus, the major effect of octopamine is to increase its overall sensitivity to wind-borne signals from sources such as flying insect prey or predators.

8-Bromo Cyclic Adenosine Monophosphate↗

Design of superior spider silk: from nanostructure to mechanical properties.

Spider dragline silk is of practical interest because of its excellent mechanical properties. However, the structure of this material is still largely unknown. In this article, we report what we believe is a new model of the hierarchical structure of silk based on scanning electron microscope and atomic force microscope images. This hierarchical structure includes beta-sheet, polypeptide chain network, and silk fibril. It turns out that an exceptionally high strength of the spider dragline silk can be obtained by decreasing the size of the crystalline nodes in the polypeptide chain network while increasing the degree of orientation of the crystalline nodes. Based on this understanding, how the reeling speed affects mechanical properties of spider dragline silk can be understood properly. Hopefully, the understanding obtained in this study will shed light on the formation of spider silk, and consequently, on the principles for the design of ultrastrong silk.

Animals↗

Spiders of soybean crops in Santa Fe province, Argentina: influence of surrounding spontaneous vegetation on lot colonization.

Trials during two consecutive soybean cycles were performed in central Santa Fe in order to determine the main spider families present in the crop and to determine the influence of spontaneous margin flora on colonization towards the lot. Samplings were done by sweeping net and pitfall traps. It was concluded that:1. Oxyopidae was the most frequent family in the herbaceous layer of both the margins and the soybean crop, and Lycosidae in the lower layer; 2. Margin strips in a soybean lot contribute to the colonization of the crop by spiders of aerial habits and also promote re-colonization following pesticide applications, since they act as shelters. The influence on spiders of terrestrial habits was somewhat lower; 3. The distribution of the populations of spiders of terrestrial habits was homogeneous in a soybean crop seeded directly and these predators had a greater capacity to control pests at all points of the lot.

Animals↗

Spider fauna in apple ecosystem of western Oregon and its field susceptibility to chemical and microbial insecticides.

Twelve families, 26 genera, and 30 identifiable spider species were found in surveys conducted in apple orchards of western Oregon. The Salticidae, Linyphiidae, Clubionidae, Philodromidae, and Theridiidae comprised 85.56% ofthe total spiders collected. The most common species in order of abundance were Metaphidippus aeneolus Curtis, Spirembolus mundus Chamberlin & Ivie, Cheiracanthium inclusum (Hentz), Philodromus spectabilis Keyserling, Eris marginata (Walckenaer), and Theridion lawrencei Gertsch & Archer. Individuals of these species were collected in 50-60% of the samples and were most abundant in the month of August. The Bacillus thuringiensis-based insecticides, DiPel (100 Million International Units/100 liters) and MVP (250 ml/100 liters), summer oil (0.5-1.0 liter/100 liters), the insect growth regulator (IGR) diflubenzuron (3-12 g/100 liters), and organophosphate Phosmet (6-60 g/100 liters) were generally harmless (P > 0.05) to these spider species. Full field rates of organophosphate azinphosmethyl (25 g/100 liters) and carbamate carbaryl (60 g/100 liters) were slightly to moderately harmful (25-75% mortality). These insecticides at reduced rates (azinphosmethyl 2.5-5.0 g and carbaryl 12 g/100 liters) applied alone or in combination with DiPel and MVP, had a negligible effect. Full rates of pyrethroids esfenvalerate (2.5 g/100 liters) and permethrin (4.0 g/100 liters) were moderately to highly harmful (50-75% mortality) and their reduced rates (esfenvalerate 0.25-0.50 g and permethrin 0.4-0.8 g/100 liters) were selective to the spiders.

Agriculture↗

Bromeliad-living spiders improve host plant nutrition and growth.

Although bromeliads are believed to obtain nutrients from debris deposited by animals in their rosettes, there is little evidence to support this assumption. Using stable isotope methods, we found that the Neotropical jumping spider Psecas chapoda (Salticidae), which lives strictly associated with the terrestrial bromeliad Bromelia balansae, contributed 18% of the total nitrogen of its host plant in a greenhouse experiment. In a one-year field experiment, plants with spiders produced leaves 15% longer than plants from which the spiders were excluded. This is the first study to show nutrient provisioning in a spider-plant system. Because several animal species live strictly associated with bromeliad rosettes, this type of facultative mutualism involving the Bromeliaceae may be more common than previously thought.

Analysis of Variance↗